The harpy eagle (Harpia harpyja) is a large neotropical species of eagle. It is also called the American harpy eagle to distinguish it from the Papuan eagle, which is sometimes known as the New Guinea harpy eagle or Papuan harpy eagle.[5] It is the largest bird of prey throughout its range,[6] and among the largest extant species of eagles in the world. It usually inhabits tropical lowland rainforests in the upper (emergent) canopy layer. Destruction of its natural habitat has caused it to vanish from many parts of its former range, and it is nearly extirpated from much of Central America. The genusHarpia, together with Harpyopsis, Macheiramphus and Morphnus, form the subfamily Harpiinae.
The harpy eagle was first described by Carl Linnaeus in his landmark 1758 10th edition of Systema Naturae as Vultur harpyja,[7] after the mythological beast harpy. It is now the only species placed in the genus Harpia that was introduced in 1816 by the French ornithologist Louis Pierre Vieillot.[8][9] The harpy eagle is most closely related to the crested eagle (Morphnus guianensis), the Papuan eagle (Harpyopsis novaeguineae) and the bat hawk (Macheiramphus alcinus), the four composing the subfamily Harpiinae within the large family Accipitridae. Previously thought to be closely related, the Philippine eagle has been shown by DNA analysis to belong elsewhere in the raptor family, as it is related to the Circaetinae.[10]
The specific name harpyja and the word "harpy" in the common name both come from Ancient Greekharpyia (ἅρπυια). They refer to the harpies of Ancient Greek mythology. These were wind spirits who flew the dead to Hades or Tartarus,
purported to have the lower body and talons of a raptor and the head of
a woman, standing anywhere from the height of a tall child to as high
as a grown man; some depictions have the creatures possessing an
eagle-like body with the exposed breasts of an elderly female human, a
giant wingspan and the head of a grotesque, sharp-toothed, mutant
eagle—something more akin to a goblin with wings.[11]
The upperside of the harpy eagle is covered with slate-black feathers, and the underside is mostly white, except for the feathered tarsi,
which are striped black. A broad black band across the upper breast
separates the gray head from the white belly. The head is pale grey, and
is crowned with a double crest. The upperside of the tail is black with
three gray bands, while the underside of it is black with three white
bands. The irises are gray or brown or red, the cere and bill are black
or blackish and the tarsi and toes are yellow. The plumage of males and females is identical. The tarsus is up to 13 cm (5.1 in) long.[12][13]
Female harpy eagles typically weigh 6 to 9 kg (13 to 20 lb).[14][15][12][16] One source states that adult females can weigh up to 10 kg (22 lb).[17] An exceptionally large captive female, "Jezebel", weighed 12.3 kg (27 lb).[18]
Being captive, however, this large female may not be representative of
the weight possible in wild harpy eagles due to differences in the food
availability.[19][20] The male, in comparison, is much smaller and may range in weight from 4 to 6 kg (8.8 to 13.2 lb).[14][12][16][15]
The average weight of adult males has been reported as 4.4 to 4.8 kg
(9.7 to 10.6 lb) against an average of 7.3 to 8.3 kg (16 to 18 lb) for
adult females, a 35% or higher difference in mean body mass.[15][21][22] Harpy eagles may measure from 86.5 to 107 cm (34.1 to 42.1 in) in total length[13][16] and have a wingspan of 176 to 224 cm (69 to 88 in).[12][13] Among the standard measurements, the wing chord measures 54–63 cm (21–25 in), the tail measures 37–42 cm (15–17 in), the tarsus is 11.4–13 cm (4.5–5.1 in) long, and the exposed culmen from the cere (the beak) is 4.2 to 6.5 cm (1.7 to 2.6 in).[12][23][24] Mean talon size is 8.6 cm (3.4 in) in males, and 12.3 cm (4.8 in) in females.[25]
It is sometimes cited as the largest eagle alongside the
Philippine eagle, which is somewhat longer on average (between sexes
averaging 100 cm (39 in)) but weighs slightly less, and the Steller's sea eagle, which is perhaps slightly heavier on average (mean of three unsexed birds was 7.75 kg (17.1 lb)).[11][22][26]
The harpy eagle may be the largest bird species to reside in Central America, though large water birds such as American white pelicans (Pelecanus erythrorhynchos) and jabirus (Jabiru mycteria) have scarcely lower mean body masses.[22]
The wingspan of the harpy eagle is relatively small, though the wings
are quite broad, an adaptation that increases maneuverability in
forested habitats and is shared by other raptors in similar habitats.
The wingspan of the harpy eagle is surpassed by several large eagles
that live in more open habitats, such as those in the Haliaeetus and Aquila genera.[12] The extinct Haast's eagle was significantly larger than all extant eagles, including the harpy.[27]
This species is largely silent away from the nest. There, the
adults give a penetrating, weak, melancholy scream, with the incubating
males' call described as "whispy screaming or wailing".[28]
The females' calls while incubating are similar, but are lower-pitched.
While approaching the nest with food, the male calls out "rapid chirps,
goose-like calls, and occasional sharp screams". Vocalization in both
parents decreases as the nestlings age, while the nestlings become more
vocal. The nestlings call chi-chi-chi...chi-chi-chi-chi,
seemingly in alarm in response to rain or direct sunlight. When humans
approach the nest, the nestlings have been described as uttering croaks,
quacks, and whistles.[29]
Relatively rare and elusive throughout its range, the harpy eagle is found from southern México (incl. Chiapas, Oaxaca and the Yucatán states) and south through Central America, into South America to as far south as Argentina. They can still be seen by tourists and locals in Costa Rica and Panamá. As their preferred habitat is rainforest, they nest and hunt predominantly in the emergent layer. The eagle is most common in Brazil, where it is found across the entire country.[30] With the exception of some areas of the aforementioned Panamá and Costa Rica, the species is nearly extinct in Central America,
likely due to the logging industry’s decimation of much of the
Meso-American rainforests. Their habitat is expected to decline further
due to climate change.[31]
The harpy eagle prefers tropical, lowland rainforests and may also
choose to nest within such areas from the canopy to the emergent
vegetation. They typically occur below an elevation of 900 m (3,000 ft),
but have been recorded at elevations up to 2,000 m (6,600 ft).[2]
Within the forests, they hunt in the canopy or, rarely, on the ground,
and perch on emergent trees to scout for prey. They do not generally
occur in disturbed areas, avoiding humans whenever possible, but
regularly visit semi-open forest and pasture mosaic, in hunting forays.[32] Harpies, however, can be found flying over forest borders in a variety of habitats, such as cerrados, caatingas, buriti palm stands, cultivated fields, and cities.[33] They have recently been found in areas where high-grade forestry is practiced.
Feeding on small prey
A stuffed specimen of a harpy eagle preying on a macaw at the Museum für Naturkunde, Berlin
Full grown harpy eagles are at the top of a food chain.[34] They possess the largest talons of any living eagle and have been recorded as carrying prey weighing up to roughly half of their own body weight.[12]
This allows them to snatch from tree branches a live sloth and other
large prey items. Most commonly, harpy eagles use perch hunting, in
which they scan for prey activity while briefly perched between short
flights from tree to tree.[12]
Upon spotting prey, the eagle quickly dives and grabs it. Sometimes,
harpy eagles are "sit-and-wait" predators (common in forest-dwelling
raptors), perching for long periods on a high point near an opening, a
river, or a salt lick, where many mammals go to attain nutrients.[12]
On occasion, they may also hunt by flying within or above the canopy.
They have also been observed tail-chasing: pursuing another bird in
flight, rapidly dodging among trees and branches, a predation style
common to hawks (genus Accipiter) that hunt birds.[12]
A recent literature review and research using camera traps list a total of 116 prey species.[35][36] Its main prey are tree-dwelling mammals, and a majority of the diet has been shown to focus on sloths.[37] Research conducted by Aguiar-Silva between 2003 and 2005 in a nesting site in Parintins, Amazonas, Brazil,
collected remains from prey offered to the nestling by its parents. The
researchers found that 79% of the harpy's prey was accounted for by
sloths from two species: 39% brown-throated sloth (Bradypus variegatus), and 40% Linnaeus's two-toed sloth (Choloepus didactylus).[38]
Similar research in Panama, where two captive-bred subadults were
released, found that 52% of the male's captures and 54% of the female's
were of two sloth species (brown-throated sloth and Hoffmann's two-toed sloth (Choloepus hoffmanni).[39] Harpy eagles are capable of hunting all size of sloths, including full-grown adult two-toed sloths weighing up to 9 kg (20 lb).[40]
Along with sloths, monkeys, such as the tufted capuchin (Cebus appella), are one of the main prey of the harpy eagle[41]
prey on all ages and sexes, while male harpy
eagles tend to focus on juveniles.[15][45][46][47][48] In one study, breeding harpy eagles hunted Yucatán black howler (Alouatta pigra),
the largest howler monkey which can weigh between 6.4 and 11.3 kg (14
and 25 lb), although the ages of the monkeys taken by these eagles are
unknown.[49][50] Nevertheless, adults of other large monkeys can be taken by female harpy eagles, including woolly monkey (Lagothrix cana) and Peruvian spider monkey (Ateles chamek), and red-faced spider monkey (Ateles paniscus) which can weigh around 5.8 to 9.4 kg (13 to 21 lb) and possibly exceeding 10 to 11 kg (22 to 24 lb) in large males.[35][36][51][44][52]
Other partially arboreal and even land mammals are also preyed on given the opportunity. In the Pantanal, a pair of nesting eagles preyed largely on the porcupine (Coendou prehensilis) and the agouti (Dasyprocta azarae).[53] Both species of tamanduas (Tamandua mexicana & T. tetradactyla) are taken and armadillos, especially nine-banded armadillo (Dasypus novemcinctus) are also taken,[35][36] as well as carnivores such as kinkajous (Potos flavus), coatis (Nasua nasua & N. narcia), tayras (Eira barbara), and occasionally margays (Leopardus wiedii) and crab-eating foxes (Cerdocyon thous).[12][35] In one instant, an adult greater grison (Galictis vittata) was killed and partly consumed by subadult female harpy eagle.[54] Those carnivoran prey species usually weigh around 1.4 to 7.2 kg (3.1 to 15.9 lb),[55][52] but there is a report that harpy eagles prey on possibly larger carnivores such as ocelot (Leopardus pardalis) and adult crab-eating raccoon respectively.[51][15] Other mammals, such as young peccaries, deer fawns, squirrels and opossums are additionally taken.[12]
The eagle may also attack bird species such as macaws: At the Parintins research site, the red-and-green macaw (Ara chloropterus) made up for 0.4% of the prey base, with other birds amounting to 4.6%.[38][56] Other parrots have also been preyed on, as well as cracids such as curassows and other birds like seriemas.[12] In one occasion, dependent juvenile male eagle quickly learned how to hunt black vultures (Coragyps atratus) and accounted for 9 of our 10 records of harpy predation on vultures.[36] Additional prey items reported include reptiles such as iguanas, tegus, snakes, and amphisbaenids.[12][16] In Suriname, green iguanas (Iguana iguana) can be important prey source, and predation on yellow-footed tortoise (Chelonoidis denticulata) have been recorded twice.[35]
The eagle has been recorded as taking domestic livestock, including chickens, lambs, goats, and young pigs, but this is extremely rare under normal circumstances.[12] They control the population of mesopredators
such as capuchin monkeys, which prey extensively on bird's eggs and
which (if not naturally controlled) may cause local extinctions of
sensitive species.[57]
Males usually take relatively smaller prey, with a typical range
of 0.5 to 2.5 kg (1.1 to 5.5 lb) or about half their own weight.[12]
The larger females take larger prey, with a minimum recorded prey
weight of around 2.7 kg (6.0 lb). Adult female harpies regularly grab
large male howler or spider monkeys or mature sloths weighing 6 to 9 kg
(13 to 20 lb) in flight and fly off without landing, an enormous feat of
strength.[12][58][59] Prey items taken to the nest by the parents are normally medium-sized, having been recorded from 1 to 4 kg (2.2 to 8.8 lb).[12]
The prey brought to the nest by males averaged 1.5 kg (3.3 lb), while
the prey brought to the nest by females averaged 3.2 kg (7.1 lb).[29]
In another study, floaters (i.e. birds not engaging in breeding at that
time) were found to take larger prey, averaging 4.24 kg (9.3 lb), than
those that were nesting, for which prey averaged 3.64 kg (8.0 lb), with
prey species estimated to weigh a mean of 1.08 kg (2.4 lb) (for common opossum) to 10.1 kg (22 lb) (for adult crab-eating raccoon).[15]
Overall, harpy eagle prey weigh between 0.3 and 6.5 kg (0.66 and
14.33 lb), with the mean prey size equalling 2.6 ± 0.8 kg (5.7 ± 1.8 lb)
[60]
Harpy eagle in flight
In
ideal habitats, nests would be fairly close together. In some parts of
Panama and Guyana, active nests were located 3 km (1.9 mi) away from one
another, while they are within 5 km (3.1 mi) of each other in
Venezuela. In Peru, the average distance between nests was 7.4 km
(4.6 mi) and the average area occupied by each breeding pairs was
estimated at 4,300 ha (11,000 acres). In less ideal areas, with
fragmented forest, breeding territories were estimated at 25 km (16 mi).[16]
The female harpy eagle lays two white eggs in a large stick nest, which
commonly measures 1.2 m (3 ft 11 in) deep and 1.5 m (4 ft 11 in) across
and may be used over several years. Nests are located high up in a
tree, usually in the main fork, at 16 to 43 m (52 to 141 ft), depending
on the stature of the local trees. The harpy often builds its nest in
the crown of the kapok tree,
one of the tallest trees in South America. In many South American
cultures, cutting down the kapok tree is considered bad luck, which may
help safeguard the habitat of this stately eagle.[61] The bird also uses other huge trees on which to build its nest, such as the Brazil nut tree.[62] A nesting site found in the Brazilian Pantanal was built on a cambará tree (Vochysia divergens).[63]
No display is known between pairs of eagles, and they are
believed to mate for life. A pair of harpy eagles usually only raises
one chick every 2–3 years. After the first chick hatches, the second egg
is ignored and normally fails to hatch unless the first egg perishes.
The egg is incubated around 56 days. When the chick is 36 days old, it
can stand and walk awkwardly. The chick fledges at the age of 6 months,
but the parents continue to feed it for another 6 to 10 months. The male
captures much of the food for the incubating female and later the
eaglet, but also takes an incubating shift while the female forages and
also brings prey back to the nest. Breeding maturity is not reached
until birds are 4 to 6 years of age.[12][29][32] Adults can be aggressive toward humans who disturb the nesting site or appear to be a threat to their young.[64]
Although the harpy eagle still occurs over a considerable range, its
distribution and populations have dwindled considerably. It is
threatened primarily by habitat loss
due to the expansion of logging, cattle ranching, agriculture, and
prospecting. Secondarily, it is threatened by being hunted as an actual
threat to livestock and/or a supposed one to human life, due to its
great size.[65]
Although not actually known to prey on humans and only rarely on
domestic stock, the species' large size and nearly fearless behaviour
around humans reportedly make it an "irresistible target" for hunters.[16] Such threats apply throughout its range, in large parts of which the bird has become a transient sight only; in Brazil, it was all but wiped out from the Atlantic rainforest and is only found in appreciable numbers in the most remote parts of the Amazon basin;
a Brazilian journalistic account of the mid-1990s already complained
that at the time it was only found in significant numbers in Brazilian
territory on the northern side of the Equator.[66] Scientific 1990s records, however, suggest that the harpy Atlantic Forest population may be migratory.[67]
Subsequent research in Brazil has established that, as of 2009, the
harpy eagle, outside the Brazilian Amazon, is critically endangered in Espírito Santo,[68]São Paulo and Paraná, endangered in Rio de Janeiro, and probably extirpated in Rio Grande do Sul (where a recent (March 2015) record was set for the Parque Estadual do Turvo) and Minas Gerais[69] – the actual size of their total population in Brazil is unknown.[70]
Globally, the harpy eagle is considered vulnerable by IUCN[2] and threatened with extinction by CITES (appendix I). The Peregrine Fund
until recently considered it a "conservation-dependent species",
meaning it depends on a dedicated effort for captive breeding and
release to the wild, as well as habitat protection, to prevent it from
reaching endangered
status, but now has accepted the near threatened status. The harpy
eagle is considered critically endangered in Mexico and Central America,
where it has been extirpated in most of its former range; in Mexico, it
used to be found as far north as Veracruz, but today probably occurs only in Chiapas in the Selva Zoque.
It is considered as near threatened or vulnerable in most of the South
American portion of its range; at the southern extreme of its range, in
Argentina, it is found only in the Parana Valley forests at the province of Misiones.[71][72] It has disappeared from El Salvador, and almost so from Costa Rica.[73]
Various initiatives for restoration of the species are in place in various countries. Since 2002, the Peregrine Fund initiated a conservation and research program for the harpy eagle in the Darién Province.[74] A similar—and grander, given the dimensions of the countries involved—research project is occurring in Brazil, at the National Institute of Amazonian Research, through which 45 known nesting locations (updated to 62, only three outside the Amazonian basin
and all three inactive) are being monitored by researchers and
volunteers from local communities. A harpy eagle chick has been fitted
with a radio transmitter that allows it to be tracked for more than three years via a satellite signal sent to the Brazilian National Institute for Space Research.[75] Also, a photographic recording of a nest site in the Carajás National Forest was made for the Brazilian edition of National Geographic Magazine.[76]
Adult at São Paulo Zoo, Brazil
In Panama, the Peregrine Fund carried out a captive-breeding and release project that released a total of 49 birds in Panama and Belize.[77] The Peregrine Fund
has also carried out a research and conservation project on this
species since the year 2000, making it the longest-running study on
harpy eagles.[14][78]
In Belize, the Belize Harpy Eagle Restoration Project began in 2003 with the collaboration of Sharon Matola, founder and director of the Belize Zoo and the Peregrine Fund. The goal of this project was the re-establishment of the harpy eagle within Belize. The population of the eagle declined as a result of forest fragmentation, shooting, and nest destruction, resulting in near extirpation of the species. Captive-bred harpy eagles were released in the Rio Bravo Conservation and Management Area in Belize, chosen for its quality forest habitat and linkages with Guatemala and Mexico. Habitat linkage with Guatemala and Mexico
were important for conservation of quality habitat and the harpy eagle
on a regional level. As of November 2009, 14 harpy eagles have been
released and are monitored by the Peregrine Fund, through satellite telemetry.[79]
In January 2009, a chick from the all-but-extirpated population in the Brazilian state of Paraná was hatched in captivity at the preserve kept in the vicinity of the Itaipu Dam by the Brazilian/Paraguayan state-owned company Itaipu Binacional.[80]
In September 2009, an adult female, after being kept captive for 12
years in a private reservation, was fitted with a radio transmitter
before being restored to the wild in the vicinity of the Pau Brasil National Park (formerly Monte Pascoal NP), in the state of Bahia.[81]
In Colombia,
as of 2007, an adult male and a subadult female confiscated from
wildlife trafficking were restored to the wild and monitored in Paramillo National Park in Córdoba, and another couple was being kept in captivity at a research center for breeding and eventual release.[83] A monitoring effort with the help of volunteers from local Native American communities is also being made in Ecuador, including the joint sponsorship of various Spanish universities[84]—this effort being similar to another one going on since 1996 in Peru, centred around a native community in the Tambopata Province, Madre de Dios Region.[85] Another monitoring project, begun in 1992, was operating as of 2005 in the state of Bolívar, Venezuela.[86]
The harpy eagle is rare throughout its range, which extends from Mexico to Brazil (throughout its territory)[4] and Argentina (only the north). (note: map distribution in Trinidad and Tobago and ABC islands is erroneous)
The crowned eagle, also known as the African crowned eagle or the crowned hawk-eagle (Stephanoaetus coronatus), is a large bird of prey found in sub-Saharan Africa; in Southern Africa, it is restricted to more easterly areas.[2] Its preferred habitats are principally riparian woodlands and various forests.[2] The crowned eagle is the only extant member of the genusStephanoaetus. A second species, the Malagasy crowned eagle (Stephanoaetus mahery), went extinct after early humans settled on Madagascar.[3]
Although the crowned eagle's long tail imparts an overall length
of up to 90 cm (35 in), it is lighter by weight, and has a considerably
shorter wingspan, than Africa's largest eagle, the martial eagle (Polemaetus bellicosus). It is, nevertheless, considered Africa's most powerful eagle with respect to its prey’s body size.[6] It has been known to prey on ungulates as large as bushbuck (Tragelaphus scriptus), which can weigh up to 30 kg (66 lb), albeit usually much less.[6][7]
The crowned eagle possesses unusually large talons and strong legs, and
may kill by simply crushing prey’s skull. The eagle is also bold and
ferocious; records documented from beneath a nest show the remains of a
large male sooty mangabey weighing 11 kg (24 lb).[8]
Due to their ecological and behavioral similarities, the crowned
eagle is considered to be the African counterpart of the Central and
South American harpy eagle (Harpia harpyja).
Thanks to its bold and highly conspicuous behavior, it is exceptionally
well-studied for a large, forest-dwelling eagle. Due to a relatively
high level of habitat adaptability, it was until recently considered to
be faring well by the standards of large, forest-dependent raptors.[9]
However, today it is generally thought that it is decreasing far more
than was previously perceived due to the almost epidemic destruction of
native tropical African forest.[1] It is now listed by the IUCN as Near Threatened.
Taxonomy and systematics
An adult crowned eagle in flight.
This species was first described by Carl Linnaeus in the 12th edition of Systema Naturae, published in 1766, as Falco coronatus.
Since birds were grouped largely on superficial characteristics at that
time, many unrelated species were grouped by Linnaeus in the genus Falco.
The actual taxonomic alignment of the crowned eagle is apparent due to
its feathering over its tarsus, which is generally rare in unrelated
accipitrids. The crowned eagle is actually part of the diverse "booted eagle" group, which has sometimes been considered a distinct subfamily (Aquilinae). Included in this grouping are the genus Aquila and all species described as "hawk eagles" including the genera Spizaetus and Nisaetus. Other assorted monotypical genera included amongst "booted eagles" are Lophaetus, Polemaetus, Lophotriorchis and Ictinaetus.
At one time, the genus Stephanoaetus was considered a "specialized offshoot" of the Spizaetus hawk-eagles based on morphological attributes.[10] DNA sequences utilizing one mitochondrial and three nuclear genes indicated the crowned eagle is a sister species to the Asian hawk-eagles, which are now considered a separate genus, Nisaetus, that are not closely related to the neotropical hawk-eagles, which are retained in Spizaetus.[11]
However, another recent study, this time of sequences of two
mitochondrial and one nuclear genes, did not reveal a close relationship
of this eagle to any other accipitrid, including the Nisaetus species, and the genus was found to be genetically highly divergent from other "booted" eagles".[12] In a case of convergent evolution, the much heavier harpy eagle, which is outside of the "booted eagle" group, has a similar skeletal morphology to the crowned eagle.[6] Two less well-known, probably distantly related species, the mountain hawk-eagle (Nisaetus nipalensis) and the black-and-chestnut eagle (Spizaetus isidori), have also been found comparable to the crowned eagle.[10]
While both are slimmer and smaller, these eagles are also large-bodied,
strong-footed offshoots of the evolutionary radiation of
forest-dwelling booted eagles, respectively distributed in East Asia and South America.
The adult crowned eagle even has somewhat intermediate appearance
between these birds, sharing the variable patterning of the mountain
hawk-eagle and some of the colouring of the black-and-chestnut.[6][10] Until possibly up to 1500, another crowned eagle species, the Malagasy crowned eagle (S. mahery) existed.[13] Similar in size and form to the extant crowned eagle, the Malagasy crowned eagle probably filled a similar niche in Madagascar, but was likely to have preyed on lemurs in place of monkeys.[3][13]
Apparently, the Malagasy crowned eagle became extinct due mainly to the
loss of prey and habitat change, attributable to early humans on the
island.[14][15]
To date, the living crowned eagle has no recognized subspecies.
However, Simon Thomsett noted from field experience possible racial
differences between crowned eagles in limited woodland habitats in East
and South Africa (called by him the "bush eagles"), which have
historically been the main populations studied, and those that live in
denser West African rainforest, in the central part of the species
distribution. The latter population, he noted, appeared smaller but
relatively larger footed, seemed chestier in build and appeared to have
deeper eyebrows than the bush eagle; behaviorally the rainforest eagles
seemed bolder and louder, which is reinforced in other accounts of the
species.[7][16]
Description
Immature Kakamega Forest, KenyaA captive crowned eagle, showing the extended crest and the permanent fierce stance of the species
The crowned eagle is a very large eagle. Measuring from 80 to 99 cm
(31 to 39 in) in length, it is the fifth longest extant eagle in the
world.[7]
The female, at a weight of 3.2–4.7 kg (7.1–10.4 lb), is around 10–15%
larger than the male, at a weight of 2.55–4.12 kg (5.6–9.1 lb).[6][7] An average body mass of 3.64 kg (8.0 lb) was given in one account.[17] Elsewhere, an average of 3.8 kg (8.4 lb) has been claimed.[8]
Slightly smaller weights were found in a South African survey where 5
males averaged 2.65 kg (5.8 lb) and 8 females averaged 3.71 kg (8.2 lb).[18] Overall, they are the 9th heaviest living eagle species.[7]
The wingspan typically ranges from 1.51 to 1.81 m (4 ft 11 in to 5 ft
11 in). The largest authenticated wingspan for a female was 1.9 m (6 ft
3 in),[19] with a claim of wingspans of up to 2 m (6 ft 7 in) needing confirmation.[20] This eagle's wingspan is quite short for the bird's size, being around the same mean width as that of a tawny eagle (Aquila rapax) or a short-toed snake eagle (Circaetus gallicus),
species that weigh about half as much as a crowned eagle. However, the
somewhat boxy and rounded wings are quite broad, being broader than, for
example, the much longer-winged golden eagle (Aquila chrysaetos).[21] The wing morphology of the species gives it maneuverability in its densely wooded environment. The wing chord measures 445–532 mm (17.5–20.9 in), with a median of 467 mm (18.4 in) in males and 512 mm (20.2 in) in females.[7][10] In South Africa, 5 males averaged 475.2 mm (18.71 in) in wing chord length and 7 females averaged 506.9 mm (19.96 in).[18] While it, on average, is less heavy and has a smaller wingspan than the often sympatric martial eagle,
its average total length exceeds that of the martial eagle thanks to
its much longer tail. The crowned eagle's tail is from 300 to 410 mm (12
to 16 in) long, with a median of 315 mm (12.4 in) in males and 348 mm
(13.7 in) in females.[7][10] South African males averaged 320.4 mm (12.61 in) in tail length in a sample of 4 and females averaged 352.4 mm (13.87 in).[18] The bill
is of a medium size relative to its body size, with one large museum
specimen's bill measuring 55 mm (2.2 in) in length from the gape, 45 mm
(1.8 in) in culmen length and 33 mm (1.3 in) in bill depth.[22]
In South Africa, culmen length averaged 50 mm (2.0 in) in 4 males and
54.9 mm (2.16 in) in 7 females, with a range in both 46.5 to 61.4 mm
(1.83 to 2.42 in).[18]
A zoo-kept adult crowned eagle in Hungary illustrates its formidable talons.
The tarsus is of a modest length for a raptor of its size, at 8.5–10.3 cm (3.3–4.1 in), and is clearly shorter than that of martial eagle.[7] However, the feet and legs are visibly thicker and heavier than those of the martial eagle and the talons
are apparently quite massive in both length and width. While few
comprehensive measurements of the talon size of wild crowned eagles are
known, one female museum specimen reportedly had a hallux-claw (or hind claw, which is the largest talon on accipitrids) of 6.2 cm (2.4 in),[22]
while another female was measured at 5 years of age, the age of sexual
maturity, 5.74 cm (2.26 in) in the hallux-claw and an adult male
measured 4.9 cm (1.9 in).[23]
In South Africa, hallux-claw length averaged 52.4 mm (2.06 in) in 5
males and 60 mm (2.4 in) in 7 females with a range in both of 48.6 to
61.4 mm (1.91 to 2.42 in).[24] These figures put their talon size as around the same size as the largest golden eagles and half the size of a harpy eagle.[25][26]
Some captive crowned eagles have been credited with a hallux-claw
length of up to 10 cm (3.9 in), although, much like a single report of
captive harpy eagles with a 13 cm (5.1 in) hallux-claw, no such outsized
talons are known to have been confirmed.[16][27]
In a small sampling of large, forest-dwelling raptors, the front-left
talon of the crowned eagle, at 4.74 cm (1.87 in), was around one cm less
than that of a harpy eagle or the huge, recently extinct Haast's eagle (Hieraaetus moorei) and slightly smaller than those of the Philippine eagle (Pithecophaga jefferyi).[28]
Considering a big female of these species can weigh up to twice as much
as an average crowned eagle may illustrate the relative largeness of
the crowned eagle's talons.[17]
The adult crowned eagle is quite strikingly plumaged. Its crown
is dark to rufous-tinged brown with a prominent, oft-raised black-tipped
double crest, which can give the head a somewhat triangular appearance.[7][10]
The upperparts of an adult are a blackish brown-grey color, with a
variable tinge of blue. The throat is brown while the belly and breast
are white overlaid densely with blackish bars and blotches, variably
marked with cream or rich buff-rufous coloration. The wing primaries
are white at the base, broadly tipped with black and crossed by two
black bars. The tail is black with brownish-grey bands. The thighs and
legs are barred and closely spotted with black and white. The underwing
coverts of adults have a bold chestnut coloration, spotted lightly with
black.[7][10]
The adult crowned eagle has eyes that can range from yellow to almost
white, a cere and feet of an ochre-yellow color and black talons.[10] In the wild, misidentification of an adult is improbable thanks to the species' bearing and voice.[7] The strongly barred outer wings and tail are all diagnostic in flight.[2]
Further simplifying identification, details such as the crest, the
bird's upright perching posture and large size are unique to this
animal.[7]
While they do differ somewhat in size, the genders' sexual dimorphism
by size is relatively modest and eagles are unlikely to be sexed by this
alone. However, the male may be distinguished by his more rapid wing
beats (4 or 5 per second) from the more sluggish female (3 or 4 per
second).[10]
As seen in about half of the "booted eagle" group, the juvenile
crowned eagle has strikingly different looking plumage compared to the
adults. Much variation occurs as the maturation process occurs. A great
majority of juveniles have a white head and underside, which contrast
with the thighs and legs, which are heavily spotted with black.[7]
The juvenile eagle's back is light brown or grayish-brown, with pale
feather edgings that often give the back a scaled appearance, especially
on the upper-wing coverts. There is often a pinkish red wash on the
upper chest.[29]
Just-fledged chicks tend have dark patched faces, freckled bibs and
slightly barred chests and spotted legs. Less common juvenile crowned
eagle plumages, possible even when they are under a year of age and
still under parental care, may include eagles so stripy that they which
one could easily have aged as two- to three-year-olds.[16]
The tail of the juvenile is black with three pale bars and a narrow
black tip. The juvenile eagle's cere is grey and the feet are dull
yellow.[10]
By 4 months post-fledgling, the inner thighs, previously poorly covered
with downy type feathers, are covered with small feathers. While the
pale 'morph' young just prior to leaving the nest usually have unmarked
tarsus, they soon get spots on the front part of the tibio tarsal joint.
The tibio tarsal pad is still bare and obvious up until it is a year
old, whereupon it vanishes only to return to incubating females.[16]
Eye color is variable too with some having khaki light brown just prior
to fledging and others with adult-like yellow ochre eyes.[16]
Up to 15 months after leaving the nest, the immature eagles more
closely resemble the plumage they have at first independence than the
adult's plumage.[10] The juvenile may be confused with the similarly colored juvenile martial eagle, especially in flight.[7]
It is distinguished from the martial species in having a much longer,
more heavily barred tail, much shorter wings and spotted thighs.[2]
Distribution and habitat
Humid Congolian forests along the Dja River, typical habitat near the center of the crowned eagle's range
The crowned eagle inhabits mainly dense woodlands, including those deep within rainforest, but will sometimes also be found in relict patches, wooded escarpments, riparian strips of Acacia,
heavily wooded hillsides, and rocky outcrops throughout its range. The
crowned eagle may be found from an altitude of sea-level to at least
3,000 m (9,840 ft). Owing to lack of current suitable habitat, the
eagle's range is often somewhat discontinuous.[7] In the Democratic Republic of the Congo,
the crowned eagle has been confirmed to survive at relatively high
densities in protected areas that maintain dense, old-growth
rainforests.[30] In Kenya, 84% of the crowned eagles range is within rainforest with an annual rainfall amount of more than 150 cm (59 in).[31] Around stretches of East Africa
where protected areas mostly consist of fairly open habitat, crowned
eagles usually live in wooded areas of rocky hills and narrow riverine
strips, only rarely ranging into savanna surrounding the hills.[10]Southern Africa
has been subject to the most comprehensive study of crowned eagle
habitat, largely since many areas there would seem inhospitable to a
large raptor often associated with old-growth forest.[32] In southern Africa, its distribution south of the Limpopo River coincides largely with montane forest, although it is not restricted to that habitat and may range secondarily into plantations, usually of eucalyptus.[7][32] In South Africa, it occurs in both lowland and montane evergreen forest, dense woodland, and forested ravines and gorges in open savannas and thornveld.[33] In Zimbabwe, the crowned eagle can be found in quite open woods with Adansonia trees and may occasionally forage in savanna and secondary growth.[7] In Malawi, highland birds forage in lower miombo woodland,
and lower altitudes, breeding occurs in deciduous forest, more locally
in dense miombo, tall riparian woodlands, and in remnants close to
cultivation.[9] Crowned eagles in Zambezi,
occurs in evergreen forest in the eastern highlands, in rugged, hilly
terrain over the central plateau, in hills and escarpments in the
southeastern portions of the central watershed, and in riparian habitat
along the larger rivers.[34]
Behavior
A crowned eagle flying over its territory in Budongo, western Uganda
Typical of most raptors that breed in Africa, the crowned eagle is non-migratory and is largely sedentary.[32]
This species usually inhabits a fixed territory throughout the year
during its adult life. There is evidence that the birds move about to
some degree when circumstances require it, for example when they need to
change mates in isolated breeding areas.[10]
The greatest movement of any notable distance is usually associated
with juvenile birds, which wander relatively widely before maturation.[32][35] Forty-four birds of various ages that were ringed in southern Africa and recovered were all found near the ringing localities.[32]
Small movements by crowned eagles has resulted in them being sighted in
improbable habitats, such as on an open savanna hill in Kenya, an open river bed in Lewa Wildlife Conservancy and even a golf course in Nairobi.[16]
The crowned eagle is highly vocal and has a noisy, undulating display flight. In Equatorial Africa, they often call year-around, while elsewhere they may vocalize mainly in the context of breeding and nesting activities.[7] The call is a series of loud whistles that rise and fall in pitch.[7]
The male performs an elaborate rise-and-fall display over the forest
canopy both during the breeding season and outside it as a territorial
proposition.[7]
Usually, territorial displays, which outnumber breeding displays, occur
around the periphery of the bird's home range while breeding displays
are likely to be over or at least near the nest.[10]
Displays consists of a series of steep dives and ascents, with a few
wing-flaps at the top of each climb and descending circles and figures
of eight.[10] During descents, eagles can drop as much as 60 m (200 ft) at a time before circling back up.[21] During this display, the male is noisy, uttering a shrill kewee-kewee-kewee while throwing his head back, often calling for a spell of approximately 30 seconds.[7]
The displaying male may reach heights exceeding 900 m (2,950 ft),
sometimes even near cloud level at over 2,000 m (6,560 ft) above the
ground.[10] The adult female may also perform independent display flights, uttering a lower kooee-kooee-kooee. The female seems to display less often and tends to have a mellower voice.[10]
Pairs also perform visually striking mutual displays, sometimes arising
from the first type or when the pairs come together after a brief
absence.[10] Spectacular tandems, interlocking talons and falling some distance from the sky are typical of mutual displays.[7]
Despite the spectacular and conspicuous nature of their display
flights, the crowned eagle is sometimes considered a "clumsy" flier out
in the open, lacking the grace and speed attributed to other eagles in a
high-flying context, such as large Aquila species.[21] While awaiting food at the nest, both the female and the young call out a penetrating, high kwee-kwee-kwee, that can border on incessant.[7]
In disposition, the crowned eagle is considered nervous, constantly alert and on edge.[16] They are quite different from the oft-stately open country eagles such as the Aquila species. In their training and management, crowned eagles are perhaps more reminiscent of a Goshawk (Accipiter gentilis) than Aquila eagles.[16]
Some linkage has been made between forest-dwelling habits and having a
curious and edgy disposition amongst various raptorial birds.[10][16]
Ironically considering their otherwise hyperactive behavior, the main
hunting techniques of crowned eagle require long periods of inactivity,
spent sitting on a perch.[21]
Mature crowned eagles are reportedly nearly fearless towards humans
and, unless shy from prior interactions, unusually prone to treat humans
aggressively.[21]
Crowned eagles can nest around developed areas, including in the
vicinity of quite suburban, developed areas (such as in view of
apartment or office buildings), so long as prey is abundant and
accessible and the habitat provides enough mature vegetation to
facilitate nest building and hunting activities.[21]
Some biologists consider this species highly intelligent, cautious,
independent and inquisitive when compared to other accipitrids.[16] In falconry,
crowned eagles cannot be induced to direct their hunting instinct
towards large prey by increasing their hunger, as is done with Aquila eagles, for example.[16] In the context of human interaction, wild, adult crowned eagles are quite the opposite of the martial eagle, which are typically very wary and tends to shun any variety of human activity.[7]
However, young crowned eagles in their post-fledging stage differ
greatly in behavior from independent or adult eagles. Amongst
post-fledging eagles in a semi-captive state, it has been noted that
they border on helpless in terms of feeding and defending themselves
compared to other accipitrids and are even described as "cowardly",
unwilling to even simulate attacking prey until many months after
fledging. This implies a learning element occurs in wild crowned eagles
during their exceptionally long post-fledging period.[16] Crowned eagles are reportedly variable in temperament as individuals to a degree greater than that found in most other raptors.[16]
Crowned
eagles have one of the most prolonged breeding cycles of any bird. It
is common for raptors that live around the tropics to have a relatively
elongated breeding period.[7] Crowned eagle pairs breed once every two years; a single breeding cycle is approximately 500 days in duration.[10] Most other eagle species complete a breeding cycle in under six months, or in about 35% of the time it takes the crowned eagle.[21] While the incubation and nestling stages are about average for a tropical eagle (for example the black-breasted snake-eagle (Circaetus pectoralis),
about half the weight of this species, has an incubation/nestling cycle
of a similar length), it is the extraordinary post-fledging period of
9–11 months that makes the crowned eagles' breeding cycle so long.[21] In harpy and Philippine eagle,
although these are less extensively studied, it may take a similar or
even longer amount of time for the young to attain full independence.[36][37]
A case of crowned eagle pairs that reportedly bred every year in South
Africa are unsubstantiated but may be due to an apparently high
population loss rate among juvenile eagles near areas that are heavily
populated by humans.[38][39]
Breeding can occur almost year-around throughout the range, though egg
laying seems to peak roughly around the end of the African wet season or the early dry season, from July to November.[7] Territories or home ranges are maintained vigorously. In Zimbabwe, individual home ranges can vary from 140 to 200 km2 (50 to 80 sq mi) in size.[34] Near the city of Nelspruit in South Africa, home ranges averaged 30 km2 (10 sq mi) in size.[38] In southern Africa, the mean distance between active nest sites can range from 2 to 19.5 km (0 to 10 mi).[33]
After engaging in the breeding display described above, the pair
collaborate in building a massive nest in a fork of a large forest tree,
typically from 12 to 45 m (40 to 150 ft) above the ground. While the
female fetches more nesting material, the male tends to be more active
in nest construction.[10] In East Africa, many nests appear to be close to a forest river.[10][40] Generally, crowned eagles seem to be attracted to the taller trees in the forest.[38] On the Nyika Plateau in Malawi, the favorite nesting trees are the large emergent Aningeria adolfi-friederici and Gambeya gorungosana, and a pair in the Lower Shire used a Sterculia appendiculata tree.[9] In Zimbabwe, Newtonia buchananii are reportedly one of the most used tree species for nesting.[34] Exceptional crowned eagle nests have been observed on sheer cliff faces.[41] In southern Africa, the species nest in drier and denuded terrain than expected such as Adansonia
stands on semi-arid hillsides. Despite the relative sparseness of this
habitat, these sites have a varied and convoluted terrain, with nooks
and crannies, valleys, overhangs and hideaways that allow a crowned
eagle to exercise its particular hunting skills. In Kenya, similar
fractured landscapes can also be utilized by crowned eagles, such as the
black gigantic volcanic rubble fields of Tsavo West National Park, the lower Chyulu Hills, Kibwezi and Soysambu Conservancy. These are jungles of boulders covered with low growth interspersed (in the past) with high trees.[16]
A nest built from scratch may take up to 5 months to construct, however
existing nests are often repaired and re-used during successive
breeding seasons, a process that can take as much as 3 months.[10]
It is typical for an eagle pair to use a nest for more than five years
and, unlike several other booted eagles, crowned eagle pairs rarely
build more than one nest for alternative use.[21]
Most large eagles build a very large nest and the crowned eagle is no
exception, as it builds one of the largest nests of any eagle. In the
first year they build a nest, it may measure 1.5 m (4.9 ft) across and
50 cm (20 in) deep. However, a larger nest, usually after several years
of usage, may measure up to 2.5 m (8.2 ft) across and up to 3 m (9.8 ft)
deep. The nest consists of both dead and greener branches and has a
light coverage of leaves and animal matter.[7]
Copulation takes place in the nest, several times a day. Reportedly
copulation can occur up to a year before laying, although these may be
exceptional cases of mating for non-fertilization purposes (which, in
other eagles, has been thought to be related mainly to strengthening
pair bonds).[10]
A pre-copulation display typically occurs, wherein the male runs
repeatedly around the crouching female with wings upraised, which
displays the chestnut of the under-wing coverts and beautiful barring.[10]
In South Africa the crowned eagle lays its eggs from September to October; in Zimbabwe, it lays from May to October; mainly nearer to October around the Congo River; anywhere from June to November in Kenya, with a peak in August through October; in Uganda from December to July; and in West Africa, laying peaks in October.[10] The clutch of the crowned eagle either contains 1 or 2 eggs. Often in East Africa, just one egg is laid.[10]
Eggs are usually just white, though may sometimes be overlaid with
sparse red-brown markings. The eggs are moderate in size, averaging
68.2 mm × 53.6 mm (2.69 in × 2.11 in), with ranges of 60.9–75.5 mm
(2.40–2.97 in) in length and 50.8–57.9 mm (2.00–2.28 in) in width.[10] When a natural disaster befalls a nest, a replacement may be made in 2 months time.[10] Incubation lasts for approximately 49 days. 80–90% of egg incubation is done by the female during the day.[10]
Food is mainly brought to the nest by the male in the early stages of
breeding, though sometimes both sexes may deliver food. The male brings
food to the incubating female every 3 to 5 days. When they initially
hatch, the young tend to be quite quiet. If two eggs are laid, the
younger one dies by starvation after being outcompeted for food by the
older one or even directly killed by its older sibling.[7]
No nest of wild crowned eagles has been known to successfully produce
more than one fledgling, though in captivity two have been known to
survive with human assistance (supplementally feeding the younger chick
or taking it out of the nest).[16][38] In cases where the older nestling dies, the younger one may be fed more regularly and survive.[7]
After hatching, the male kill rate rises to one kill every 1.5
days. Pair behavior while raising chicks is very variable, some males
are very attentive to their young, while others leave virtually all
brooding to the female. After 40 days of age, the young is capable of
feeding itself, though is often still fed. The first feathers through
the white down emerge when the crowned eagle chick is 40 days old, with
the feathers ultimately covering the down in 76 days. After 76 days, the
main feather development is in the tail or the wings. Wing flapping
begins at 45 to 50 days, increasing after around 75 days.[10]
The young fledge at 90 to 115 days, with an average 110.6 days and any
period of time less than 100 days is considered unusually soon. On
average, male chicks tend to be more active wing-flappers and usually
will first fly around 10 days earlier than female chicks.[7]
After fledging, females are attentive 95% of the day and brood 50–75%
of the day, the amount decreasing slightly with each day. The female
does much of the prey capture and a majority of the nest defense after
the young fledge. After fledging, the young remain in the neighborhood
of the parent's nest and are fed every 3 to 5 days by either parent for
their first 270–350 days of life.[10]
The rate of food-delivery varies from several times a day to every 3
days on average during the post-fledging period. The fledged juvenile
will solicit adults (apparently even unrelated adults) for food but does
not actually take the prey unless this occurs around the nest site.[10]
The first recorded kill for a young crowned occurred 61 days after
fledging, although this is considered exceptionally early by the
standards of this species.[10]
Flights increase incrementally through the post-fledging period,
although the young do not engage in rising flights until they are fully
independent. Independence appears to be triggered by the increased
indifference of parents to bringing food. Due to the loud vocal
interplay between the parents and the fledging eagle, the adults seem to
take it as a sign that their offspring has sought independence if they
return to the nesting area and hear no begging auditory response.[21]
The young eagle usually remains in the care of its parents for a total
of up to 11 months after it fledges, longer than is known in almost any
other raptor. The advantage of this prolonged stretch to independence is
that it may make for a stronger young eagle when compared to other
accipitrids which have almost no post-fledging dependency period.[21]
In 34 possible cases, 18 resulted in eggs being laid. Fledging success
is approximately 83% and almost all young that leave the nest also reach
independence.[10]
It is estimated that most crowned eagles will reach breeding maturity at
around five years old, as is typical for other large eagle species.[21]
The
crowned eagle is often described as the most powerful raptor in Africa,
even more so than the two slightly heavier species endemic to Africa,
the martial eagle and the Verreaux's eagle (Aquila verreauxii).[10][41]
One listing included the crowned eagle as the only bird in a ranking of
the 10 strongest living land creatures (pound for pound).[42] Elsewhere, the harpy eagle is listed as the overall most powerful living eagle and bird of prey.[7][43]
Since there are no known actual tests in any African raptor of the
pressure exerted via their grip, as has been done with some other large
eagles, their power has been estimated from the size of the feet and
talons and from the prey they typically select.[10]
The crowned eagle's staple diet is mostly mammalian. One estimate
of the typical prey range posited that the weight range is from 1 to
5 kg (2.2 to 11.0 lb), which is, based on the species ecology in Kenyan
hillside woodland. This prey weight range is roughly the same weight
range that's typically attributed to martial or Verreaux's eagles.[44] Perhaps unsurprisingly, that is the weight range of the rock hyrax, which all three large eagles are known to hunt regularly in East Africa.[45] In a large collection of bones in the South African Museum, it was found that 51.2% of the bones collected from Nature's Valley
in South Africa were from smallish species which weigh under 2 kg
(4.4 lb) in adulthood, 26.3% from "medium-sized" species with an adult
weight of 2 to 20 kg (4.4 to 44.1 lb) and 22.5% were from larger species
with an adult weight of over 20 kg (44 lb).[46]
However, about 91% of 87 bones that were from the relatively large
antelope species, amongst those that could be accurately sized, were
from juvenile specimens.[46] On the other hand, in the rainforest community of Taï National Park in the Ivory Coast, the estimated average weight of prey for crowned eagles was clearly higher at 5.67 kg (12.5 lb).[47]
Of all other living eagles, only the female harpy eagle has been
credited with an average prey weight range that is comparably high and,
at the species level, the crowned eagle's prey size from the latter
study is the largest of any known for all extant accipitrids.[7] The crowned eagle is perhaps the only living accipitrid to routinely attack prey weighing in excess of 9 kg (20 lb).[21]
According to some authorities typical prey sizes for crowned eagle may
be considered to range to at least 20 kg (44 lb) in body mass.[8][48][24] Reportedly, the crowned eagle can lift more than its own body weight in flight, though verified accounts of this are sparse.[49]
In a deep forest, an adult eagle may cover a hunting range of up to 6.5 to 16 km2 (0 to 10 sq mi), with the home ranges being smaller for those that inhabit rocky hills and cliffs abounding in hyraxes.[10] Eagles start hunting soon after dawn and mainly kill early in the morning and in the evening prior to sundown.[10]
Being a forest-dwelling species, the crowned eagle has no need to
travel great distances to hunt, nor employ a great deal of active
hunting flight (such as soaring seen in savannah-dwelling species). Rather, it tends to hunt passively.[7] Crowned eagles may locate a suitable hunting spot by listening (such as via the call of the noisy vervet monkey) or watching for prey activity, though may also use habitual hunting perches where they've previously had hunting successes.[7][10]
Although this behavior is unconfirmed, some crowned eagles have been
reported to let out a soft whistle, unlike their other vocalizations
which, for some reason, is attractive to monkeys and will then attack
the first monkey to enter their line of sight.[10]
These eagles often still-hunt, wherein they drop or stoop onto prey
from a branch perch. Following the sighting of suitable prey, the eagle
quickly and stealthily maneuvers itself through the forest towards its
prey, a certain element of surprise inherent in its final approach.[7] A majority of the crowned eagle's kills are made on the forest floor. Arboreal prey may be forced to the ground during an attack.[21] The sharp, powerful talons may produce sufficient force to kill the prey on impact; if not, death from trauma or asphyxiation soon follows.[7] Several prey items have been killed by ramming the talons into the skull and penetrating the brain.[50]
Having killed on the ground, it has the ability to fly almost
vertically upwards to a branch while carrying its prey before feeding,
though it will tear up prey into manageable pieces on the ground when it
is exceedingly heavy.[10]
While they both attack somewhat similar prey in often similar habitat,
the considerable difference in body weight and wing-loading between
crowned and harpy eagles have been attributed to load-carrying while
hunting, as harpies tend to capture and carry off most prey in active
flight rather than an attack on the ground and dismember if necessary.[10]
On rare occasions, crowned eagles may also hunt on the wing,
flying slightly over the canopy and causing a cacophony amongst monkey
groups until they detect and capture their prey, often a monkey or tree
hyrax.[10]
Crowned eagles are believed to take uneaten portions of prey up into
the trees to cache around the nest or habitual perches so that the
pieces can be consumed over the course of the next several days.[7] If the prey is too heavy for taking flight with, even after dismemberment, for example, a bushbuck, crowned eagles have been known to cache food at the thickly vegetated base of a tree and only carry limbs to the nest.[21] Pairs may collaborate in capturing prey, with one bird flushing the prey so the other can glide in unseen and ambush it.[7] Female eagles may target male monkeys more often than males, which are more likely to hunt female or young monkeys.[16] In one case, a female crowned eagle stalked a bushbuck
calf over the course of two days but was repeatedly foiled when it went
in for the attack, either by the mother bushbuck or an associating
troop of yellow baboons (Papio cyanocephalus).
However, the crowned eagle one day assaulted the bushbuck calf quickly,
leaving it with a gaping wound on its flank, and flew off to observe
from a distance. Within a few more days, the bleeding, wounded calf was
unable to keep pace with its mother and was tracked and killed by the
assaulting eagle.[16] Another assault, this one on an adult vervet monkey (Chlorocebus pygerythrus) apparently had a similar outcome as the bushbuck attack.[16] This type of strike-and-wait hunting technique may be used by diverse predators, from Komodo dragons (Varanus komodoensis) to great white sharks (Carcharodon carcharias),
which tend to track their victims by scent after biting them rather
than sight and sound, but is virtually unprecedented in birds.[16] Crowned eagles have been recorded to consume carrion but this behavior has only rarely been observed.[10]
Primate
A Diana monkey (Cercopithecus diana), a typical and located favored monkey in the crowned eagle's dietCrowned eagle with a smallish prey item.
The crowned eagle occupies a unique niche, as it is the only bird in which primates are the most commonly taken prey at the species level.[4] While at least a dozen other accipitrids opportunistically hunt small or young monkeys, only the harpy eagle and, possibly, crested eagle of the neotropics have diets where primates may locally outnumber other prey. However, New World monkeys are generally smaller and less formidable than Old World monkeys. Also, both other eagles may prefer different prey where available: harpy eagles primarily take sloths, which can be as heavy as a colobus or mangabey
but are much slower and less able to defend themselves, while crested
eagles take a variety of tropical birds and arboreal mammals. Other
large eagles have been confirmed to hunt adult Old World monkeys, including martial eagles,[51]Verreaux's eagles,[52]mountain hawk-eagles[53][54] and Philippine eagles, whose generic scientific name and old common was even the monkey-eating eagle,[55] but all are believed to rely on non-primate prey for the majority of their diet.[7]
The favored group in the crowned eagle's diet is certainly the genus Cercopithecus. In Kibale National Park, Uganda, the red-tailed monkey (Cercopithecus ascanius) was the most represented prey species, making up 40% of the remains. Larger monkeys, such as western red colobus (Piliocolobus badius), mantled guereza (Colobus guereza) and grey-cheeked mangabey (Lophocebus albigena) were secondary in the prior study and, altogether, primates made up 82.2% of the remains from two nests there.[56] A total of 88% of the remains found around crowned eagle nests in the Ituri Rainforest in the Democratic Republic of the Congo were from primates: blue monkey (Cercopithecus mitis), red-tailed monkey, Wolf's mona monkey (C. wolfi), western red colobus and a mixture of mangabeys and black-and-white colobus.[57] In 16 nests in the Ivory Coast's Taï Forest, more than 60% of remains around the eyries were monkeys and more than 45% of remains were from Cercopithecus monkeys. About half of the Cercopithecus remains were from Diana monkey (C. diana; 44 items from 28 adults and 16 immatures), with the other half from Campbell's mona monkey (C. campbelli) and lesser spot-nosed monkey (C. petaurista), which were indistinguishable as remains. Other represented monkeys in the Taï Forest included the Western red colobus, the olive colobus (Procolobus verus) and the king colobus (Colobus polykomos).[8] The diet is, by necessity, more diverse in Kiwengwa/Pongwe Forest Reserve in the Matumbi Hills of Tanzania but the blue monkey
was still the most represented prey species, making up 20% of the
remains. In one nest in the prior study, the Blue monkey made up more
than 90% of the remains.[58] Where they overlap in range, the vervet monkey
can be quite prominent as prey for crowned eagles, as their relatively
small size, diurnal and terrestrial habits may make them more
vulnerable.[59][60]
Struhsaker and Leakey found that among both small and large
species, adult male monkeys were often better represented in the diet
than adult females.[56]
This may be a result of the behavior of male primates. In some species,
only adult males often travel separately from social groups, only adult
males take aggressive action against eagles, and only adult males have
loud calls that may attract the attention of eagles.[56][61] The variation in behavior of primates is illustrated so that in red colobus species, in which adult males patrol in mixed groups, the males are rarely attacked; meanwhile in the black-and-white colobus, in which adult males often patrol alone, males are regularly attacked and killed by crowned eagles.[4] However, adult monkeys (52%) were found in almost equal measure as juvenile monkeys (at 48%) in Uganda.[4]
As various species and ages of primates are taken, primate prey
varies in size. All diurnal African monkeys weigh in excess of 2 kg
(4.4 lb) in adulthood.[61] African primates weighing under 2 kg (4.4 lb), are almost entirely arboreal and nocturnal.[62] However, such primates, are actually allies of lemurs rather than monkeys, may occasionally also be hunted by crowned eagles. Potto (Perodicticus potto) are notable secondary prey in the Tai Forest and various galagos have been found across the range, such as in the Kiwengwa/Pongwe Forest Reserve where they made up 7.5% of the remains.[8][58]
Although no one has seen the eagles catch these primates, it is thought
pottos and galagos are likely to be taken if discovered by an eagle
while sleeping in dense foliage during the day.[4] Among monkeys, Cercopithecine monkeys are on the small side. Full-grown female Cercopithecus
can range in average weight from 2.7 to 4.26 kg (6.0 to 9.4 lb) and
males from 4.1 to 6.9 kg (9.0 to 15.2 lb) depending on species.[62][56][8] Other major prey, such as vervet monkeys typically weighing 5.5 kg (12 lb) in males and 4.1 kg (9.0 lb) in females, but up to 8 kg (18 lb).[63][64]Mangabeys and most colobus monkeys are larger, weighing considerably in excess of 5 kg (11 lb) at maturity.[62] The adults of these species taken by crowned eagles usually weigh 10.5 kg (23 lb)in males and 7 kg (15 lb) in females,[56] but olive colobus are smaller, weighing less than 5 kg (11 lb),[8] and large adults of western red colobus and Sooty mangabeys can weigh up to 11–12 kg (24–26 lb).[65] Larger king colobus, mantled guereza and black-and-white colobus can weigh up to 13.5 kg (30 lb) in adult males.[66][67] Based on observation and literature, any monkey of a weight of up to 10 to 15 kg (22 to 33 lb) may reportedly be hunted.[68][69]
The crowned eagle's diet may extend to the young and juveniles of baboons and similar species. yellow baboons, olive baboons (Papio anubis), chacma baboons (P. ursinus), drills (Mandrillus leucophaeus) and mandrills (M. sphinx), all falling into the above maximum primate weight bracket and all have been successfully hunted.[4][16][70][71] In South Africa, remains of three adults or subadult chacma baboons were found in the nest remain, suggesting adult female or subadult baboons can be possibly preyed upon.[72]
In many cases, baboons and drills may be actively avoided, as the adult
males of these species, at up to twice as heavy as the female, are
probably invulnerable to hunting and tend to be notoriously
violent-tempered. In one case, olive baboons destroyed the nest of a
crowned eagle pair after one of the eagles killed a baby in the baboon
group.[73]
When experimentally exposed to an image of a crowned eagle, a dominant
male mandrill reacted aggressively towards it, implying that they will
keep eagles at bay to protect the more vulnerable members of their
troop.[74] The crowned eagle is considered a potential predator of infant chimpanzees (Pan troglodytes) and bonobos (P. paniscus), although this is unconfirmed.[75] Although rarely observed, some monkeys will "bait" crowned eagles, which harass and provoke them. Leslie Brown described Sykes' monkeys (Cercopithecus albogularis) as rarely baiting eagles "with the insolent impunity of an expert tolero with a bull".[21]
Ungulate
A full-grown female bushbuck (Tragelaphus scriptus) is the largest animal known to be hunted by crowned eagles.
Outside of the rainforest, the crowned eagle's diet tends to be
somewhat more diverse. While they may take monkeys on a relatively small
scale, other families, largely the small antelopes and suids (specifically, the red river hog (Potamochoerus porcus)), are the leading prey.[21][6] In some wooded hillside nests in Kenya, about half of the remains were from small antelope. The predominant prey species there is the Suni (Neotragus moschatus).[76] In the Matumbi Hills of Tanzania, antelope make up about 30% of the dietary intake at nests, much of this being the Suni again.[58] Adults antelope of a similarly small size to the Suni, around 5 kg (11 lb) or slightly less, are readily hunted, such as Kirk's dik-dik (Madoqua kirkii) and the blue duiker (Philantomba monticola).[77][78]
Larger antelopes are also taken, mainly calves but occasionally adults
weighing around or more than 10 kg (22 lb) can be taken, including klipspringer (Oreotragus oreotragus), steenbok (Raphicerus campestris), Cape grysbok (R. melanotis), Sharpe's grysbok (R. sharpei) and about a half dozen small to medium sized duikers, especially the red forest duiker (Cephalophus natalensis) and larger common duiker (Sylvicapra grimmia).[58][72][79][80] National Geographic recorded a video depicting a crowned eagle stalking a water chevrotain (Hyemoschus aquaticus) (the only African representative of a small-bodied, deer-like family) along a rainforest river but show the chevrotain evading the eagle by submerging and swimming away from it.[81] Even bigger bushbuck and greater kudu, mainly calves, are sometimes preyed upon regularly.[6]
Among accurately weighed bushbuck to be killed by a crowned eagle, some
scaled 15.9 kg (35 lb), 20 kg (44 lb), or even 28.8 kg (63 lb) and
30 kg (66 lb) respectively in case of young ram, which about up to eight
times the eagle's weight.[21][49][82][83] An even more extreme body size estimated for a bushbuck kill was 50 kg (110 lb).[84] Crowned eagles are known to hunt fawns of other largish antelopes including Thomson's gazelle (Eudorcas thomsonii), grey rhebok (Pelea capreolus) and impala (Aepyceros melampus), and trained eagles from re-introduction programme managed to kill adult female impalas and bushbucks.[16][38]
The taking of ungulates on a large scale, unlike primates, is not
unique to the crowned eagle. The martial eagle was reported in Tsavo East National Park to hunt mainly dik-diks and elsewhere has exceptionally killed large duiker weighing up to 37 kg (82 lb).[7][85] Equally or even more impressive feats of ungulate-hunting have been credited to the wedge-tailed eagle (Aquila audax) and especially the golden eagle, despite these being considered less powerful than the crowned eagle.[16] Both Aquila take mainly neonatal lambs and deer fawns but can attack adult sheep and other large prey (i.e. full-grown kangaroos, emus, pronghorns, reindeer, etc.).[86][87] Even a bald eagle (Haliaeetus leucocephalus), usually target aquatic preys, was reportedly witnessed killing a pregnant adult sheep ewe.[88] The largest prey attacked by any living raptor is probably the up to 114 kg (251 lb) domestic calves attacked and killed by golden eagles.[89]
Hyraxes are known to be a major source of prey outsides of the rainforest.[72][80]
All six currently recognized species of hyrax have been hunted by
crowned eagles. The greatest level of predation for this family has been
directed at Southern tree hyrax (Dendrohyrax arboreus), but when they become locally abundant, rock hyrax (Procavia capensis) can become the species' preferred prey.[21] Other mammals have been recorded as opportunistic prey, including bats, hares (Lepus sp.), springhares (Pedetes sp.), cane rats (Thryonomys sp.), giant pouched rats (Cricetomys sp.), sun squirrels (Heliosciurus sp.), African giant squirrels (Protoxerus sp.), brush-tailed porcupines (Atherurus sp.), four-toed elephant shrews (Petrodromus tetradactylus) and tree pangolins (Manis tricuspis).[4][7][16][90][6] Despite their obvious defenses and nocturnal habits, small Cape porcupines (Hystrix africaeaustralis) have reportedly been taken in South Africa.[38]
These assorted mammals, generally smaller than primates and ungulates,
are typically taken when preferred prey species are locally scarce.[10] Mammalian carnivores are sometimes also hunted ranging from smaller types such as yellow mongoose (Cynictis penicillata), banded mongoose (Mungos mungo), cusimanses, African palm civet (Nandinia binotata) or genets to larger varieties such as adult African wildcats (Felis lybica) and black-backed jackals (Canis mesomelas).[16][56][72][58]
Adult eagles will only resort to hunting large birds when mammals are scarce, but in southern Africa, they can be a fairly common component of the diet. Bird prey can include guineafowl, francolins, ibis, pigeons, ostrich chicks (Struthio camelus) and the fledglings of herons and storks.[7][38] One nest even had the remains of a marabou stork (Leptoptilos crumeniferus), which is a formidable species not usually susceptible to avian predators.[16]Hornbills may be the most widely represented bird in the diet and the black-casqued wattled hornbill (Ceratogymna atrata)
reacts strongly to both the call of crowned eagles and the alarm calls
of monkeys specified to the eagles (which are separately identifiable
from the monkey's leopard-related alarm calls to both humans and,
apparently, hornbills).[4][91] In Kenya, snakes, including venomous varieties, may regularly supplement the diet.[7]Monitor lizards may also be hunted and, as in the martial eagle, the crowned eagle may attack even the largest African monitors, the adult Nile monitor (Varanus niloticus) and the rock monitor (V. albigularis).[92][72][58]
Domestic animals, including chicken (Gallus gallus domesticus), turkeys (Meleagris gallopavo), cats (Felis catus), small to medium-sized dogs (Canis familiaris), small pigs (Sus domesticus), lambs (Ovis aries), and goats (Capra hircus), are taken only when wild prey is greatly depleted.[7][38] Juveniles and subadults, may take unconventional prey more frequently than adults.
In
the rainforest interior, the crowned eagle occupies a unique niche and
it is, by far, the largest and most dominant raptorial bird in such
areas. Other large predators that may exploit similar prey in the same
forested habitats include leopard (Panthera pardus), African golden cat (Profelis aurata), Nile crocodile (Crocodylus niloticus), dwarf crocodile (Osteolaemus tetraspis), African rock python (Python sebae), chimpanzee and larger monkeys, like mandrill and baboons.[16][21][51]
All of these competitors are much heavier bodied than a crowned eagle,
ranging in size from the 10 kg (22 lb) golden cat to the 225 kg (496 lb)
Nile crocodile.[93][94] While the reptiles usually hunt on the ground or near water, felids, large monitor lizards and baboons can be assured thieves who will climb trees and take crowned eagle kills.[21]
In a comparison of the monkey-based diets of rainforest crowned eagles
with leopards and chimpanzees, the big cat was estimated to take prey
averaging 11.27 kg (24.8 lb), about twice the average estimated prey
weight for crowned eagles in the same ecosystem, and the great ape
6.9 kg (15 lb), about a kilogram more than that of the crowned eagle.[47] In South Africa, it is reported that Cape porcupines and bushpigs (Potamochoerus larvatus)
are attracted to trees used for prey consumption by crowned eagles, in
order to scavenge the sinew and bone that's discarded to the ground.[38]
In more mixed eastern and southern habitats, the diversity of large
predators is higher and the crowned eagle, despite its great power, is
not assured at the top of the avian food chain. Amongst the more
formidable raptors, the martial, the Verreaux's, and the crowned eagles,
may live on the same hillside and all hunt hyraxes. While the Verreaux's eagle is something of a rock hyrax
specialist, the martial eagle, even more so than the crowned eagle, has
an extremely broad prey base. All these eagles will readily steal the
prey caught by the other raptors.[95]
However, the great eagles are segregated by both their habitat
preferences and main hunting techniques, which make it possible for the
species to successfully nest within a few kilometers of one another.
While the crowned eagle lives in denser woods and hunts from a perch,
the martial eagle tends to live in more open wooded savanna habitats and
tends to hunt on the wing at a high flying height (thanks to its superb
vision) and the Verreaux's eagle lives in precipitous mountain habitats
and tends to contour-hunt, hugging the uneven contours of the rocks
while flying, only a few meters high. Like the martial eagle, the
crowned eagle has been known to prey on smaller raptorial birds.[7]
Young and inexperienced crowned eagles may be killed as prey by
large carnivores. Two eagles reintroduced into the wild were killed by
predators, one by a leopard that surprised a male on a monkey kill in
the rain, and the other by a crocodile that took a female as she ate a
young bushbuck kill near the water's edge.[16] In Kenya, cases of predation on nestlings and fledglings have reportedly involved honey badgers (Mellivora capensis) and cobras.[16] In one extraordinary case, an adult male Sanje mangabey (Cercocebus sanjei),
estimated to weigh 10 to 12 kg (22 to 26 lb), attacked an adult female
crowned eagle that was trying to hunt his troop, jumping onto her back
when she was in mid-flight and killing her with a strong bite.[96]
This is the first confirmed instance of a monkey killing an adult
crowned eagle. In one case, a female who tried to hunt an adult female
baboon was found seriously wounded after a male baboon interceded,
though the eagle was captured, medically treated, and lived in
captivity.[16] In another case involving potential prey turning the tables, an eagle that was trying to hunt an incubating female Egyptian goose (Alopochen aegyptiacus) was quickly attacked by her mate, who flew at the eagle and bit repeatedly at it, causing the eagle to quickly withdraw.[16] The average life expectancy for crowned eagles is 14 to 16 years, although some specimens live more than 30 years.[10][16][97]
While
several smaller raptorial birds will attack humans if they come too
close to the nesting site, usually these have minor consequences for the
human victim. Mother crowned eagles, in the post-fledging stage,
readily attack any human who comes close to the nest. Adult males may
also attack humans before fledging but only do so rarely.[21] Despite the size and power of the eagle,
attacks by the eagles may have minor consequences as well since the
attacks are meant only to displace and not kill or seriously maim the
intruding animal. However, nest-defense attacks may nonetheless possibly
result in deep, painful, open wounds, which can lead to risk of
infection or the need for stitches.
The crowned eagle is perhaps the only extant raptorial bird which
has been believed to attack human children as prey, outside of few
exceptional circumstances in the case of the Wedge-tailed eagle and the Martial eagle. [87][98][99]
In one case, a 7-year-old boy, of a weight of approximately 20 kg
(44 lb), was ambushed by a crowned eagle, who gouged its talons through
the boy's throat and chest. The attack was ended by a woman who came
upon them and rescued the child by bludgeoning the eagle to death with a
hoe.[41] In another case, the skull of a human child was found in the nest of a crowned eagle pair.[41]
In yet another instance, when assisting in the investigation of the
disappearance of a four-year-old girl, Simon Thomsett came to believe
she was the victim of a crowned eagle after the severed arm of a child
was found in a tall tree that was inaccessible to leopards and known to
be used as a crowned eagle cache.[16]
One other living eagle, the martial eagle (Polemaetus bellicosus), has been confirmed to take a human child in a possible predation attempt, a four-year-old boy in Ethiopia.
Unlike the crowned eagle, the martial eagle is not a specialized
primate hunter, and preys mainly on large ground-dwelling birds. The
eagle in question attacked three children, killing one of them, before
being shot by a villager.[100] In the famed Taung deposit in South Africa, a skull from a child Australopithecus africanus, a possible ancestor of human beings, led to considerable speculation. Ultimately referred to as the Taung Child
and estimated to weigh 9 to 11 kg (20 to 24 lb), the child became the
type specimen for its species. The child appeared to have died from a
clean row of piercing to its skull. Scholarly examination of the
piercings has led scientists to believe that the specimen was seemingly
killed by an eagle, of which the crowned eagle is the most likely
candidate.[101][65] This predatory relationship has led to much hypothesizing as to whether Stephanoaetus
eagles may have partially shaped human evolution, with small early
primate ancestors having evolved towards larger body sizes and larger
brains due to the reduced probability of eagle predation with these
features.[8][16][101]
The crowned eagle is fairly common in suitable habitat, though at the
population level, its numbers have shown a decline in sync with deforestation. Declines appear to be widespread and may be increasing due to the often fevered pace of clear-cutting.[7]
This species main habitat is rich, high-canopy forest, which is a major
target of timber companies, agriculturists, palm oil and biofuel
plantations and miners as well as slash and burn farmers. A charcoal-based economy outdoes mineral-based economics in Democratic Republic of the Congo and Zaire, both fuel wars and makes inroads deep into previously virgin forests.[16] As two of central Africa's largest businesses, this has a devastating effect on forests and wildlife. Charcoal taken from Kenya and Ethiopia is often thought to finance Somali warlords.[16]Tanzania,
which is more heavily developed for agriculture (largely today for
biofuel) than adjacent Kenya, has even more reduced forest habitat.[16]
The crowned eagle is far more common in protected areas and reserves
than elsewhere in its range, though is still recorded consistently
outside of these areas. Biologists in Africa now suspect that the
crowned eagles adaptability to small, fragmented tracts of woodland has
been exaggerated in the past.[16]
Some habitat losses have been offset by the establishment of exotic
tree plantations, where this species can nest, but which generally lack a
sufficient prey base.[46] The crowned eagle in Ethiopia
is certainly at very low density and restricted to protected areas. It
may be obliged to utilize exotics stands, but it is unlikely to ever be
capable of surviving in the complete absence of indigenous (and thus
prey productive) forests.[16] Certain southern African countries, such as Zambia, Zimbabwe and Malawi, have almost no extensive stands of native forest today, while other countries such as Namibia were never heavily forested.[16] In 2012 the species status was changed to Near Threatened by the IUCN.[1][102] Like the martial eagle,
the crowned has throughout modern history been persecuted by farmers,
who maintain that the bird is a threat to their livestock.[7] In fact, both the crowned and martial eagles only rarely attack livestock.[6] In some cases, however, crowned eagles have actually been killed while attempting to hunt domestic animals.[16]
Another cause for persecution of the species is that crowned eagles are
considered competitors in the illegal bushmeat and poaching trades.
Within the forest land-locked countries of Africa, the bushmeat trade is the largest source of animal protein for humans.[103]
It is a multibillion-dollar business with some 5 million tons (mostly
small antelopes and monkeys, the crowned eagle's staple diet) being
killed each year.[104] In just 500 million acres of the Congo Basin owned by 8 countries the weight equivalent to 40.7 million humans is removed each year (or 740,000 bull elephants).[105]
The effect of this unsustainable culling is to severely depress or
remove the large, medium and small wildlife species of the forests.
Crowned eagles require some 430 kg (950 lb) of "bush meat" a year and
thus directly compete with the industry.[16][70][103] The red colobus,
a monkey that typifies the optimal forest quality and is a main food
species for crowned eagles, has been singled out as one of the fastest
declining and most endangered monkeys in the world due primarily to the
bushmeat trade.[106]
In some cases, crowned eagles have reportedly even been shot by primate
conservationists in a misguided attempt to mitigate their predation of
declining primate species.[16]
It is estimated that 90% of the global distribution of the crowned
eagle may be subject to habitual persecution or is even killed and eaten
itself as bushmeat.[16][103]
On the other hand, some educated foresters and fruit-growers actually
encourage protection of populations, due to the controlling effect
crowned eagles have on populations of potentially harmful mammals.[7][38] In April 1996 the world's first captive-born crowned eagle hatched at the San Diego Zoo. Among ISIS registered zoos, only San Diego Zoo, San Francisco Zoo, Los Angeles Zoo, Fort Worth Zoo and Lowry Park Zoo house this species.[107] Several wildlife rehabilitation centers in Africa house crowned eagles.[16]
Due to their high-strung dispositions, tendency for aggression towards
humans and resistance to hunting prey via coercion and hunger, the
crowned eagle is often considered to be poorly suited for falconry.
However, there are several eagles of this species used as such in
England and sometimes in Africa, where they have been reportedly used to
cull locally overpopulated feral dogs.[16]Crowned eagle range
Subadult in Belize Zoo
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_immature.jpg)
_with_prey_..._(31850656390).jpg)
_female.jpg)
