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Figure 1 . Formerly extirpated from much of original range by synthetic organic chemicals such as DDT (see below and Conservation and management, below); reoccupancy and restoration still incomplete in 2001, especially in central and s. Canada and midwestern and e. U.S., where much of distribution is urban, but progressing rapidly (Enderson et al. 1995a). Thus, distribution as given is subject to change. Following information as of 2000–2001, provided by federal, provincial, and state agencies, recovery-team members, and individuals documenting recovery.
Alaska, Canada, and Greenland. In the west from w. Aleutian Is. (Attu 53°N, 172°35'E) east through Aleutians and Alaska Peninsula, then north along coastal w. Alaska, where spotty but with local concentrations such as Norton Sound (64°N; absent from Nunivak, St. Matthew, Pribilof, St. Lawrence, and Diomede Is.), to North Slope of Alaska (about 70°N; locally, largely riverine, but moving into coastal plain around lakes north of 70°; R. Ritchie pers. comm.); Yukon Territory, Northwest Territories, and Nunavut locally north to about 73°30'–74°N in parts of Banks I., Victoria I., Melville I. to 75°N; Bothia Peninsula to about 72°N, to Baffin I. and then to ice-free parts of w. Greenland from Thule (Qaanaaq), about 76–77°N (scarce; Burnham 1996) south around tip and north on east side at least to Angmagssalik (65°40'N), although seen as high as Thomsen Land (75°N) and Germania Land (76°50'–77°N; Boertman 1994), in breeding season. Then southward through Alaska, Yukon (spotty and local), Northwest Territories, Nunavut, British Columbia (poorly documented inland), n. and central Alberta, Saskatchewan (Saskatoon and Regina, where urban breeders), Manitoba (Winnipeg and Brandon; urban sites), s. Ontario (e.g., along Great Lakes), s. Quebec (along St. Lawrence River), Labrador (mainly coastal; Todd 1963a; Canadian Wildlife Service 1988; G. Chilton, G. A. Court, U. Ban-asch, G. Holroyd pers. comm.). Absent as breeder from Newfoundland.
United States. Locally (see Figure 1) through northern tier of states, most midwestern and eastern states. Many introduced into urban areas (e.g., Milwaukee, WI; Chicago, IL; Fort Wayne, IN; New York, NY); spotty and local (urban) in midwestern states (e.g., Nebraska, Missouri, Iowa); spotty and local in most eastern states (e.g., New England, Pennsylvania, New York, New Jersey, Maryland, Delaware, Virginia, N. Carolina, S. Carolina, Alabama; Burnham 1995; J. Castrale, M. Amaral, C. Koppie pers. comm.). Widely in western states (e.g., most of Utah, Arizona, w. Colorado, w. and n. California; see Figure 1), but absent from N. Dakota (although bred in urban Fargo, ND, in 2001; B. Tordoff pers. comm.), Florida, Oklahoma, Arkansas, Alabama, Mississippi, Louisiana, Texas (except sw. Rio Grande region), and Nevada (except along borders with Utah, California, and Arizona; Burnham 1995, TJC, CMW, WGH).
Mexico. Baja California and islands of Gulf of California (absent from Guadalupe I.) and Sierra Madre Occidental and Oriental in Sonora, Chihuahua, w. Coahuila (locally), Durango (but poorly defined); south to near Ciudad Victoria, Tamaulipas (23°44'N; W. Wimsatt in Hickey 1969, Porter et al. 1988, D. Lan-ning pers. comm., WGH); perhaps at latitude of s. San Luis Potosí on Mexican Plateau (22°30'N; R. Munro-Wilson pers. comm.).
Middle America and Caribbean. Extralimital nesting in 1999 confirmed for Holguín Province of e. Cuba (Regalado and Cables 2000) and one record given for Dominica (details unavailable; Raffaele et al. 1998). A mid-twentieth–century record suggested for cliff on Lago de Managua, Nicaragua (M. W. Nelson pers. comm.). Southern subspecies cassini over most of South America but absent as breeder from Central America, most of Amazon and Orinoco River basins.
Nearctic. Northern limits of range (Figure 1) variable between coasts, more northward (except for Greenland) on west coast, throughout Aleutian Is. and Gulf of Alaska (Kodiak I.); on east coast north to Gaspé Peninsula, limit of range apparently function of urban locations with good prey populations or areas of abundant prey (e.g., open water or edge of ice sheet); seemingly shifted northward since reintroduction began in 1970s (see American Ornithologists' Union 1983, where range given as south from central and s. U.S.). Inland records for mid-Dec or Jan (Christmas Bird Counts; presumed wintering) from s. British Columbia (at least 49°N); in Alberta (generally south of 52°N) at such locations as Calgary, Cochrane, Bruce, and Monarch (Semenchuk 1992); environs of Winnipeg, Manitoba (B. Jones, W. Neily pers. comm.); rarely extreme s. Saskatchewan (since 1942, seen 1980, 1981, 1988; M. Williams pers. comm.); s. Ontario; Montreal (D. Bird, U. Banasch pers. comm.) and Gaspé Peninsula, Quebec; s. New Brunswick, and Nova Scotia (Halifax); occasionally in s. Greenland (older specimen records; Salomonsen 1950). In most major cities from Washington State, Minnesota, Wisconsin (Septon 2000), coastal Maine south through U.S.
Neotropical. Spotty, with concentrations on coasts, in cities, and in wetlands (perhaps function of observer distribution; e.g., Risebrough et al. 1990). Throughout Caribbean (West Indies) south through Mexico, Central America to at least Chiloe I., Chile (42°30'S); high numbers coastally in at least Ecuador to n. Chile; n. South America and Netherlands Antilles (rare; Voous 1983) south throughout most of Brazil (except perhaps Brazilian Shield of s.-central Brazil). Of 117 locations in 18 Brazilian states where recorded, most in agricultural and urban habitats of ne. and s. Brazil (perhaps a function of observer concentration; Albuquerque 1978, Silva E Silva 1996), then south through Bolivia, Uruguay, Paraguay, and Argentina at least to Bahía Blanca at 38°S (W. G. Vasina pers. comm.) but probably to Gulfo San Matias at 42°S. West coast of South America to 40°S and about 4,100 m in Andes (Fjeldså and Krabbe 1990).
Worldwide breeder, as far east in Southern Hemisphere as Fiji and south to cape of South Africa and Tasmania, but absent from most of Saharan Africa, parts of central and e. Asian steppes, New Zealand, Iceland, and central Pacific Ocean, and Antarctica. In Palearctic, northern populations mainly migratory moving south to cape region of South Africa, Indian subcontinent, and Indonesia. Most southern populations and island endemics resident (Cade 1982b, White and Boyce 1988, White et al. 1994b).
Most of breeding range except n. Scandinavia and tundra of Asia. Found in United Kingdom, most of Europe south of about 58°N, Asia south of 50°N, throughout Africa, Indonesia, Philippines, New Guinea and Australia, and eastward to Vanuatu, New Caledonia, and Fiji (Cramp and Simmons 1980a, Stepanyan 1990, White et al. 1994b).
Widespread during migration; e.g., mountains at 3,800 m above sea level (far above breeding range), offshore Atlantic routes (Kerlinger et al. 1983), mid-Pacific islands (perhaps not from North America; Clapp and Woodward 1968, Woodward 1972b). In schematic outline, continental populations migrate in more or less "leap-frog" fashion (Schmutz et al. 1991a, Mcgrady et al. 2002), but not uniformly. Northernmost tundra breeders generally move farthest south to central Argentina and Chile, with breeders farther south traveling shorter distances, but arctic and boreal nesters overlap considerably in winter distribution; some mid- to southern populations are resident, others at high elevations in Rocky Mtns. move south into Mexico are resident (White White 1968a, White 1968b; Yates et al. in Cade et al. 1988; Enderson et al. 1991; Schmutz et al. 1991a). Individuals on nonbreeding (wintering) grounds in coastal e. Mexico and Central America with satellite transmitters moved to breed in Canadian Arctic and w. Greenland (Mcgrady et al. 2002, T. Maechtle pers. comm.), these birds winter-ing farther north than migrants from Wyoming and Colorado wintering in Sinoloa, Mexico (Molina and Cade 1990). Pale-colored falcons, presumably from tundra populations (not based on specimens), migrate south to at least Chiloe I., Chile (42°30'S) where they are seen at least Nov through Feb (R. Schlatter via C. Sadler pers. comm.). Most breeding populations not above 75°N and thus some one-way migration 12,400-13,300 km or more; one banded in Northwest Territories, Canada recovered 14,500 km south in Chaco, Argentina, less than 4 mo latter (Kuyt 1967). Two records of tundrius from Iceland, 1 on 30 Jul 1961, 1 on 16 Oct 1985 (British Birds 1997)
Despite broad-front migration and widespread nature of movements (Anderson et al. and Yates et al. in Cade et al. 1988), there are clearly defined routes where Peregrines concentrate: along leading lines or coastal areas of prime habitat (barrier islands), particularly on Eastern Seaboard (Chincoteague and Assateague Is., MD-VA), Gulf Coast (Padre I., TX), and e. Mexico (Veracruz; Thiollay 1980, Heintzelman 1986, Hunt et al. 1975, Hunt and Ward 1988, Ward et al. 1988, Chavez-Ramirez et al. null, HawkWatch International pers. comm.). Lesser concentrations along shores of Great Lakes, West Coast of U.S., w. Mexico, and eastern front of Rocky Mtns. In South America, large numbers on west coast from Ecuador to Chile (Schoonmaker et al. 1985; Risebrough et al. 1990; C. M. Anderson, C. Gonzales, O. Beingolea pers. comm.).
Typical migration from w. Greenland is west across Davis Strait, south in Canada and along U.S. East Coast either through Florida Keys to Caribbean (West Indies) and Central and South America or west around Gulf of Mexico, through Texas and Mexico to Central and South America. Large numbers now seen at off-shore oil rigs in Gulf of Mexico, associated with transgulf flights of passerines and shorebirds (A. Wormington in Pendergrass 2000, R. W. Russell pers. comm.). At autumn banding sites, adults tend to arrive prior to first-year birds; ratios about 1:1.7 along Great Lakes, 1:5.7 and 1:3.9 on Atlantic and Gulf Coasts, respectively, perhaps reflecting age difference in staging behavior (Mueller et al. 2000).
Timing elsewhere generally similar; e.g., central Alberta spring migrants between 20 Apr-31 May, mid-dates for adults about 8 May, immatures 15 May (Dekker 1984b). Missouri, spring peak last week of Apr, with autumn migration starting late Aug (peak late Sep-early Oct; seldom after Oct; Robbins and Easterla 1992). Ohio (mainly along Lake Erie) in spring, most 20 Apr-15 May; autumn, most 25 Sep-20 Oct, some as late as mid-Nov (Peterjohn 1989b). Cape May, NJ, common in autumn, with total of 1,503 in 1996; daily maxima 206 on 1 Oct 1996 (96 in 1 h); fewer in spring, with peak first week of May (Sibley 1997). Bermuda, mainly autumn, 13 Sep-late Nov; rare in spring, late Mar to mid-Apr (Amos 1991). Costa Rica, mainly on Caribbean coast, occasionally Pacific lowlands; spring, Mar-early May, autumn, mid-Sep through Oct (Stiles and Skutch 1989). Throughout West Indies, but uncommon to rare, Oct-Apr (Raffaele et al. 1998).
Until recently, most data came from collected specimens and banding returns (see Cade et al. 1988). More recently, Peregrines tracked with radio transmitters by aircraft (W. W. Cochran, private compilation from Illinois Nat. Hist. Sur. Repts. no. 304, 305, 312, 313) or satellite (Henny et al. 1996, Seegar et al. 1996, Hendricks 1997a, T. Maechtle pers. comm.). Selected literature data from such techniques: Adult female trapped in mid-Oct 1993 on Assateague I., MD, flew to Guatemala, Honduras, Nicaragua, Ecuador, and then Bolivia; crossed Andes Mtns. to headwaters of Amazon River, finally ending up 6 Nov at 3,950 m at a saline playa, Salar de Antofalla, in the altiplano of Argentina, staying at oasis in the salar . Northward-migrating female trapped 22 Apr 1994 on Padre I., TX; stayed on eastern front of Rocky Mtns. in U.S., crossed Yukon Territory, and arrived at breeding grounds on Yukon River, AK, 31 May; left there 31 Aug, following a similar southward but more easterly route, arriving on Yucatán Peninsula 22 Sep, leaving there 3 Oct for Panama; then left Panama 19 Oct, crossed Andes, and arrived in Argentina 21 Nov. Lastly, 1 immature northward migrating female trapped 22 Apr on Padre I., TX, moved directly north through central Midwest to King William I., Nunavut, arriving 26 May; left 12 Oct, taking different route south, along west coast of Hudson Bay, leaving area of Churchill, Manitoba, 8 Nov, then south through e. Midwest to Gulf Coast east of New Orleans and to Yucatán Peninsula, apparently moving across Gulf of Mexico, by 25 Nov (Henny et al. 1996).
Extralimital returns: A first year female (perhaps tundrius) trapped 14 Oct 1991 on Padre I., TX (C. M. Anderson, T. L. Maechtle pers. comm.), recovered 8 Dec 1993 at Misono-cho, Wakayama Pref., Japan (caught alive but died). One banded, also with color band, as nestling (anatum) on Colorado River, Glen Canyon NRA, AZ (M. Britten pers. comm.), photographed several times 17 Jan 1993 to 7 Feb 1993 in central Japan area of Mishima-shi. Both returns thought to have been assisted by marine vessel from west coast of North America to Japan where marine traffic is heavy.
Data from radio transmitters from 7 individuals (see compilation from W. W. Cochran as above, Cochran 1975; also see Timing and routes of migration, above). Typical 24-h period included 17 h on perch, 6 h in migratory flight (range 1-9), 1 h hunting. When not migrating because of weather, loafing increased to 23 h/d. Migration on about 6 of 7 d, generally mid-morning to late afternoons. On average, flights 192 km at 33 km/h, but one individual went from Chicago to Tennessee in 1 d-560 km at 64 km/h. Efficiency in hunting during migration not necessarily a function of falcon's age. A first-year bird averaged 53 min/d hunting, while one adult spent 93 min/d hunting before prey capture over a 5-d period. Chavez-Ramirez et al. (Chavez-Ramirez et al. null) found spring migrants flew more hours on average (12.02 h/d) than fall migrants (8.75 h/d).
Considered a low-altitude migrant (mostly <100 m above ground level up to 900 m; Kerlinger 1989a), but soars more than generally recognized. Average ground speed for flapping flight about 49 km/h (Cochran and Applegate 1986). Adults may use different routes from immatures over part of the migration. Telemetry data show spring migrants remain on Padre I. for average of 8.4 d (range 3-28; Hunt and Ward 1988); autumn migrants also stage on Padre I. In coastal Peru, ratio of adults to immatures 2:1, with most immatures seen in early winter (but differences may also suggest greater immature overwintering mortality; Schoonmaker et al. 1985). No apparent difference in time of arrival of the sexes on breeding grounds in northern latitudes (Court et al. 1988b, Kerlinger 1989a), although male often appears to arrive at cliff sites first.
Not documented or clearly understood. Complex because migration distances vary, in part with breeding latitude. However, somewhat regular appearance at observation sites (e.g., barrier islands of Atlantic Seaboard and Gulf of Mexico) suggest Peregrines are less affected by passing of cold-weather fronts, which are irregular, than some other species. Alternative explanations for migration: movements of main prey items; photoperiod (e.g., predictable appearance at some observations sites and regular movements of transequatorial migrants). Some remain at certain northern latitudes (e.g., n. Pacific) where prey is available, while other populations at same or similar latitudes (e.g., Greenland) leave as prey moves; so in this example, photoperiod not regulating migra-tions equally. Some interior populations more likely to move than coastal populations where climate is moderate and prey species remain. Some genetic control may be involved, but many birds of northern origin raised and released in midlatitudes (35-40°N; G. Septon pers. comm.) do not migrate, as do birds of similar wild genetic arctic stocks (e.g., 3 adult females in downtown Milwaukee, WI, that wintered consistently there, differing from wintering behavior of original populations that bred nearby in Wisconsin and Minnesota; see papers in Cade et al. 1988 for above data).
Many terrestrial biomes in the Americas; none seems to be preferred (although perhaps greater densities in tundras and coastally). Subtropical and tropical habitats occupied sparsely or not at all; most commonly occupied habitats contain cliffs, for nesting, with open gulfs of air (rather than in confined areas; see Breeding: nest site, below) and generally open landscapes for foraging (examples in Enderson and Craig 1979, Ellis 1982a, Willey 1986a, Grebence and White 1989). May breed to 3,600 m in Rocky Mtns. (J. Enderson pers. comm., CMW). In some regions, dispersion dendritic along rivers or shorelines of coasts and lakes as nest sites usually associated with water. In addition to natural habitats, many artificial habitats now used (urban, human-built environments such as towers, buildings, etc.). See also Breeding: nest site and Food habits: microhabitat for foraging, below.
Broad array of habitats, including urban. Partial to leading lines such as barrier islands, seacoasts, lake edges, or mountain ranges; also at sea. See Migration, above.
Extreme habitat variability because of enormous geographical range. Other than resident populations, which occupy breeding habitats, may occur in open-relief habitat devoid of cliffs, as in midwest U.S.; man-grove, coastal, or wetland areas, as in Sinaloa, Mexico (Enderson et al. 1991); major river valleys and lake shores; pasture lands; featureless terrain devoid of cover and containing waterbirds or pigeons and doves (e.g., Atacama Desert of coastal Peru); and especially urban areas (Risebrough et al. 1990, Bird et al. 1996).
Historically bred continentwide, from tundra south to nearly southern edge of Mexican Plateau, with good records from 1800s and beginning of 1900s; possible changes on mainland Mexico, but not recorded. Formerly bred in trees from Louisiana (1 nest known; Lowery 1974) north through parts of Mississippi River drainage (Tennessee, Indiana, Illin-ois) and west to Kansas (Goss 1898), and on gravelly/sandy river banks in midcontinent (Hickey 1969), but this distribution never well documented. Clearing of forests in East in 1800s may have broadened distribution, but tree removal along Mississippi River drainage eliminated tree nesters; last nest in Tennessee probably in late 1940s (Spofford 1947b). By early to mid-1900s, sites easily disturbed by humans were gone (north to at least 55°N), but this not well documented; generally most breeding then restricted to areas of larger cliffs throughout this range.
Most complete documentation of former distribution in Hickey 1942, Bond 1946, and Cade 1960, until advent of major decline in numbers and loss of range after about 1950 (Hickey 1969). Well established that this range reduction and population decline resulted primarily from persistent synthetic chlorinated hydrocarbons (see Conservation and management, below). Many events in decline, and eventual recovery of species (mainly anatum), well documented or reconstructed (Hickey 1969, Cade et al. 1988, Enderson et al. 1995a). Changes in distribution too extensive to list in detail. Maximum extent of reduction in distribution may have occurred by 1972–1975 (variable in different places). In U.S., gone from the following locations: east of Rocky Mtns., Nevada, probably Montana, Wyoming, Canadian Maritime Provinces, most of Labrador, Quebec south of about 56°N, Ontario, Manitoba, Saskatchewan, most of Alberta, and n. Yukon Territory (Cade and Fyfe 1970, Fyfe et al. 1976b). Elsewhere distribution and numbers reduced by varying degrees; e.g., California from >100 to 5 pairs, w. Texas and adjacent Mexico from 14 known to 11. Persistence in Mexican highlands through DDT era likely due largely to nonmigratory prey base (e.g., Band-tailed Pigeon [Columba fasciata], other nonmigrants) and lack of mechanized agriculture. Possibly little reduced in Arizona and adjacent Colorado Plateau of s. Utah, but historical distribution in Utah well known only in northern third of state (Porter and White 1973). Knowledge of distributional changes in Mexico marginal and best known in Baja California and Gulf of California; probably disappeared from Pacific coast of Baja (Banks 1969).
Recovery and reoccupancy of historical range well on way by 1980 (Murphy 1990, White et al. 1990), in part because of release of captive-bred falcons by several organizations and breeders in Canada and U.S. (see Enderson et al. 1995a and Conservation and management, below, for details). Known distributional range expanded through 1980s–1990s across tundra and taiga of Alaska, and density even doubled over historically known numbers of 1950s in well-documented places like Colville (32 pairs in 1952 to 57 in 1997) and Yukon (about 20 in 1950 to 47 in 1997) Rivers, AK. In n. Alaska, distributional change has resulted from greater occupancy of tributaries of Colville and smaller rivers to east and on coastal plain (e.g., Barter I.; see Swem and Ambrose 1994; R. J. Ritchie, T. Swem, R. Ambrose pers. comm.; TJC; CMW). By 1996, even though somewhat clumped and spottily distributed, most of historical range reoccupied, although some pairs are urban (e.g., Iowa, Indiana, Illinois, Ohio) and not in original habitat. Distribution in parts of Midwest (e.g., Wisconsin) still largely urban or on human-made structures (Septon et al. 1996). Southern part of Mississippi River drainage (e.g., Tennessee, Louisiana) and most of n. Great Plains not reoccupied by 2000.
No Peregrine Falcon fossils predate Pleistocene (about 1.6 million to 12,000 ybp). Most fossils are mid- to late Pleistocene and widespread at scores of locations in Pleistocene and Holocene prehistoric sites: e.g., Australia, New Caledonia, throughout Europe, Mediterranean region, Caucasus Region (Azerbaijan), North and South America (Brodkorb 1964a, Malez 1988, Weesie 1988, Vickers-Rich et al. 1991). A recent origin is supported by cytochrome- b molecular data suggesting Peregrines separated from an ancestral stock around 2 mybp (Wink 1995a). Within Americas, fossils from Ecuador (Campbell 1976b), Peru (13,900 ybp; Campbell 1979) and several North American Pleistocene and Holocene prehistoric sites; e.g., Alaska, California (tar pits of Rancholabrean age 17,000–27,000 ybp), Arizona (11,940 ybp), New Mexico, Utah, Idaho, Ohio, Arkansas, Illinois, Penn-sylvania, Florida (Emslie 1988c, S. D. Emslie pers. comm.). Several western fossil sites containing Peregrines date from about 40,000 to 29,700 ybp (Porter and White 1973). Some sites in California and Utah containing Peregrine fossils associated with human occupation (Howard 1929, Parmalee 1980); this falcon seemingly used by early humans as food. Those from Utah associated with human sites occupied by Archaic people (9,500–3,000 ybp; Parmalee 1980) around edge of Pleistocene Lake Bonneville that last existed about 10,000–11,000 ybp. Peregrine bones far outnumber Prairie Falcon remains at those midden sites, and Lake Bonneville ecosystem should have been prime habitat for Peregrines based on reconstructed envi-ronment and distribution and density of contemporary and historical peregrine eyries relative to extinct shoreline of Lake Bonneville.