Peregrine Falcon

Peregrine Falcon
Adult Peregrine Falcon (Falco peregrinus) of subspecies pealei or tundrius in Alaska
Conservation status
Least Concern (IUCN 3.1)[1]
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Aves
Order:	Falconiformes
Family:	Falconidae
Genus:	Falco
Species:	F. peregrinus
Binomial name
Falco peregrinus Tunstall, 1771
Subspecies
17–19, see text
Global range Yellow: Breeding summer visitor Green: Breeding resident Blue: Winter visitor Light blue: Passage visitor
Synonyms
Falco atriceps Hume Falco kreyenborgi Kleinschmidt, 1929 Falco pelegrinoides madens Ripley & Watson, 1963 Rhynchodon peregrinus (Tunstall, 1771) and see text

The Peregrine Falcon (Falco peregrinus), also known as the Peregrine,^[2] and historically as the Duck Hawk in North America,^[3] is a widespread bird of prey in the family Falconidae. A large, crow-sized falcon, it has a blue-gray back, barred white underparts, and a black head and "moustache". Typical of bird-eating raptors, Peregrine Falcons are sexually dimorphic, with females being considerably larger than males.^[4][5] The Peregrine is renowned for its speed, reaching over 325 km/h (202 mph) during its characteristic hunting stoop,^[6] making it the fastest member of the animal kingdom.^[7][8]

The Peregrine's breeding range includes land regions from the Arctic tundra to the tropics. It can be found nearly everywhere on Earth, except extreme polar regions, very high mountains, and most tropical rainforests; the only major ice-free landmass from which it is entirely absent is New Zealand. This makes it the world's most widespread bird of prey.^[9] Both the English and scientific names of this species mean "wandering falcon", referring to the migratory habits of many northern populations. Experts recognize 17 to 19 subspecies which vary in appearance and range; there is disagreement over whether the distinctive Barbary Falcon is represented by two subspecies of Falco peregrinus, or is a separate species, F. pelegrinoides.

While its diet consists almost exclusively of medium-sized birds, the Peregrine will occasionally hunt small mammals, small reptiles, or even insects. Reaching sexual maturity at one year, it mates for life and nests in a scrape, normally on cliff edges or, in recent times, on tall human-made structures.^[10] The Peregrine Falcon became an endangered species in many areas because of pesticides, especially DDT. Since the ban on DDT from the early 1970s, populations have recovered, supported by large-scale protection of nesting places and releases to the wild.^[11]

Description

The Peregrine Falcon has a body length of 34 to 58 centimetres (13–23 in) and a wingspan of around 80 to 120 centimetres (31–47 in).^[4][12] The male and female have similar markings and plumage, but as in many birds of prey the Peregrine Falcon displays marked reverse sexual dimorphism in size, with the female measuring up to 30 percent larger than the male.^[13] Males weigh 440 to 750 grams (0.97–1.7 lb) and the noticeably larger females weigh 910 to 1,500 grams (2.0–3.3 lb); for variation in weight between subspecies, see under that section below.

The back and the long pointed wings of the adult are usually bluish black to slate gray with indistinct darker barring (see "Subspecies" below); the wingtips are black.^[12] The white to rusty underparts are barred with thin clean bands of dark brown or black.^[14] The tail, colored like the back but with thin clean bars, is long, narrow and rounded at the end with a black tip and a white band at the very end. The top of the head and a "mustache" along the cheeks are black, contrasting sharply with the pale sides of the neck and white throat.^[15] The cere is yellow, as are the feet, and the beak and claws are black.^[16] The upper beak is notched near the tip, an adaptation which enables falcons to kill prey by severing the spinal column at the neck.^[4][5][6] The immature bird is much browner with streaked, rather than barred, underparts, and has a pale bluish cere and orbital ring.^[4]

Taxonomy and systematics

Peregrine Falcons, illustration by John James Audubon

Falco peregrinus was first described under its current binomial name by English ornithologist Marmaduke Tunstall in his 1771 Ornithologia Britannica.^[17] The scientific name Falco peregrinus is a Medieval Latin phrase that was used by Albertus Magnus in 1225. The specific name taken from the fact that juvenile birds were taken while journeying to their breeding location rather than from the nest, as falcon nests were difficult to get at.^[18] The Latin term for falcon, falco, is related to falx, the Latin word meaning sickle, in reference to the silhouette of the falcon's long, pointed wings in flight.^[6]

The Peregrine Falcon belongs to a genus whose lineage includes the hierofalcons^[19] and the Prairie Falcon (F. mexicanus). This lineage probably diverged from other falcons towards the end of the Late Miocene or in the Early Pliocene, about 8–5 million years ago (mya). As the Peregrine-hierofalcon group includes both Old World and North American species, it is likely that the lineage originated in western Eurasia or Africa. Its relationship to other falcons is not clear; the issue is complicated by widespread hybridization confounding mtDNA sequence analyses; for example a genetic lineage of the Saker Falcon (F. cherrug) is known^[20] which originated from a male Saker producing fertile young with a female Peregrine ancestor.^{[citation needed]}

Today, Peregrines are regularly hybridized in captivity with other species such as the Lanner Falcon (F. biarmicus) to produce the "perilanner", a somewhat popular bird in falconry as it combines the Peregrine's hunting skill with the Lanner's hardiness, or the Gyrfalcon to produce large, strikingly colored birds for the use of falconers. As can be seen, the Peregrine is still genetically close to the hierofalcons, though their lineages diverged in the Late Pliocene (maybe some 2.5–2 mya in the Gelasian).^[21]

Subspecies

Numerous subspecies of Falco peregrinus have been described, with 19 accepted by the 1994 Handbook of the Birds of the World,^[4][5][22] which considers the Barbary Falcon of the Canary Islands and coastal north Africa to be two subspecies (pelegrinoides and babylonicus) of Falco peregrinus, rather than a distinct species, F. pelegrinoides. The following map shows the general geographical ranges of these 19 subspecies:

Breeding ranges of the subspecies

Peregrine Falcon (Falco peregrinus anatum) in flight, Morro Bay, California

• Falco peregrinus anatum, described by Bonaparte in 1838,^[23] is known as the American Peregrine Falcon, or "Duck Hawk"; its scientific name means "Duck Peregrine Falcon". At one time, it was partly included in leucogenys. It is mainly found in the Rocky Mountains today. It was formerly common throughout North America between the tundra and northern Mexico, where current reintroduction efforts seek to restore the population.^[23] Most mature anatum, except those that breed in more northern areas, winter in their breeding range. Most vagrants that reach western Europe seem to belong to the more northern and strongly migratory tundrius, only considered distinct since 1968. It is similar to peregrinus but is slightly smaller; adults are somewhat paler and less patterned below, but juveniles are darker and more patterned below. Males weigh 500 to 700 grams (1.1–1.5 lb), while females weigh 800 to 1,100 grams (1.8–2.4 lb).^[24]

Painting of Falco peregrinus babylonicus by John Gould

• Falco peregrinus babylonicus, described by P.L. Sclater in 1861, is found in eastern Iran along the Hindu Kush and Tian Shan to Mongolian Altai ranges. A few birds winter in northern and northwestern India, mainly in dry semi-desert habitats.^[25] It is paler than pelegrinoides, and somewhat similar to a small, pale Lanner Falcon (Falco biarmicus). Males weigh 330 to 400 grams (12 to 14 oz), while females weigh 513 to 765 grams (18.1 to 27.0 oz).^[5]

• Falco peregrinus brookei, described by Sharpe in 1873, is also known as the Mediterranean Peregrine Falcon or the Maltese Falcon.^[26] It includes caucasicus and most specimens of the proposed race punicus, though others may be pelegrinoides, Barbary Falcons (see also below), or perhaps the rare hybrids between these two which might occur around Algeria. They occur from the Iberian Peninsula around the Mediterranean, except in arid regions, to the Caucasus. They are non-migratory. It is smaller than the nominate subspecies, and the underside usually has rusty hue.^[14] Males weigh around 445 grams (0.98 lb), while females weigh up to 920 grams (2.0 lb).^[5]

• Falco peregrinus calidus, described by John Latham in 1790, was formerly called leucogenys and includes caeruleiceps. It breeds in the Arctic tundra of Eurasia, from Murmansk Oblast to roughly Yana and Indigirka Rivers, Siberia. It is completely migratory, and travels south in winter as far as South Asia and sub-Saharan Africa. It is often seen around wetland habitats.^[27] It is paler than peregrinus, especially on the crown. Males weigh 588 to 740 grams (1.30–1.6 lb), while females weigh 925 to 1,333 grams (2.04–2.94 lb).^[5]

• Falco peregrinus cassini, described by Sharpe in 1873, is also known as the Austral Peregrine Falcon. It includes kreyenborgi, the Pallid Falcon^[28] a leucistic morph occurring in southernmost South America, which was long believed to be a distinct species.^[29] Its range includes South America from Ecuador through Bolivia, northern Argentina and Chile to Tierra del Fuego and Falkland Islands.^[14] It is non-migratory. It is similar to nominate, but slightly smaller with a black ear region. The variation kreyenborgi is medium grey above, has little barring below, and has a head pattern like the Saker Falcon, but the ear region is white.^[29]

• Falco peregrinus ernesti, described by Sharpe in 1894, is found from Indonesia to Philippines and south to Papua New Guinea and the nearby Bismarck Archipelago. Its geographical separation from nesiotes requires confirmation. It is non-migratory. It differs from the nominate subspecies in the very dark, dense barring on its underside and its black ear coverts.

• Falco peregrinus furuitii, described by Momiyama in 1927, is found on the Izu and Ogasawara Islands south of Honshū, Japan. It is non-migratory. It is very rare, and may only remain on a single island.^[4] It is a dark form, resembling pealei in color, but darker, especially on tail.^[14]

• Falco peregrinus japonensis, described by Gmelin in 1788, includes kleinschmidti and pleskei, and harterti, and seems to refer to intergrades with calidus. It is found from northeast Siberia to Kamchatka (though it is possibly replaced by pealei on the coast there) and Japan. Northern populations are migratory, while those of Japan are resident. It is similar to peregrinus, but the young are even darker than those of anatum.

Falco peregrinus macropus, Australia

• Falco peregrinus macropus, described by Swainson in 1837, is the Australian Peregrine Falcon. It is found in Australia in all regions except the southwest. It is non-migratory. It is similar to brookei in appearance, but is slightly smaller and the ear region is entirely black. The feet are proportionally large.^[14]

• Falco peregrinus madens, described by Ripley and Watson in 1963, is unusual in having some sexual dichromatism. If the Barbary Falcon (see below) is considered a distinct species, it is sometimes placed therein. It is found in the Cape Verde Islands, and is non-migratory;^[14] it is endangered with only six to eight pairs surviving.^[4] Males have a rufous wash on crown, nape, ears and back; underside conspicuously washed pinkish-brown. Females are tinged rich brown overall, especially on the crown and nape.^[14]

Falco peregrinus minor, illustration by Keulemans, 1874

• Falco peregrinus minor, first described by Bonaparte in 1850. It was formerly often perconfusus.^[30] It is sparsely and patchily distributed throughout much of sub-Saharan Africa and widespread in Southern Africa. It apparently reaches north along the Atlantic coast as far as Morocco. It is non-migratory, and small and dark.

• Falco peregrinus nesiotes, described by Mayr in 1941,^[31] is found in Fiji and probably also Vanuatu and New Caledonia. It is non-migratory.^[32]

Captive Falco peregrinus pealei

• Falco peregrinus pealei, described by Ridgway in 1873, is also known as Peale's Falcon, and includes rudolfi.^[33] It is found in the Pacific Northwest of North America, northwards from the Puget Sound along the British Columbia coast (including the Queen Charlotte Islands), along the Gulf of Alaska and the Aleutian Islands to the far eastern Bering Sea coast of Russia,^[33] and may also occur on the Kuril Islands and the coasts of Kamchatka. It is non-migratory. It is the largest subspecies, and it looks like an oversized and darker tundrius or like a strongly barred and large anatum. The bill is very wide.^[34] Juveniles occasionally have pale crowns. Males weigh 700 to 1,000 grams (1.5–2.2 lb), while females weigh 1,000 to 1,500 grams (2.2–3.3 lb).^[24]

• Falco peregrinus pelegrinoides, first described by Temminck in 1829, is found in the Canary Islands through north Africa and the Near East to Mesopotamia. It is most similar to brookei, but is markedly paler above, with a rusty neck, and is a light buff with reduced barring below. It is smaller than the nominate subspecies; females weigh around 610 grams (1.3 lb).^[5]

• Falco peregrinus peregrinator, described by Sundevall in 1837, is known as the Indian Peregrine Falcon, Black Shaheen, Indian Shaheen ^[35] or Shaheen Falcon.^[36] It was formerly sometimes known as Falco atriceps or Falco shaheen. Its range includes South Asia from Pakistan across India and Bangladesh to Sri Lanka and Southeastern China. In India, the Shaheen is reported from all states except Uttar Pradesh, mainly from rocky and hilly regions. The Shaheen is also reported from the Andaman and Nicobar Islands in the Bay of Bengal.^[25] They have a clutch size of 3 to 4 eggs, with the chicks fledging time of 48 days with an average nesting success of 1.32 chicks per nest. In India, apart from nesting on cliffs, it has also been recorded as nesting on man-made structures such as buildings and cellphone transmission towers.^[25] A population estimate of 40 breeding pairs in Sri Lanka was made in 1996.^[37] It is non-migratory, and is small and dark, with rufous underparts. In Sri Lanka this species is found to favour the higher hills while the migrant calidus is more often seen along the coast.^[38]

• Falco peregrinus peregrinus, the nominate (first-named) subspecies, described by Tunstall in 1771, breeds over much of temperate Eurasia between the tundra in the north and the Pyrenees, Mediterranean region and Alpide belt in the south.^[23] It is mainly non-migratory in Europe, but migratory in Scandinavia and Asia. Males weigh 580 to 750 grams (1.3–1.7 lb), while females weigh 925 to 1,300 grams (2.04–2.9 lb).^[5] It includes brevirostris, germanicus, rhenanus, and riphaeus.

• Falco peregrinus radama, described by Hartlaub in 1861, is found in Madagascar and Comoros. It is non-migratory.^[14]

• Falco peregrinus submelanogenys, described by Mathews in 1912, is the Southwest Australian Peregrine Falcon. It is found in southwest Australia and is non-migratory.

• Falco peregrinus tundrius, described by C.M. White in 1968, was at one time included in leucogenys It is found in the Arctic tundra of North America to Greenland, and migrates to wintering grounds in Central and South America.^[34] Most vagrants that reach western Europe belong to this subspecies, which was previously united with anatum. It is the New World equivalent to calidus. It is smaller than anatum. It is also paler than anatum; most have a conspicuous white forehead and white in ear region, but the crown and "moustache" are very dark, unlike in calidus.^[34] Juveniles are browner, and less grey, than in calidus, and paler, sometimes almost sandy, than in anatum. Males weigh 500 to 700 grams (1.1–1.5 lb), while females weigh 800 to 1,100 grams (1.8–2.4 lb).^[24]

Barbary Falcon

Main article: Barbary Falcon

Two of the subspecies listed above (Falco peregrinus pelegrinoides and F. p. babylonicus) are often instead treated together as a distinct species, Falco pelegrinoides (Barbary Falcon),^[5] although they were included within F. peregrinus in the 1994 Handbook of the Birds of the World.^[4] These birds inhabit arid regions from the Canary Islands along the rim of the Sahara through the Middle East to Central Asia and Mongolia.

Barbary Falcons have a red neck patch but otherwise differ in appearance from the Peregrine proper merely according to Gloger's Rule, relating pigmentation to environmental humidity.^[39] The Barbary Falcon has a peculiar way of flying, beating only the outer part of its wings like fulmars sometimes do; this also occurs in the Peregrine, but less often and far less pronounced.^[5] The Barbary Falcon's shoulder and pelvis bones are stout by comparison with the Peregrine, and its feet are smaller.^[40] Barbary Falcons breed at different times of year than neighboring Peregrine Falcon subspecies,^{[5][22][41][42][43][44][45]} but there are no postzygotic reproduction barriers in place.^[46] There is a 0.6–0.7% genetic distance in the Peregine-Barbary Falcon ("peregrinoid") complex.^[41]

Another subspecies of Falco peregrinus, madens, has also sometimes been treated instead within a separately recognized F. pelegrinoides.^[14]

Ecology and behavior

Closeup of head showing nostril tubercle

Silhouettes of Peregrine Falcons in normal flight (left) and at the start of a stoop

The Peregrine Falcon lives mostly along mountain ranges, river valleys, coastlines, and increasingly in cities.^[14] In mild-winter regions, it is usually a permanent resident, and some individuals, especially adult males, will remain on the breeding territory. Only populations that breed in Arctic climes typically migrate great distances during the northern winter.^[47]

The Peregrine Falcon is often stated to be the fastest animal on the planet in its hunting dive, the stoop,^[7] which involves soaring to a great height and then diving steeply at speeds commonly said to be over 320 km/h (200 mph), and hitting one wing of its prey so as not to harm itself on impact.^[6] The air pressure from a 200 mph (320 km/h) dive could possibly damage a bird's lungs, but small bony tubercles on a falcon's nostrils guide the powerful airflow away from the nostrils, enabling the bird to breathe more easily while diving by reducing the change in air pressure.^[48] To protect their eyes, the falcons use their nictitating membranes (third eyelids) to spread tears and clear debris from their eyes while maintaining vision. A study testing the flight physics of an "ideal falcon" found a theoretical speed limit at 400 km/h (250 mph) for low altitude flight and 625 km/h (390 mph) for high altitude flight.^[49] In 2005, Ken Franklin recorded a falcon stooping at a top speed of 389 km/h (242 mph).^[50] A video of one of the dives can be seen in this link.

The life span of Peregrine Falcons in the wild is up to 15.5 years.^[5] Mortality in the first year is 59–70%, declining to 25–32% annually in adults.^[5] Apart from such anthropogenic threats as collision with human-made objects, the Peregrine may be killed by large eagles or large owls.^[51]

The Peregrine Falcon is host to a range of parasites and pathogens. It is a vector for Avipoxvirus, Newcastle disease virus, Falconid herpesvirus 1 (and possibly other Herpesviridae), and some mycoses and bacterial infections. Endoparasites include Plasmodium relictum (usually not causing malaria in the Peregrine Falcon), Strigeidae trematodes, Serratospiculum amaculata (nematode), and tapeworms. Known Peregrine Falcon ectoparasites are chewing lice,^[52] Ceratophyllus garei (a flea), and Hippoboscidae flies (Icosta nigra, Ornithoctona erythrocephala).^[53]

Feeding

Immature Peregrine Falcon using a USFWS ship as a perch on which to eat its prey

The Peregrine Falcon feeds almost exclusively on medium-sized birds such as pigeons and doves, waterfowl, songbirds, and waders.^[16] Worldwide, it is estimated that between 1,500 and 2,000 bird species (up to roughly a fifth of the world's bird species) are predated somewhere by these falcons. In North America, prey has varied in size from 3-g hummingbirds to a 3.1-kg Sandhill Crane (killed by a peregrine in a swoop).^[54] Prey also include a small raptor, the American Kestrel.^[55] Other than bats taken at night,^[56] the Peregrine rarely hunts small mammals, but will on occasion take rats, voles, hares, shrews, mice and squirrels. Coastal populations of the large subspecies pealei feed almost exclusively on seabirds.^[15] In the Brazilian mangrove swamp of Cubatão, a wintering falcon of the subspecies tundrius was observed while successfully hunting a juvenile Scarlet Ibis.^[57] Insects and reptiles make up a small proportion of the diet, which varies greatly depending on what prey is available.^[16] In urban areas, the main component of the Peregrine's diet is the Rock or Feral Pigeon, which comprise 80% or more of the dietary intake for peregrines in some cities. Other common city birds are also taken regularly, including Mourning Doves, Common Wood Pigeons, Common Swifts, Northern Flickers, Common Starlings, American Robins, Common Blackbirds, and corvids (such as magpies or Carrion, House and American Crows).^[56]

The Peregrine Falcon hunts at dawn and dusk, when prey are most active, but also nocturnally in cities, particularly during migration periods when hunting at night may become prevalent. Nocturnal migrants taken by Peregrines include species as diverse as Yellow-billed Cuckoo, Black-necked Grebe, Virginia Rail and Common Quail.^[56] The Peregrine requires open space in order to hunt, and therefore often hunts over open water, marshes, valleys, fields, and tundra, searching for prey either from a high perch or from the air.^[58] Once prey is spotted, it begins its stoop, folding back the tail and wings, with feet tucked.^[15] Prey is struck and captured in mid-air; the Peregrine Falcon strikes its prey with a clenched foot, stunning or killing it with the impact, then turns to catch it in mid-air.^[58] If its prey is too heavy to carry, a Peregrine will drop it to the ground and eat it there. Prey is plucked before consumption.^[48]

Reproduction

Peregrine at nest, France

The Peregrine Falcon is sexually mature at the end of the first year of age, but in healthy populations they breed after two to three years of age. A pair mates for life and returns to the same nesting spot annually. The courtship flight includes a mix of aerial acrobatics, precise spirals, and steep dives.^[12] The male passes prey it has caught to the female in mid-air. To make this possible, the female actually flies upside-down to receive the food from the male's talons.

During the breeding season, the Peregrine Falcon is territorial; nesting pairs are usually more than 1 km (0.62 mi) apart, and often much farther, even in areas with large numbers of pairs.^[59] The distance between nests ensures sufficient food supply for pairs and their chicks. Within a breeding territory, a pair may have several nesting ledges; the number used by a pair can vary from one or two to seven in a 16 year period.

The Peregrine Falcon nests in a scrape, normally on cliff edges. The female chooses a nest site, where she scrapes a shallow hollow in the loose soil, sand, gravel, or dead vegetation in which to lay eggs. No nest materials are added.^[12] Cliff nests are generally located under an overhang, on ledges with vegetation, and south-facing sites are favored.^[15] In some regions, as in parts of Australia and on the west coast of Northern North America, large tree hollows are used for nesting. Before the demise of most European peregrines, a large population of peregrines in central and western Europe used the disused nests of other large birds.^[16] In remote, undisturbed areas such as the Arctic, steep slopes and even low rocks and mounds may be used as nest sites. In many parts of its range, Peregrines now also nest regularly on tall buildings or bridges; these human-made structures used for breeding closely resemble the natural cliff ledges that the Peregrine prefers for its nesting locations.^[4][59]

The pair defends the chosen nest site against other Peregrines, and often against ravens, herons, and gulls, and if ground-nesting, also such mammals as foxes, wolverines, felids, bears and wolves.^[59] Both nests and (less frequently) adults are predated by larger-bodied raptorial birds like eagles, large owls, or Gyrfalcons. Peregrines defending their nests have managed to kill raptors as large as Golden Eagles and Bald Eagles (both of which they normally avoid as potential predators) that have come close to the nest.^[60]

Peregrine chick

The date of egg-laying varies according to locality, but is generally from February to March in the Northern Hemisphere, and from July to August in the Southern Hemisphere, although the Australian subspecies macropus may breed as late as November, and equatorial populations may nest anytime between June and December. If the eggs are lost early in the nesting season, the female usually lays another clutch, although this is extremely rare in the Arctic due to the short summer season. Generally three to four eggs, but sometimes as few as one or as many as five, are laid in the scrape.^[61] The eggs are white to buff with red or brown markings.^[61] They are incubated for 29 to 33 days, mainly by the female,^[15] with the male also helping with the incubation of the eggs during the day, but only the female incubating them at night. The average number of young found in nests is 2.5, and the average number that fledge is about 1.5, due to the occasional production of infertile eggs and various natural losses of nestlings.^[4][48][51]

After hatching, the chicks (called "eyases"^[62]) are covered with creamy-white down and have disproportionately large feet.^[59] The male (called the "tiercel") and the female (simply called the "falcon") both leave the nest to gather prey to feed the young.^[48] The hunting territory of the parents can extend a radius of 19 to 24 km (12–15 miles) from the nest site.^[63] Chicks fledge 42 to 46 days after hatching, and remain dependent on their parents for up to two months.^[64]

Relationship with humans

Falconry

Main article: Falconry

Tethered Falco peregrinus

The Peregrine Falcon has been used in falconry for more than 3,000 years, beginning with nomads in central Asia.^[59] Due to its ability to dive at high speeds, it is highly sought-after and generally used by experienced falconers.^[13] Peregrine Falcons are also occasionally used to scare away birds at airports to reduce the risk of bird-plane strikes, improving air-traffic safety,^[65] and were used to intercept homing pigeons during World War II.^[66]

Until 2004 nearly all Peregrines used for falconry in the U.S. were captive-bred from the progeny of falcons taken before the U. S. Endangered Species Act was enacted and from those few infusions of wild genes available from Canada and special circumstances. Peregrine Falcons were removed from the United States' endangered species list in 1999 due largely to the effort and knowledge of falconers in collaboration with the Peregrine Fund and state and federal agencies – through a technique called hacking.^{[citation needed]} Finally, after years of close work with the US Fish and Wildlife Service, a limited take of wild Peregrines was allowed in 2004, the first wild Peregrines taken specifically for falconry in over 30 years. Since Peregrine eggs and chicks are still often targeted by illegal collectors,^[67] it is common practice not to publicize unprotected nest locations.^[68]

Captive breeding

Peregrine Falcons have been successfully bred in captivity, both for falconry and for release back into the wild.^[69]

The first successful captive breeding of peregrine falcons in North America occurred in the early 1970s by The Peregrine Fund, professor and falconer Heinz Meng, and other private falconer/breeders such as Dave Jamieson and Les Boyd who bred the first peregrines by means of artificial insemination. In Great Britain, falconer Phillip Glasier of the Falconry Centre in Newent, Gloucestershire, was successful in obtaining young from more than 20 species of captive raptors. A cooperative effort began between various government agencies, non-government organizations, and falconers to supplement various wild raptor populations in peril. This effort was strongest in North America where significant private donations along with funding allocations through the Endangered Species Act of 1972 provided the means to continue the release of captive-bred Peregrines and various other raptors. By the mid-1980s, falconers had become self-sufficient as regards sources of birds to train and fly, in addition to the immensely important conservation benefits conferred by captive breeding.

Decline due to pesticides

The Peregrine Falcon became an endangered species because of the use of organochlorine pesticides, especially DDT, during the 1950s, 60s, and 70s.^[70] Pesticide biomagnification caused organochlorine to build up in the falcons' fat tissues, reducing the amount of calcium in their eggshells. With thinner shells, fewer falcon eggs survived to hatching.^[58][71] In several parts of the world, such as the eastern United States and Belgium, this species became extirpated (locally extinct) as a result.^[64]

Recovery efforts

Peregrine Falcon on the Ambassador Bridge in Detroit-Windsor.

In the United States, Canada, Germany and Poland, wildlife services in Peregrine Falcon recovery teams breed the species in captivity.^[72] The chicks are usually fed through a chute or with a hand puppet mimicking a Peregrine's head, so they cannot see to imprint on the human trainers.^[47] Then, when they are old enough, the rearing box is opened, allowing the bird to train its wings. As the fledgling gets stronger, feeding is reduced forcing the bird to learn to hunt. This procedure is called hacking back to the wild.^[73] To release a captive-bred falcon, the bird is placed in a special cage at the top of a tower or cliff ledge for some days or so, allowing it to acclimate itself to its future environment.^[73] Worldwide recovery efforts have been remarkably successful.^[72] The widespread restriction of DDT use eventually allowed released birds to breed successfully.^[47] The Peregrine Falcon was removed from the U.S. Endangered Species list on August 25, 1999.^[47][74]

Some controversy has existed over the origins of captive breeding stock used by The Peregrine Fund in the recovery of peregrine falcons throughout the contiguous United States. Several peregrine subspecies were included in the breeding stock, including birds of Eurasian origin. Due to the extirpation of the Eastern anatum (Falco peregrinus anatum), the near extirpation of the anatum in the Midwest, and the limited gene pool within North American breeding stock, the inclusion of non-native subspecies was justified to optimize the genetic diversity found within the species as a whole.^[75] Such strategies are common in endangered species re-introduction scenarios,^{[citation needed]} where dramatic population declines result in a genetic bottleneck and the loss of genetic diversity.

Current status

Populations of the Peregrine Falcon have bounced back in most parts of the world. In Britain, there has been a recovery of populations since the crash of the 1960s. This has been greatly assisted by conservation and protection work led by the Royal Society for the Protection of Birds. The RSPB has estimated that there are 1,402 breeding pairs in the UK.^[76][77] Peregrines now breed in many mountainous and coastal areas, especially in the west and north, and nest in some urban areas, capitalising on the urban Feral Pigeon populations for food.^[78] In many parts of the world Peregrine Falcons have adapted to urban habitats, nesting on cathedrals, skyscraper window ledges, tower blocks,^[79] and the towers of suspension bridges. Many of these nesting birds are encouraged, sometimes gathering media attention and often monitored by cameras.^[80][81]

Cultural significance

Idaho state quarter

Due to its striking hunting technique, the Peregrine has often been associated with aggression and martial prowess. Native Americans of the Mississippian culture (c. 800–1500) used the Peregrine, along with other several birds of prey, in imagery as a symbol of "aerial (celestial) power" and buried men of high status in costumes associating to the ferocity of "raptorial" birds.^[82] In the late Middle Ages, the Western European nobility that used Peregrines for hunting, considered the bird associated with princes in formal hierarchies of birds of prey, just below the Gyrfalcon associated with kings. It was considered "a royal bird, more armed by its courage than its claws". Terminology used by Peregrine breeders also used the Old French term gentil, "of noble birth; aristocratic", particularly with the Peregrine.^[83]

Since 1927, the Peregrine Falcon has been the official mascot of Bowling Green State University in Bowling Green, Ohio.^[84] The 2007 U.S. Idaho state quarter features a Peregrine Falcon.^[85]

Birds portal

References

1. ^ BirdLife International (2009). Falco peregrinus. In: IUCN 2008. IUCN Red List of Threatened Species. Downloaded on 2009-11-03.
2. ^ Heinzel, H.; Fitter, R.S.R.; Parslow, J. (1995), Birds of Britain and Europe (5 ed.), London: HarperCollins, ISBN 0-00-219894-0 
3. ^ Friedmann, H. (1950), "The birds of North and Middle America", U.S. National Museum Bulletin 50 (11): 1–793 
4. ^ ^{a b c d e f g h i j} White, C.M. et al. (1994), "Family Falconidae", in del Hoyo, J., Elliot, A. and Sargatal, J., Handbook of Birds of the World: New World Vultures to Guinea fowl, 2, Barcelona: Lynx Edicions, pp. 216–275, plates 24–28, ISBN 84-87334-15-6 
5. ^ ^{a b c d e f g h i j k l m} Snow, D.W. et al. (1998), The Complete Birds of the Western Palaearctic on CD-ROM, Oxford University Press, ISBN 0-19-268579-1 
6. ^ ^{a b c d} U.S. Fish and Wildlife Service (1999), All about the Peregrine falcon, archived from the original on 2008-04-16, http://web.archive.org/web/20080416195055/http://www.fws.gov/endangered/recovery/peregrine/QandA.html#fast, retrieved 2007-08-13 
7. ^ ^{a b} "Wildlife Finder – Peregrine Falcon". BBC. http://www.bbc.co.uk/nature/species/Peregrine_falcon. Retrieved 2010-03-18. 
8. ^ "The World's Fastest Animal Takes New York". Smithsonian. http://www.smithsonianmag.com/science-nature/The-Worlds-Fastest-Animal-Takes-New-York.html. Retrieved 2010-11-08. 
9. ^ Ferguson-Lees, J.; Christie, D.A. (2001), Raptors of the World, London: Christopher Helm, ISBN 0-7136-8026-1 
10. ^ Cade, T.J. et al. (1996), "Peregrine Falcons in Urban North America", in Bird, D.M., D.E. Varland & J.J. Negro, Raptors in Human Landscapes, London: Academic Press, pp. 3–13, ISBN 0-12-100130-X 
11. ^ Cade, T.J. et al. (1988), Peregrine Falcon Populations – Their management and recovery, The Peregrine Fund, Boise, Idaho, ISBN 0-9619839-0-6 
12. ^ ^{a b c d} Dewey, T. and Potter, M. (2002), Animal Diversity Web: Falco peregrinus, University of Michigan Museum of Zoology, http://animaldiversity.ummz.umich.edu/site/accounts/information/Falco_peregrinus.html, retrieved 21 May 2008 
13. ^ ^{a b} Scholz, F. (1993), Birds of Prey, Stackpole Books, ISBN 0-8117-0242-1 
14. ^ ^{a b c d e f g h i j} Ferguson-Lees, J. and Christie, D. (2001), Raptors of the World, Houghton Mifflin Field Guides, ISBN 0-618-12762-3 
15. ^ ^{a b c d e} Terres, J.K. (1991), The Audubon Society Encyclopedia of North American Birds, Wings Books, New York, ISBN 0-517-03288-0 
16. ^ ^{a b c d} Beckstead, D. (2001)
17. ^ (Latin) Tunstall, Marmaduke (1771), Ornithologia Britannica: seu Avium omnium Britannicarum tam terrrestrium, quam aquaticarum catalogus, sermone Latino, Anglico et Gallico redditus: cui subjuctur appendix avec alennigenas, in Angliam raro advenientes, complectens, London, J. Dixwell 
18. ^ Merriam-Webster, Inc (1991), The Merriam-Webster new book of word histories, Merriam-Webster, pp. 365, ISBN 0877796033, http://books.google.com/?id=IrcZEZ1bOJsC&pg=PA365&dq=peregrinus+meaning+falcon#v=onepage&q=peregrinus%20meaning%20falcon&f=false 
19. ^ Contra Helbig et al. (1994), Wink et al. (1998). The supposed basal position of the hierofalcons was due to them having a cytochrome b numt: see Wink & Sauer-Gürth (2000)
20. ^ Helbig et al. (1994), Wink et al. (1998)
21. ^ Helbig et al. (1994), Wink et al. (1998), Griffiths (1999), Wink & Sauer-Gürth (2000), Groombridge et al. (2002), Griffiths et al. (2004), Nittinger et al. (2005)
22. ^ ^{a b} Vaurie (1961)
23. ^ ^{a b c} American Ornithologists' Union (1910):p.164
24. ^ ^{a b c} White, Clayton M.; Clum, Nancy J.; Cade, Tom J.; Hunt, W. Grainger (2002). "Peregrine Falcon (Falco peregrinus)". The Birds of North America Online (A. Poole, Ed.). Ithaca: Cornell Lab of Ornithology. http://bna.birds.cornell.edu/bna/species/660. Retrieved 13 May 2011. 
25. ^ ^{a b c} Pande, Satish; Yosef, Reuven; Mahabal, Anil (2009), "Distribution of the Peregrine Falcon (Falco peregrinus babylonicus, F. p. calidus and F. p. peregrinator) in India with some notes on the nesting habits of the Shaheen Falcon", in Sielicki, Janusz, Peregrine Falcon populations – Status and Perspectives in the 21st Century, Mizera, Tadeusz, European Peregrine Falcon Working Group and Society for the Protection of Wild animals "Falcon", Poland and Turl Publishing & Poznan University of Life Sciences Press, Warsaw-Poznan, p. 800, ISBN 978-83-920969-6-2 
26. ^ Charles V, Holy Roman Emperor levied a rent of these birds on the Knights Hospitaller when he donated the Island of Malta to them. Source of the name for Dashiell Hammett's novel.
27. ^ Rasmussen, PC & JC Anderton (2005), Birds of South Asia. The Ripley Guide. Volume 2, Smithsonian Institution and Lynx Edicions, p. 116 
28. ^ Also called "Kleinschmidt's Falcon", but this might equally refer to F. p. kleinschmidti which is a junior synonym of japonensis,
29. ^ ^{a b} Ellis, David H.; Garat, Cesar P. (1983), "The Pallid Falcon Falco kreyenborgi is a color phase of the Austral Peregrine Falcon (Falco peregrinus cassini)", Auk 100 (2): 269–71, http://elibrary.unm.edu/sora/Auk/v100n02/p0269-p0271.pdf, retrieved 24 May 2008. 
30. ^ Vaurie, 1961
31. ^ Mayr (1941)
32. ^ Peters, J. L.; Mayr, E. & Cottrell, W. (1979):p.423
33. ^ ^{a b} American Ornithologists' Union (1910):p.165
34. ^ ^{a b c} Proctor, N. & Lynch, P. (1993):p.13
35. ^ The shaheen (شاهین) of Arabic and Persian writers are usually Barbary Falcons; those in Indian (शाहीन) and Pakistani (شاہین) sources normally refer to peregrinator.
36. ^ Gehan de Silva Wijeratne; Deepal Warakagoda & T.S.U. de Zylva (2007), "Species description", A Photographic Guide to Birds of Sri Lanka, New Holland Publishers (UK) Ltd, p. 37 and/or 144, ISBN 9781859745113 
37. ^ Döttlinger,Hermann; Hoffmann,Thilo W (1999), "Status of the Black Shaheen or Indian Peregrine Falcon Falco peregrinus peregrinator in Sri Lanka", J. Bombay Nat. Hist. Soc. 96 (2): 239–243. 
38. ^ Döttlinger & Nicholls (2005)
39. ^ Döttlinger, 2002
40. ^ (Vaurie, 1961)
41. ^ ^{a b} Wink et al. (2000)
42. ^ Helbig et al. (1994)
43. ^ Wink et al. (1998)
44. ^ Wink & Sauer-Gürth (2000)
45. ^ Wink et al. (2004)
46. ^ Blondel (1999)
47. ^ ^{a b c d} U.S. Fish and Wildlife Service (1995), Peregrine Falcon, http://ecos.fws.gov/speciesProfile/SpeciesReport.do?spcode=B050, retrieved 2008-05-22 
48. ^ ^{a b c d} Wisconsin Department of Natural Resources
49. ^ Tucker (1998)
50. ^ Harpole, Tom (2005-03-01). "Falling with the Falcon". Smithsonian Air & Space magazine. http://www.airspacemag.com/flight-today/falcon.html. Retrieved 2008-09-04. 
51. ^ ^{a b} Michigan Department of Natural Resources (2007)
52. ^ Colpocephalum falconii which was described from specimens found on the Peregrine Falcon, Colpocephalum subzerafae, Colpocephalum zerafae and Nosopon lucidum (all Menoponidae), Degeeriella rufa (Philopteridae), Laemobothrion tinnunculi (Laemobothriidae). All are known from other Falco species too.(Dewey & Potter 2002, Dagleish 2003)
53. ^ Raidal et al. (1999), Raidal & Jaensch (2000), Dewey & Potter (2002), Dalgleish (2003)
54. ^ "Birds of North America Online". Bna.birds.cornell.edu. http://bna.birds.cornell.edu/bna/species/660/articles/foodhabits. Retrieved 2011-08-30. 
55. ^ C.Michael Hogan, ed. 2010. American Kestrel. Encyclopedia of Earth, U.S. National Council for Science and the Environment, Ed-in-chief C.Cleveland
56. ^ ^{a b c} Drewitt, E.J.A.; Dixon, N. (February 2008), "Diet and prey selection of urban-dwelling Peregrine Falcons in southwest England", British Birds 101: 58–67 
57. ^ Olmos, F.; Silva e Silva, R. (2003), Guará: Ambiente , Fauna e Flora dos Manguezais de Santos-Cubatão, Brasil, São Paulo: Empresa das Artes, p. 111, ISBN 85-89138-06-2 
58. ^ ^{a b c} Ehrlich, P.; Dobkin, D.; Wheye, D. (1992), Birds in Jeopardy: The Imperiled and Extinct Birds of the United States, Stanford University Press, ISBN 0-8047-1981-0 
59. ^ ^{a b c d e} Blood, D. and Banasch, U. (2001), Hinterland Who's Who Bird Fact Sheets: Peregrine Falcon, http://www.hww.ca/hww2.asp?id=60, retrieved 2008-05-22 
60. ^ "— Birds of North America Online". Bna.birds.cornell.edu. http://bna.birds.cornell.edu/bna/species/660/articles/behavior. Retrieved 2011-08-30. 
61. ^ ^{a b} Peterson, R. T (1976):p.171
62. ^ "Falcon Facts". Raptorresource.org. http://www.raptorresource.org/facts.htm. Retrieved 2011-08-31. 
63. ^ Towry (1987)
64. ^ ^{a b} Snow (1994)
65. ^ Kuzir, S. and Muzini, J. (1999), "Birds and air traffic safety on Zagreb airport (Croatia)", The Environmentalist 18 (4): 231–237, doi:10.1023/A:1006541304592 
66. ^ Enderson, James; Katona, Robert (illustrator) (2005), Peregrine Falcon: Stories of the Blue Meanie, University of Texas Press, p. 175, ISBN 0-292-70624-3 
67. ^ "Falco peregrinus". Scientific-web.com. http://www.scientific-web.com/en/Biology/Animalia/Chordata/Aves/FalcoPeregrinus01.html. Retrieved 2011-08-31. 
68. ^ American Birding Association (2005), Code of Birding Ethics, American Birding Association, http://www.aba.org/about/ethics.html, retrieved 2008-05-26 
69. ^ "SCPBRG: Captive Breeding Program". .ucsc.edu. http://www2.ucsc.edu/scpbrg/captive_breeding.htm. Retrieved 2011-08-30. 
70. ^ T. J. Cade, J. H. Enderson, C. G. Thelander & C. M. White (Eds): Peregrine Falcon Populations – Their management and recovery. The Peregrine Fund, Boise, Idaho, 1988. ISBN 0-9619839-0-6.
71. ^ Brown (1976)
72. ^ ^{a b} Cassidy, J. and Reader's Digest Editors (2005). "Book of North American Birds". Reader's Digest (via Google Books). p. 34. ISBN 0-89577-351-1. http://books.google.com/?id=eKU-5_7lfXMC&dq. Retrieved 2008-05-26. 
73. ^ ^{a b} Aitken, G. (2004), A New Approach to Conservation, Ashgate Publishing, p. 126, ISBN 0-7546-3283-0, http://books.google.com/?id=IAu-MzZbAPMC&dq, retrieved 2008-05-26 
74. ^ Henny, Charles; Nelson, Morlan W. (1981), "Decline and Present Status of Breeding Peregrine Falcons in Oregon", The Murrelet (Society for Northwestern Vertebrate Biology) 62 (2): 43–53, doi:10.2307/3534174, JSTOR 3534174, "The records of Richard M. Bond and William E. Griffee, and the recollections of Larry L. Schramm and Merlin A. McColm were critical in putting the Peregrine back off the endangered list." 
75. ^ Cade, T.J., Burnham W. 2003 The Return of the Peregrine:a North American sage of tenacity and teamwork. The Peregrine Fund
76. ^ "Rare peregrine falcons raise four chicks in Nottingham". Bbc.co.uk. 2011-05-11. http://www.bbc.co.uk/news/uk-england-nottinghamshire-13358253. Retrieved 2011-08-30. 
77. ^ "Peregrine". The Rspb. 2011-08-22. http://www.rspb.org.uk/wildlife/birdguide/name/p/peregrine/index.aspx. Retrieved 2011-08-30. 
78. ^ The Royal Society for the Protection of Birds (2003), Peregrine Falcon: Threats, http://www.rspb.org.uk/wildlife/birdguide/name/p/peregrine/threats.asp, retrieved 2008-05-26 
79. ^ "London | Falcon eggs hatch on tower block". BBC News. 2005-06-03. http://news.bbc.co.uk/1/hi/england/london/4605621.stm. Retrieved 2011-08-30. 
80. ^ Navarro, Mireya (2009-02-12). "Record Number of Peregrine Falcons in New York State". The New York Times. http://www.nytimes.com/2009/02/13/nyregion/13falcon.html. Retrieved 2009-02-13. 
81. ^ "Santa Cruz Predatory Bird Group WebCam". Santa Cruz Predatory Bird Research Group. 2011-03-11. http://www2.ucsc.edu/scpbrg/nestcamSF.htm. Retrieved 2011-03-11. 
82. ^ Krech (2009), pp. 92–95
83. ^ Evans (1990), pp. 79–86
84. ^ "Office Of The Dean Of Students :: Bowling Green State University History And Traditions". Bowling Green State University. http://www.bgsu.edu/offices/sa/deanofstudents/traditions/. Retrieved 2011-08-31. 
85. ^ Shalaway, Scott (2007-09-02). "Quarters Reflect High Interest in Nature". The Charleston Gazette. 

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• Döttlinger, H. (2002): The Black Shaheen Falcon. Books on Demand. ISBN 3-8311-3626-2
• Döttlinger, H. & M. Nicholls (2005): Distribution and population trends of the 'black shaheen' Peregrine Falcon Falco peregrinus peregrinator and the eastern Peregrine Falcon F. p. calidus in Sri Lanka. Forktail 21: 133–138 PDF fulltext
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• Groombridge, J. J.; Jones, C. G.; Bayes, M. K.; van Zyl, A.J.; Carrillo, J.; Nichols, R. A. & Bruford, M. W. (2002): A molecular phylogeny of African kestrels with reference to divergence across the Indian Ocean. Molecular Phylogenetics and Evolution 25(2): 267–277. doi:10.1016/S1055-7903(02)00254-3 (HTML abstract)
• Helbig, A.J.; Seibold, I.; Bednarek, W.; Brüning, H.; Gaucher, P.; Ristow, D.; Scharlau, W.; Schmidl, D. & Wink, M. (1994): Phylogenetic relationships among falcon species (genus Falco) according to DNA sequence variation of the cytochrome b gene. In: Meyburg, B.-U. & Chancellor, R.D. (eds.): Raptor conservation today: 593–599. PDF fulltext
• Krech, Shepard (2009) Spirits of the Air: Birds & American Indians in the South. University of Georgia Press
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• Nittinger, F.; Haring, E.; Pinsker, W.; Wink, M. & Gamauf, A. (2005): Out of Africa? Phylogenetic relationships between Falco biarmicus and other hierofalcons (Aves Falconidae). Journal of Zoological Systematics and Evolutionary Research 43(4): 321–331. doi:10.1111/j.1439-0469.2005.00326.x PDF fulltext
• Peters, J. L.; Mayr, E. & Cottrell, W. (1979): Check-list of Birds of the World. Museum of Comparative Zoology.
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• Wink, M. & Sauer-Gürth, H. (2000): Advances in the molecular systematics of African raptors. In: Chancellor, R.D. & Meyburg, B.-U. (eds): Raptors at Risk: 135–147. WWGBP/Hancock House, Berlin/Blaine. PDF fulltext
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• Wink, M.; Döttlinger, H.; Nicholls, M. K. & Sauer-Gürth, H. (2000): Phylogenetic relationships between Black Shaheen (Falco peregrinus peregrinator), Red-naped Shaheen (F. pelegrinoides babylonicus) and Peregrines (F. peregrinus). In: Chancellor, R.D. & Meyburg, B.-U. (eds): Raptors at Risk: 853–857. WWGBP/Hancock House, Berlin/Blaine. PDF fulltext
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External links

External videos
	Peregrine Falcon Banding, Metropolitan Transportation Authority Bridges and Tunnels; June 3, 2010; 3-minute YouTube video clip
	Throgs Neck Bridge Peregrine Banding 2011, Metropolitan Transportation Authority Bridges and Tunnels; May 27, 2011; 10:54 YouTube video clip

• Identification guide (PDF) by Javier Blasco-Zumeta

Conservation organizations

• Arctic Raptors – Ongoing research with raptors in the Canadian Arctic
• Falcon Research Group
• Peregrine Falcon Fund
• The Canadian Peregrine Foundation
• Peregrine Falcon Recovery Project (Manitoba)

Video and other media of Peregrines

• Science on the SPOT: Peregrine Falcons Up Close – KQED QUEST
• Falcon Fascination – KQED QUEST
• Derby Cathedral Peregrine Project, UK. Links to webcams and video sequences
• Peregrine Falcon videos on the Internet Bird Collection
• The Raptor Resource Project. Links to Peregrine Falcon webcams
• Peregrines on Brussels Cathedral
• Photo documentation of Peregrines returning to south California beach cliffs after over 50 years absence
• Nottingham Trent University, where peregrines return to breed on the top of the Newton building every year. Includes images and webcam.
• Worcester Peregrine Falcon Project, UK. Includes feeds from 'Peregrines in Worcester' Facebook Fan page, YouTube & Flickr photo groups