Peregrine Falcon

Peregrine Falcon
Adult with prey in Nova Scotia, Canada
Conservation status
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-grey back, barred white underparts, and a black head and "moustache". As is 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 (high speed dive),[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]

Contents

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% 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 below.

The back and the long pointed wings of the adult are usually bluish black to slate grey 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, coloured 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 "moustache" 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

Falco peregrinus was first described under its current binomial name by English ornithologist Marmaduke Tunstall in his 1771 work 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, and the descendants further breeding with Sakers.[21]

Today, Peregrines are regularly paired 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 coloured 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).[22]

Subspecies

Numerous subspecies of Falco peregrinus have been described, with 19 accepted by the 1994 Handbook of the Birds of the World,[4][5][23] 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 ranges of these 19 subspecies:

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.[41] 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.[42] Barbary Falcons breed at different times of year than neighboring Peregrine Falcon subspecies,[5][23][43][44][45][46][47] but there are no postzygotic reproduction barriers in place.[48] There is a 0.6–0.7% genetic distance in the Peregine-Barbary Falcon ("peregrinoid") complex.[43]

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

Ecology and behavior

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.[49]

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.[50] 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.[51] In 2005, Ken Franklin recorded a falcon stooping at a top speed of 389 km/h (242 mph).[52] 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 eagles or large owls.[53]

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,[54] Ceratophyllus garei (a flea), and Hippoboscidae flies (Icosta nigra, Ornithoctona erythrocephala).[55]

Feeding

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).[56] Prey also include a small raptor, the American Kestrel.[57] Other than bats taken at night,[58] 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.[59] 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).[58]

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.[58] 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.[60] 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.[60] 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.[50]

Reproduction

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.[61] 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][61]

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.[61] 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.[62]

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.[63] The eggs are white to buff with red or brown markings.[63] 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][50][53]

After hatching, the chicks (called "eyases"[64]) are covered with creamy-white down and have disproportionately large feet.[61] The male (called the "tiercel") and the female (simply called the "falcon") both leave the nest to gather prey to feed the young.[50] The hunting territory of the parents can extend a radius of 19 to 24 km (12–15 miles) from the nest site.[65] Chicks fledge 42 to 46 days after hatching, and remain dependent on their parents for up to two months.[66]

Relationship with humans

Falconry

The Peregrine Falcon has been used in falconry for more than 3,000 years, beginning with nomads in central Asia.[61] 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,[67] and were used to intercept homing pigeons during World War II.[68]

Peregrine Falcons have been successfully bred in captivity, both for falconry and for release back into the wild.[69] Until 2004 nearly all Peregrines used for falconry in the US were captive-bred from the progeny of falcons taken before the US 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. The successful recovery program was aided by the effort and knowledge of falconers – in collaboration with The Peregrine Fund and state and federal agencies – through a technique called hacking. 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,[70] it is common practice not to publicize unprotected nest locations.[71]

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.[72] 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.[60][73] In several parts of the world, such as the eastern United States and Belgium, this species became extirpated (locally extinct) as a result.[66] An alternate point of view is that populations in the eastern North America had vanished due to hunting and egg collection.[26]

Recovery efforts

In the United States, Canada, Germany and Poland, wildlife services in Peregrine Falcon recovery teams breed the species in captivity.[74] 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.[49] 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.[75] 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.[75]

Worldwide recovery efforts have been remarkably successful.[74] The widespread restriction of DDT use eventually allowed released birds to breed successfully.[49] The Peregrine Falcon was removed from the U.S. Endangered Species list on August 25, 1999.[49][76]

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.[77]

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.[78][79] 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.[80] In many parts of the world Peregrine Falcons have adapted to urban habitats, nesting on cathedrals, skyscraper window ledges, tower blocks,[81] and the towers of suspension bridges. Many of these nesting birds are encouraged, sometimes gathering media attention and often monitored by cameras.[82][83]

Cultural significance

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.[84] 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.[85]

Since 1927, the Peregrine Falcon has been the official mascot of Bowling Green State University in Bowling Green, Ohio.[86] The 2007 U.S. Idaho state quarter features a Peregrine Falcon.[87]

Birds portal


References

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  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)
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  30. ^ Also called "Kleinschmidt's Falcon", but this might equally refer to F. p. kleinschmidti which is a junior synonym of japonensis,
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  37. ^ The shaheen (شاهین) of Arabic and Persian writers are usually Barbary Falcons; those in Indian (शाहीन) and Pakistani (شاہین) sources normally refer to peregrinator.
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  47. ^ Wink et al. (2004)
  48. ^ Blondel (1999)
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  54. ^ 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)
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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
Conservation organizations
Video and other media of Peregrines