Peregrine Falcon

Peregrine Falcon
Adult of subspecies pealei or tundrius, 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 cosmopolitan bird of prey in the family Falconidae. It is a large, crow-sized falcon, with a blue-gray back, barred white underparts, and a black head and "moustache". It can reach speeds over 320 km/h (200 mph) in a stoop,[4] making it one of the fastest creatures on the planet.[5] As is common with bird-eating raptors, the female is much bigger than the male.[6][7] Experts recognize 17 to 19 subspecies which vary in appearance and range; there is disagreement over whether the distinctive Barbary Falcon is a subspecies or a distinct species.

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 fact makes it the world's most widespread bird of prey.[8] Both the English and scientific names of this species mean "wandering falcon", referring to the migratory habits of many northern populations.

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.[9] The Peregrine Falcon became an endangered species in many areas due to the use of pesticides, especially DDT. Since the ban on DDT from the beginning of the 1970s onwards, the populations recovered, supported by large scale protection of nesting places and releases to the wild.[10]

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).[6][11] 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.[12] 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.[11] The white to rusty underparts are barred with thin clean bands of dark brown or black.[13] 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.[14] The cere is yellow, as are the feet, and the beak and claws are black.[15] 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][6][7] The immature bird is much browner with streaked, rather than barred, underparts, and has a pale bluish cere and orbital ring.[6]

Taxonomy and systematics

F. p. anatum in flight, Morro Bay, California

This species was first described by Marmaduke Tunstall in his 1771 Ornithologia Britannica under its current binomial name.[16] The scientific name Falco peregrinus, means "wandering falcon" in Latin.[17] Indeed, the species' common name refers to its wide-ranging flights in most European languages.[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.[4]

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.

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

Breeding ranges of the subspecies

Numerous subspecies of the Peregrine have been described, with 19 accepted by the Handbook of the Birds of the World.[6][7][22]

Australian race F. p. macropus
Captive bird of the subspecies pealei
Subspecies minor, illustration by Keulemans, 1874
Painting of subspecies babylonicus by John Gould

These last two races are often split as Barbary Falcon Falco pelegrinoides.[7] There is a 0.6–0.7% genetic distance in the Peregine-Barbary Falcon ("peregrinoid") complex.[36] These birds inhabit arid regions from the Canary Islands along the rim of the Sahara through the Middle East to Central Asia and Mongolia. They have a red neck patch but otherwise differ in appearance from the Peregrine proper merely according to Gloger's Rule.[37] 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.[7] The Barbary Falcon's shoulder and pelvis bones are stout by comparison with the Peregrine, and its feet are smaller.[38] They have no postzygotic reproduction barriers in place,[39] but they breed at different times of year than neighboring Peregrine Falcon subspecies.[7][22][36][40][41][42][43]

Ecology and behavior

Silhouettes 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.[13] 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.[44] The Peregrine Falcon is often stated to be the fastest animal on the planet in its hunting dive, the stoop,[5] 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.[4] 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.[45] In 2005, Ken Franklin recorded a falcon stooping at a top speed of 389 km/h (242 mph).[46] A video of one of the dives can be seen in this link.

The life span in the wild is up to 15.5 years.[7] Mortality in the first year is between 59–70%, declining to between 25–32% in adults.[7] Apart from anthropogenic threats like collision with human-made objects, the Peregrine may be killed by large eagles or large owls.[27] 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[47] Ceratophyllus garei (a flea), and Hippoboscidae flies (Icosta nigra, Ornithoctona erythrocephala).[48]

Feeding

Immature 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 doves, waterfowl, songbirds, waders and pigeons.[15] 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 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).[49] Prey also include the small raptor, the American Kestrel.[50]Other than bats taken at night,[51] it rarely hunts small mammals, but will on occasion take rats, voles, hares, mice and squirrels; the coastal populations of the large subspecies pealei feed almost exclusively on seabirds.[14] In the Brazilian mangrove swamp of Cubatão, a wintering falcon of the subspecies tundrius was observed while successfully hunting a juvenile Scarlet Ibis.[52] Insects and reptiles make up a small proportion of the diet, which varies greatly depending on what prey is available.[15] In urban areas, the main item 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, such as Mourning Doves, Common Swifts, Northern Flickers, Common Starlings, American Robins and various corvids.[51]

The Peregrine Falcon hunts at dawn and dusk, when prey are most active, but in cities also nocturnally, 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.[51] It requires open space in order to hunt, and therefore often hunts over open water, marshes, valleys, fields and tundra. It searches for prey either from a high perch or from the air.[53] Once prey is spotted, it begins its stoop, folding back the tail and wings, with feet tucked.[14] 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.[54] To protect their eyes, the falcons use their nictitating membranes (third eyelids) to spread tears and clear debris from their eyes while maintaining vision. Prey is struck and captured in mid-air; the Peregrine Falcon strikes its prey with a clenched foot, stunning or killing it, then turns to catch it in mid-air.[53] The Peregrine will drop it to the ground and eat it there if it is too heavy to carry. Prey is plucked before consumption.[54]

Reproduction

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. The 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.[11] 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. The Peregrine Falcon is territorial during the breeding season; nesting pairs are usually more than 1 km (0.6 miles) apart, and often much farther, even in areas with large numbers of pairs.[55] 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 pair defends the chosen nest site against other Peregrines, and often against ravens, herons, gulls and (in ground nest) mammals like foxes, wolverines, felids, bears and wolves.[55] 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.[56]

The Peregrine Falcon nests in a scrape, normally on cliff edges or, today regularly in many parts of its range, on tall buildings or bridges. Cliff nests are generally located under an overhang, on ledges with vegetation, and south-facing sites are favored.[14] 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, there was a large population of peregrines in central and western Europe using the disused nests of other large birds.[15] 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.[11] In remote, undisturbed areas such as the Arctic, steep slopes and even low rocks and mounds may be used as nest sites. The human-made structures used for breeding closely resemble the natural cliff ledges that the Peregrine prefers for its nesting locations.[6][55]

Chick

Mostly three to four eggs (range 1–5) are laid in the scrape.[57] The eggs are white to buff with red or brown markings.[57] They are incubated for 29 to 33 days, mainly by the female.[14] The male also helps with the incubation of the eggs over day, but at night only the female incubates. 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 (the Australian subspecies macropus may breed as late as November and equatorial populations may nest anytime between June and December). The female generally lays another clutch if the eggs are lost early in the nesting season, though this is extremely rare in the Arctic owing to the short summer season. As a result of some infertile eggs and natural losses of nestlings, the average number of young found in nests is 2.5, and the average number that fledges is about 1.5.[6][27][54]

After hatching, the eyases[58], or chicks, are covered with creamy-white down and have disproportionately large feet.[55] The male, which is called the "tiercel", and the female, which is simply called the "falcon", both leave the nest to gather prey to feed the young.[54] The hunting territory of the parents can extend a radius of 19 to 24 km (12–15 miles) from the nest site.[59] Chicks fledge 42 to 46 days after hatching, and remain dependent on their parents for up to two months.[60]

Relationship with humans

Pesticides

The Peregrine Falcon became an endangered species because of the use of organocholorine pesticides, especially DDT during the 1950s, 60s, and 70s.[61] 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.[53][62] In several parts of the world, such as the eastern USA and Belgium, this species became locally extinct as a result.[60]

Illegal collectors

Peregrine eggs and chicks are often targeted by black marketeers[63] and unscrupulous egg collectors, so it is normal practice not to publicize unprotected nest locations.[64]

Falconry

The Peregrine Falcon was used in falconry for more than 3,000 years, beginning with nomads in central Asia.[55] Due to its ability to dive at high speeds, it was highly sought-after and generally used by experienced falconers.[12] 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]

Recovery efforts

In the USA, Canada, Germany and Poland, Wildlife services in Peregrine Falcon recovery teams breed the species in captivity.[67] 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.[44] 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.[68] 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.[68] Worldwide recovery efforts have been remarkably successful.[67] The widespread restriction of DDT use eventually allowed released birds to breed successfully.[44] The Peregrine Falcon was removed from the U.S. Endangered Species list on August 25, 1999.[44][69]

Current status

In the USA

Many Peregrine Falcons have settled in large cities, nesting on cathedrals, skyscraper window ledges, and the towers of suspension bridges. As early as 1946, Peregrine Falcons were nesting atop Philadelphia City Hall, which is believed to be among the first artificial structures in the world to be used as a nest site by this species.[70] In Virginia, state officials working with students from the Center for Conservation Biology of the College of William and Mary in Williamsburg successfully established nesting boxes high atop the George P. Coleman Memorial Bridge on the York River, the Benjamin Harrison Memorial Bridge and Varina-Enon Bridge on the James River, and at other similar locations. Thirteen new chicks were hatched in this Virginia program during a recent year. Over 250 falcons have been released through the Virginia program.[71] The New York State Department of Environmental Conservation reported that there were 67 pairs of Peregrine Falcons in the state during 2008.[72]

In Canada

Peregrine Falcon on the Ambassador Bridge in Windsor, Ontario

As in the USA, Peregrine Falcons have moved into major Canadian cities.

In Winnipeg, Manitoba, the first post-population crash wild nest occurred in 1989 on the Delta Winnipeg Hotel (now the Radisson Hotel Downtown-Skyview) as a result of recovery efforts begun in 1980-1981 by the Peregrine Falcon Recovery Project (Manitoba). The Radisson site has been occupied continuously since 1989 and offspring from this nest can be found across the Canadian Prairie Provinces and the US Midwest States. The popular CBC Manitoba/Shaw FalconCam has been following the lives (and deaths) of the Radisson peregrines since 2006[73]. In 1993, peregrines began nesting at the McKenzie Seeds Building in Brandon, Manitoba and have likewise had a similarly long and successful nesting history.

In Hamilton, there have been a pair of birds nesting on a ledge overhanging a window of the downtown Sheraton hotel every year since 1994.[74] The Hamilton Community Peregrine Project was setup in 1995 to observe the falcons and provide care should they need it.[75] This includes banding birds that have successfully fledged and caring for birds which crash onto the busy streets below the nest. They also have a camera aimed at the nesting site, which can be viewed on their website.[76]

The University of Alberta in Edmonton, Alberta also houses a pair of breeding Peregrine Falcons atop the Clinical Sciences building, amongst others on the campus who are actively monitored via webcams. The female of this pair originated in Winnipeg, Manitoba while the male is from Alberta. Several birds have also been tagged with radio transmitters, permitting study of migration patterns.[77]

In Britain

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. Peregrines now breed in many mountainous and coastal areas, especially in the west and north, and nest in some urban areas, capitalizing on the urban pigeon populations for food.[78]

Cultural references

References

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  3. Friedmann, H. (1950), "i rock birds of North and Middle America", U.S. National Museum Bulletin 50 (11): 1–793 
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  18. e.g. French faucon pèlerin, German Wanderfalke, Italian falco pellegrino, Polish sokół wędrowny, Slovak sokol sťahovavý, Swedish pilgrimsfalk
  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. 22.0 22.1 Vaurie (1961)
  23. 23.0 23.1 23.2 American Ornithologists' Union (1910):p.164
  24. The shaheen (شاهین) of Arabic and Persian writers are usually Barbary Falcons; those in Indian (शाहीन) and Pakistani (شاہین) sources normally refer to peregrinator.
  25. Döttlinger & Nicholls (2005)
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  27. 27.0 27.1 27.2 Michigan Department of Natural Resources (2007)
  28. 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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  30. 30.0 30.1 American Ornithologists' Union (1910):p.165
  31. 31.0 31.1 31.2 Proctor, N. & Lynch, P. (1993):p.13
  32. Vaurie, 1961
  33. 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.
  34. Mayr (1941)
  35. Peters, J. L.; Mayr, E. & Cottrell, W. (1979):p.423
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  38. (Vaurie, 1961)
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  43. Wink et al. (2004)
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  45. Tucker (1998)
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  47. 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)
  48. Raidal et al. (1999), Raidal & Jaensch (2000), Dewey & Potter (2002), Dalgleish (2003)
  49. http://bna.birds.cornell.edu/bna/species/660/articles/foodhabits
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  56. http://bna.birds.cornell.edu/bna/species/660/articles/behavior
  57. 57.0 57.1 Peterson, R. T (1976):p.171
  58. Taken from http://www.raptorresource.org/facts.htm
  59. Towry (1987)
  60. 60.0 60.1 Snow (1994)
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  62. Brown (1976)
  63. Trade in wild-caught Peregrine Falcons and their eggs and young is illegal in most jurisdictions. Falconers are advised to demand valid documentation even if they are able to legally purchase this species.
  64. American Birding Association (2005), Code of Birding Ethics, American Birding Association, Inc., http://www.aba.org/about/ethics.html, retrieved 2008-05-26 
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External links

Conservation organizations
Video and other media of Peregrines