Empidonax traillii - (Audubon, 1828)
Willow Flycatcher
Other English Common Names: willow flycatcher
Taxonomic Status: Accepted
Related ITIS Name(s): Empidonax traillii (Audubon, 1828) (TSN 178341)
French Common Names: moucherolle des saules
Spanish Common Names: Mosquero Saucero
Unique Identifier: ELEMENT_GLOBAL.2.103270
Element Code: ABPAE33040
Informal Taxonomy: Animals, Vertebrates - Birds - Perching Birds
 
Kingdom Phylum Class Order Family Genus
Animalia Craniata Aves Passeriformes Tyrannidae Empidonax
Genus Size: C - Small genus (6-20 species)
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Concept Reference
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Concept Reference: American Ornithologists' Union (AOU). 1998. Check-list of North American birds. Seventh edition. American Ornithologists' Union, Washington, D.C. [as modified by subsequent supplements and corrections published in The Auk]. Also available online: http://www.aou.org/.
Concept Reference Code: B98AOU01NAUS
Name Used in Concept Reference: Empidonax traillii
Taxonomic Comments: Sometimes treated as E. brewsteri, a junior synonym. Formerly regarded as conspecific with E. alnorum as E. traillii, Traill's Flycatcher (AOU 1998). See Phillips (1948) for a review of geographic variation in morphology, with the original descriptions of subspecies alascensis and extimus. Unitt (1987) reviewed infraspecific variation and concluded that four subspecies (brewsteri, extimus, adastus, and traillii) are recognizable. Paxton (2000) concluded that E. t. extimus is genetically distinct from other subspecies. Sedgwick (2001) demonstrated that E. t. adastus and E. t. extimus each have distinctive songs and used vocal signatures to determine distributional limits of the two subspecies. The two song types seem to be largely allopatric, separated by latitude and/or elevation. The two groups appear to be evolving independently of one another and warrant at least subspecific status.
Conservation Status
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NatureServe Status

Global Status: G5
Global Status Last Reviewed: 07Apr2016
Global Status Last Changed: 02Dec1996
Ranking Methodology Used: Ranked by inspection
Rounded Global Status: G5 - Secure
Reasons: Stable or increasing over much of the large range; has declined in the Southwest (see files for subspecies EXTIMUS) and in the Pacific states and British Columbia.
Nation: United States
National Status: N5B (19Mar1997)
Nation: Canada
National Status: N5B,N5M (26Jan2018)

U.S. & Canada State/Province Status
Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
United States Alabama (S1B), Arizona (S1), Arkansas (S1B,S3N), California (S1S2), Colorado (S4), Connecticut (S5B), Delaware (S3B), Florida (SNA), Georgia (S3), Idaho (S4B), Illinois (S5), Indiana (S4B), Iowa (S4B,S4N), Kansas (S2?B), Kentucky (S3S4B), Louisiana (SNA), Maine (S3?B), Maryland (S4B), Massachusetts (S4B), Michigan (S5), Minnesota (SNRB), Mississippi (SNA), Missouri (S3?), Montana (S4B), Navajo Nation (S1B), Nebraska (S4), Nevada (S3B), New Hampshire (S5B), New Jersey (S4B), New Mexico (S4N), New York (S5B), North Carolina (S3B), North Dakota (SNRB), Ohio (S5), Oklahoma (S4N), Oregon (S3B), Pennsylvania (S5B), Rhode Island (S3B,S3N), South Carolina (S4), South Dakota (S5B), Tennessee (S2S3), Texas (S1B), Utah (S4B), Vermont (S4B), Virginia (S4B), Washington (S4B), West Virginia (S4B), Wisconsin (S4B), Wyoming (S4B)
Canada Alberta (S3S4B), British Columbia (S5B), Manitoba (S2S3B), New Brunswick (S1S2B,S1S2M), Nova Scotia (S2B), Ontario (S5B), Quebec (S4B), Saskatchewan (S4B,S4M)

Other Statuses

Implied Status under the U.S. Endangered Species Act (USESA): PS
Comments on USESA: Subspecies extimus (southwestern willow flycatcher) of AZ, CA, CO, NM, NV, TX, UT, and Central America is listed by USFWS as Endangered (USFWS 1995). Listed as a state endangered species in California. Considered a species of high management concern in the Interior Columbia River Basin region (Saab and Rich 1997).
IUCN Red List Category: LC - Least concern

NatureServe Global Conservation Status Factors

Range Extent: >2,500,000 square km (greater than 1,000,000 square miles)
Range Extent Comments: BREEDING: central British Columbia across southern Canada and northern U.S. to New Brunswick, Prince Edward Island, and Nova Scotia, south to southern California, northern Baja California, northern Sonora, southern Arizona and New Mexico, western and central Texas, Arkansas, northern Georgia, and eastern Virginia (AOU 1998). NON-BREEDING: Nayarit and southwestern Oaxaca south to Panama and northwestern Colombia (Stiles and Skutch 1989, AOU 1998).

Subspecies brewsteri: Breeding distribution is west of the Cascades and in the Sierra Nevada from southwestern California to southwestern British Columbia (Sedgwick 2000).

Overall Threat Impact Comments: Threats include factors that destroy or degrade shrubby riparian vegetation. Riparian areas are particularly vulnerable to high-intensity livestock grazing, recreation and development pressure, flooding of nesting sites, and water diversions and flood control that prevent shrub and tree regeneration (Ohmart 1996, Saab and Rich 1997). HABITAT LOSS AND FRAGMENTATION: Habitat loss and alteration is thought to be the principle cause of decline in the West. Riparian habitats, particularly cottonwood and willow (SALIX spp.) communities, have been dramatically reduced and degraded by urban development, roads, off-road vehicle use, recreation, livestock grazing, agriculture, water development projects, channelization, willow control, and encroachment by non-native species (USFWS 1996). In California, high quality habitat has nearly disappeared and remaining habitat is widely dispersed and isolated, mostly as montane meadows in the Sierra Nevada and along a few river courses (USDA Forest Service 1994). In Arizona, as much as 90 percent of lowland riparian habitat has been lost or altered (USFWS 1996). These patterns are continuing throughout the western states. NON-NATIVE VEGETATION: Tamarisk or saltcedar (TAMARIX spp.) and Russian olive (ELAEAGNUS ANGUSTIFOLIA) have invaded riparian areas throughout the West, and particularly the southwest, and are likely factors in the flycatcher's decline. Tamarisk has replaced some riparian communities completely, but is less common in others. Tamarisk replaces the preferred multi-layered shrub community with a monotypic stand with one shrub layer, decreases plant and insect diversity, and can increase the frequency and intensity of fire. Dams and flood control, and irrigation water high in salts also give tamarisk a competitive edge over native vegetation. Although may nest in tamarisk where it provides the right vegetation structure, tamarisk may provide poor quality habitat and some studies have documented low breeding densities and low reproductive success in tamarisk (USFWS 1995, 1996; Sogge et al. 1997). LIVESTOCK: Placement of nests near the edges of shrubs makes them vulnerable to direct disturbance or destruction by livestock (Sanders and Flett 1989). Cattle and sheep browse on shrubs in the mid-level preferred by willow flycatchers, and consume or trample young woody plants (Sanders and Flett 1989). Heavy or poorly timed livestock grazing damages deciduous shrubs and can prevent shrub regeneration, reducing flycatcher habitat. Cattle prefer willow and cottonwood shoots to tamarisk and other non-natives, further depleting flycatcher habitat. Streambank trampling and soil compaction also adversely affect the water table, reduce free water, and discourage shrub growth (Flett and Sanders 1987). In Oregon, populations increased after reduction in cattle grazing and cessation of poisoning and removal of riparian willows (Taylor and Littlefield 1986). BROOD PARASITISM: Is a common host to brood parasitism by the brown-headed cowbird (MOLOTHRUS ATER; Bent 1942, King 1955, Walkinshaw 1966, Flett and Sanders 1987, Ehrlich et al. 1988, Sedgwick and Knopf 1988, Sanders and Flett 1989). Brood parasitism may pose a significant threat, particularly in its western range where habitat is limited and fragmented, and where livestock are often present in meadows and riparian habitats. In California, parasitism rates are high in the lowlands (e.g., 13 of 19 nests; Harris 1991), but much less in the Sierra Nevada (e.g., 1 of 22 nests; Sanders and Flett 1989). High parasitism rates (11 of 27 nests) were recorded at high elevations in northcentral Colorado, but cowbird eggs were accepted by the hosts at only 2 of these nests (Sedgwick and Knopf 1988). In Arizona, cowbirds were documented at all 12 known breeding locations and parasitism documented at 50 percent of the sites (USFWS 1996). In the Grand Canyon, cowbirds occur at all flycatcher breeding sites and typically 50 percent of nests are parasitized (Brown 1994, Sogge et al. 1997). In hundreds of monitored nests of southwestern willow flycatcher, brood parasitism caused either nest failure or successful rearing of only cowbird chicks (Sogge et al. 1997). There is, however, evidence of adaptive behavior toward cowbird parasitism in some populations. Parasitized nests may be abandoned or dismantled; renesting may occur in some cases (Sedgwick and Knopf 1988, Harris 1991), although fewer eggs may be laid (Holcomb 1974). Flycatchers will chase female cowbirds near nests, and some may respond to cowbird calls by becoming quieter and less conspicuous (Uyehara and Narins 1995). WATER DEVELOPMENTS: Impoundments, channelization, and water diversions have greatly reduced and modified native riparian habitats. Reservoirs flood native riparian communities, and regulated flows reduce the seasonal flooding that many riparian plants need for regeneration. Non-native plants (such as tamarisk in the southwest) often invade new habitat created along reservoir shorelines, or riparian communities of rivers with controlled or reduced flows from water diversion and dams. Channelization also alters the river system dynamics needed to maintain riparian communities (USFWS 1996). Some remaining populations in Arizona are threatened by fluctuating reservoir levels and flooding (Latta et al. 1999).

Subspecies brewsteri: While there appear to be moderate populations in early-seral upland forest habitats the species is especially threatened by population declines in valley habitats, lower nest success in these valley habitats than in early-seral forest, and continual loss of riparian habitat (Altman 2003).

Short-term Trend Comments: North American Breeding Bird Survey (BBS) data indicate a significant survey-wide population decline between 1966 and 1996 (-1.2 percent average annual decrease; P = 0.01; N = 1053 survey routes), and in the period from 1980 to 1996 (average -1.4 percent per year; P = 0.00; N = 957; Sauer et al. 1997). Significant declines evident in the western region for 1966-1996 (average -2.3 percent per year; P = 0.00; N = 311) and for 1980-1996 (average -1.9 percent per year; P = 0.00; N = 286); a likely decline indicated in the eastern region from 1980-1996 (average -0.9 percent per year; P = 0.19; N = 568). Most recent analysis for Canada indicates a stable overall trend from 1967 to 1998 (average 0.4 percent increase per year; not significant; N = 121 survey routes; Canadian Wildlife Service 1999). Mapped long-term trends for 1966-1996 show strong declines throughout the western range, from British Columbia to the Southwest, and a mosaic of declines and increases in the central and eastern states and provinces. Fewer than 500 breeding pairs of E. T. EXTIMUS are estimated to remain in widely scattered and isolated breeding sites (Arizona Partners in Flight 1999). In California, was once common and widespread but has been extirpated from most of it's range in the state; remaining populations occur in isolated areas in the Sierra Nevada mountains and along several rivers (Harris et al. 1987; USDA Forest Service 1994; USFWS 1996). In Arizona in 1997, EXTIMUS bred only in 45 sites statewide (Latta et al. 1999). Once common along the Colorado River, it is now very rare in the few undammed miles of Glen Canyon and in the Grand Canyon, and no longer breeds in the Lower Colorado River Valley (Carothers and Brown 1991; Rosenberg et al. 1991; Sogge 1995). The Grand Canyon population is dynamic, localized, and small, and only one to two sites are occupied in a given year (Sogge et al. 1997).

Long-term Trend:  
Long-term Trend Comments: Subspecies brewsteri: Once common the whole length of the Pacific Coast. Now rare to local in California mountain meadows. Only about 140-150 pairs in the Sierra Nevada. BBS data for the last 30+ years indicate significant population declines of 3.6%/yr. in western Oregon and Washington and northwestern California. In the Willamette Valley of western Oregon, populations have declined from a mean of 11-13 birds/route (BBS) in the early 1970s to 4-5 birds/route throughout the 1990s (Altman 2003).

Other NatureServe Conservation Status Information

Distribution
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Global Range: (>2,500,000 square km (greater than 1,000,000 square miles)) BREEDING: central British Columbia across southern Canada and northern U.S. to New Brunswick, Prince Edward Island, and Nova Scotia, south to southern California, northern Baja California, northern Sonora, southern Arizona and New Mexico, western and central Texas, Arkansas, northern Georgia, and eastern Virginia (AOU 1998). NON-BREEDING: Nayarit and southwestern Oaxaca south to Panama and northwestern Colombia (Stiles and Skutch 1989, AOU 1998).

Subspecies brewsteri: Breeding distribution is west of the Cascades and in the Sierra Nevada from southwestern California to southwestern British Columbia (Sedgwick 2000).

U.S. States and Canadian Provinces

Due to latency between updates made in state, provincial or other NatureServe Network databases and when they appear on NatureServe Explorer, for state or provincial information you may wish to contact the data steward in your jurisdiction to obtain the most current data. Please refer to our Distribution Data Sources to find contact information for your jurisdiction.
Color legend for Distribution Map
NOTE: The maps for birds represent the breeding status by state and province. In some jurisdictions, the subnational statuses for common species have not been assessed and the status is shown as not-assessed (SNR). In some jurisdictions, the subnational status refers to the status as a non-breeder; these errors will be corrected in future versions of these maps. A species is not shown in a jurisdiction if it is not known to breed in the jurisdiction or if it occurs only accidentally or casually in the jurisdiction. Thus, the species may occur in a jurisdiction as a seasonal non-breeding resident or as a migratory transient but this will not be indicated on these maps. See other maps on this web site that depict the Western Hemisphere ranges of these species at all seasons of the year.
Endemism: occurs (regularly, as a native taxon) in multiple nations

U.S. & Canada State/Province Distribution
United States AL, AR, AZ, CA, CO, CT, DE, FL, GA, IA, ID, IL, IN, KS, KY, LA, MA, MD, ME, MI, MN, MO, MS, MT, NC, ND, NE, NH, NJ, NM, NN, NV, NY, OH, OK, OR, PA, RI, SC, SD, TN, TX, UT, VA, VT, WA, WI, WV, WY
Canada AB, BC, MB, NB, NS, ON, QC, SK

Range Map
Note: Range depicted for New World only. The scale of the maps may cause narrow coastal ranges or ranges on small islands not to appear. Not all vagrant or small disjunct occurrences are depicted. For migratory birds, some individuals occur outside of the passage migrant range depicted. For information on how to obtain shapefiles of species ranges see our Species Mapping pages at www.natureserve.org/conservation-tools/data-maps-tools.

Range Map Compilers: NatureServe, 2002; WILDSPACETM 2002


U.S. Distribution by County Help
State County Name (FIPS Code)
AR Benton (05007), Boone (05009), Prairie (05117), Van Buren (05141)
AZ Apache (04001), Cochise (04003), Coconino (04005), Gila (04007), Graham (04009), Greenlee (04011), La Paz (04012), Maricopa (04013), Mohave (04015), Pima (04019), Pinal (04021), Santa Cruz (04023), Yavapai (04025), Yuma (04027)
CA Alpine (06003), Butte (06007), Del Norte (06015), El Dorado (06017), Fresno (06019), Humboldt (06023), Imperial (06025), Inyo (06027), Kern (06029), Lassen (06035), Los Angeles (06037), Madera (06039), Mariposa (06043), Mono (06051), Nevada (06057), Orange (06059), Placer (06061), Plumas (06063), Riverside (06065), San Bernardino (06071), San Diego (06073), Santa Barbara (06083), Shasta (06089), Sierra (06091), Siskiyou (06093), Tehama (06103), Tulare (06107), Tuolumne (06109), Ventura (06111)
CO Alamosa (08003), Dolores (08033), Gunnison (08051), Hinsdale (08053)
GA Glynn (13127)
NM Bernalillo (35001), Catron (35003), Chaves (35005)*, Cibola (35006), Dona Ana (35013), Grant (35017), Hidalgo (35023), Los Alamos (35028), Mckinley (35031), Mora (35033), Otero (35035), Rio Arriba (35039), San Juan (35045), San Miguel (35047), Sandoval (35043), Santa Fe (35049), Sierra (35051), Socorro (35053), Taos (35055), Valencia (35061)
NV Clark (32003), Lincoln (32017), Nye (32023)
UT Beaver (49001)*, Carbon (49007), Emery (49015), Garfield (49017), Grand (49019), Iron (49021)*, Kane (49025), San Juan (49037), Sevier (49041), Uintah (49047), Washington (49053), Wayne (49055)
WY Albany (56001), Big Horn (56003), Campbell (56005), Carbon (56007), Converse (56009), Crook (56011), Fremont (56013), Hot Springs (56017), Johnson (56019), Laramie (56021), Lincoln (56023), Natrona (56025), Niobrara (56027), Park (56029), Sheridan (56033), Sublette (56035), Sweetwater (56037), Teton (56039), Uinta (56041), Weston (56045)
* Extirpated/possibly extirpated
U.S. Distribution by Watershed Help
Watershed Region Help Watershed Name (Watershed Code)
03 Cumberland-St. Simons (03070203)+
08 Lower White (08020303)+
10 Yellowstone Headwaters (10070001)+, Clarks Fork Yellowstone (10070006)+, Upper Wind (10080001)+, Popo Agie (10080003)+, Lower Wind (10080005)+, Badwater (10080006)+, Upper Bighorn (10080007)+, Nowood (10080008)+, Greybull (10080009)+, Big Horn Lake (10080010)+, North Fork Shoshone (10080012)+, South Fork Shoshone (10080013)+, Shoshone (10080014)+, Little Bighorn (10080016)+, Upper Tongue (10090101)+, Middle Fork Powder (10090201)+, Upper Powder (10090202)+, Crazy Woman (10090205)+, Clear (10090206)+, Little Powder (10090208)+, Antelope (10120101)+, Dry Fork Cheyenne (10120102)+, Upper Cheyenne (10120103)+, Lance (10120104)+, Lightning (10120105)+, Angostura Reservoir (10120106)+, Beaver (10120107)+, Hat (10120108)+, Upper Belle Fourche (10120201)+, Redwater (10120203)+, Niobrara Headwaters (10150002)+, Upper North Platte (10180002)+, Medicine Bow (10180004)+, Sweetwater (10180006)+, Middle North Platte-Casper (10180007)+, Glendo Reservoir (10180008)+, Upper Laramie (10180010)+, Cache La Poudre (10190007)+, Lone Tree-Owl (10190008)+, Crow (10190009)+, Upper Lodgepole (10190015)+
11 Bull Shoals Lake (11010003)+, Little Red (11010014)+, Mora (11080004)+, Illinois (11110103)+
13 Alamosa-Trinchera (13010002)+, Upper Rio Grande (13020101)+, Rio Chama (13020102)+, Rio Grande-Santa Fe (13020201)+, Jemez (13020202)+, Rio Grande-Albuquerque (13020203)+, Rio Puerco (13020204)+, Rio San Jose (13020207)+, Rio Salado (13020209)+, Elephant Butte Reservoir (13020211)+, Caballo (13030101)+, El Paso-Las Cruces (13030102)+, Tularosa Valley (13050003)+, Pecos headwaters (13060001)+, Upper Pecos-Long Arroyo (13060007)+*, Rio Penasco (13060010)+*
14 Tomichi (14020003)+, Westwater Canyon (14030001)+, Upper Dolores (14030002)+, Lower Dolores (14030004)+, Upper Colorado-Kane Springs (14030005)+, Upper Green (14040101)+, New Fork (14040102)+, Upper Green-Slate (14040103)+, Bitter (14040105)+*, Upper Green-Flaming Gorge Reservoir (14040106)+, Blacks Fork (14040107)+, Muddy (14040108)+, Great Divide closed basin (14040200)+, Little Snake (14050003)+, Muddy (14050004)+, Lower Green-Desolation Canyon (14060005)+, Price (14060007)+, Lower Green (14060008)+, Upper Lake Powell (14070001)+*, Fremont (14070003)+, Dirty Devil (14070004)+, Escalante (14070005)+, Lower Lake Powell (14070006)+, Paria (14070007)+, Piedra (14080102)+, Middle San Juan (14080105)+, Chaco (14080106)+, Lower San Juan-Four Corners (14080201)+, Montezuma (14080203)+, Lower San Juan (14080205)+*
15 Lower Colorado-Marble Canyon (15010001)+, Grand Canyon (15010002)+, Lake Mead (15010005)+, Grand Wash (15010006)+, Upper Virgin (15010008)+, Fort Pierce Wash (15010009)+, Lower Virgin (15010010)+, White (15010011)+, Muddy (15010012)+, Meadow Valley Wash (15010013)+, Las Vegas Wash (15010015)+, Little Colorado headwaters (15020001)+, Zuni (15020004)+, Havasu-Mohave Lakes (15030101)+, Imperial Reservoir (15030104)+, Lower Colorado (15030107)+, Big Sandy (15030201)+, Santa Maria (15030203)+, Bill Williams (15030204)+, Upper Gila (15040001)+, Upper Gila-Mangas (15040002)+, San Francisco (15040004)+, Upper Gila-San Carlos Reservoir (15040005)+, San Simon (15040006)+, Middle Gila (15050100)+, Upper San Pedro (15050202)+, Lower San Pedro (15050203)+, Upper Santa Cruz (15050301)+, Rillito (15050302)+, Upper Salt (15060103)+, Tonto (15060105)+, Lower Salt (15060106)+, Upper Verde (15060202)+, Lower Verde (15060203)+, Lower Gila-Painted Rock Reservoir (15070101)+, Agua Fria (15070102)+, Hassayampa (15070103)+, Lower Gila (15070201)+
16 Central Bear (16010102)+, Upper Sevier (16030001)+, East Fork Sevier (16030002)+*, Escalante Desert (16030006)+*, Beaver Bottoms-Upper Beaver (16030007)+*, Lake Tahoe (16050101)+, Truckee (16050102)+, Upper Carson (16050201)+, Sand Spring-Tikaboo Valleys (16060014)+
17 Snake headwaters (17040101)+, Gros Ventre (17040102)+, Greys-Hobock (17040103)+, Palisades (17040104)+, Salt (17040105)+, Lower Henrys (17040203)+, Teton (17040204)+
18 Smith (18010101)+, South Fork Eel (18010106)+, Upper Klamath (18010206)+, Shasta (18010207)+, Upper Pit (18020002)+, Lower Pit (18020003)+, Mccloud (18020004)+, North Fork Feather (18020121)+, East Branch North Fork Feather (18020122)+, Middle Fork Feather (18020123)+, Upper Yuba (18020125)+, South Fork American (18020129)+*, Battle Creek (18020153)+, Thomes Creek-Sacramento River (18020156)+, Big Chico Creek-Sacramento River (18020157)+, South Fork Kern (18030002)+, Middle Kern-Upper Tehachapi- (18030003)+, Upper Kaweah (18030007)+, Upper King (18030010)+, Upper San Joaquin (18040006)+, Upper Merced (18040008)+, Upper Tuolumne (18040009)+, Upper Stanislaus (18040010)+*, Santa Ynez (18060010)+, Santa Clara (18070102)+, Santa Monica Bay (18070104)+*, Los Angeles (18070105)+*, San Gabriel (18070106)+, San Jacinto (18070202)+, Santa Ana (18070203)+, Aliso-San Onofre (18070301)+, Santa Margarita (18070302)+, San Luis Rey-Escondido (18070303)+, San Diego (18070304)+, Mono Lake (18090101)+, Crowley Lake (18090102)+, Owens Lake (18090103)+*, Upper Amargosa (18090202)+, Mojave (18090208)+, Whitewater River (18100201)+, Salton Sea (18100204)+
+ Natural heritage record(s) exist for this watershed
* Extirpated/possibly extirpated
Ecology & Life History
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Basic Description: Small flycatcher; 13.3-17 cm in length (Sedgwick 2000). Drab olive becoming brownish gray (with wear) on back. Underside is light gray; belly washed with yellow in the spring. Wings with whitish-gray wing-bars; underwing-coverts white. Indistinct (sometimes lacking) whitish eye-ring. Juveniles are browner above and yellower below, with buff or yellowish brown wing-bars.

General Description: A flycatcher with brownish-olive upperparts (slightly grayer in the east), a whitish throat that contrasts with the pale olive breast, a pale yellow belly, and two light wing bars; generally lacks a conspicuous eye ring; as in other flycatchers, the bill is depressed and wide at the base (NGS 1983).
Diagnostic Characteristics: Generally indistinguishable from the Alder Flycatcher (E. ALNORUM), but tends to lack a conspicuous eye ring (Alder tends to have one), have a slightly longer bill, and is less green above (NGS 1983). Reliably distinguished from the Alder Flycatcher only by voice. Song is a sneezy "fitz-bew," with accent on the first syllable (Alder Flycatcher song is "rrree-BEEa" or "fee-bee-o" with accent on the second syllable) (Kaufman 1990, McCabe 1991). Breeding habitats of the two species differ somewhat, with Willow Flycatcher in more southern and western regions of North America and in more open habitats and Alder Flycatcher a more northern bird, generally breeding in shrub and alder thickets of boreal forests in the eastern U.S., Canada, and Alaska (McCabe 1991).
Reproduction Comments: A late breeder, eggs usually laid in mid- to late-June and young fledge in August (USDA Forest Service 1994). Clutch size is three to four. Incubation lasts 12-15 days, by female. Young are tended by both parents, leave nest at 12-15 days. Site fidelity strong in both males and females (Walkinshaw 1966). May incur a high rate of cowbird parasitism (e.g., Sedgwick and Knopf 1988, Harris 1991, Brown 1988). Sometimes polygynous and may maintain polygynous trios, possibly a response to narrow habitats with high habitat productivity or other factors (Prescott 1986, Sedgwick and Knopf 1989). Singing, unmated males may be present on breeding grounds, and single pairs may breed in absence of other individuals (USDA Forest Service 1994). On one study in Ohio and Nebraska, 91 nests had 272 eggs from which 99 young fledged (36.4 percent success) and 39.5 percent of nests produced at least one young; 96 eggs and 41 nestlings were depredated (50.4 percent; Holcomb 1972).
Ecology Comments: BREEDING: Conduct most of their activity within their defended territory, but both male and female will also use adjacent areas, especially when feeding young; territory defense declines once young are fledged (USDA Forest Service 1994). In Ontario, territory size ranged from about 0.1 hectares to 0.47 hectares and averaged 0.35 hectares (Prescott and Middleton 1988); in southern Michigan, territories averaged 0.7 hectares (Walkinshaw 1966). In California, territories ranged from 0.1 hectares to 0.9 hectares, and averaged 0.2 hectares in Fresno County and 0.4 hectares on the Truckee River (USDA Forest Service 1994). Where breeding range overlaps with alder flycatchers (EMPIDONAX ALNORUM), may show territorial defense toward the other species (Prescott 1987).

NON-BREEDING: In Panama, winter home range estimated to be about 1100 square meters (Gorski 1969).

Non-Migrant: N
Locally Migrant: N
Long Distance Migrant: Y
Mobility and Migration Comments: Usually arrives on U.S. nesting grounds by May-June (Terres 1980). Present in California from late April to September (Biosystems Analysis 1989). Migrates through southern Arizona mainly in the first half of June and August-September (Phillips et al. 1964). Arrives in Washington in late May or early June. Fairly common migrant in Costa Rica, mid-August to late October (peak late September) and mid-March to late May (Stiles and Skutch 1989). In Ontario, tends to have an earlier and longer migration period in spring than Alder flycatcher (EMPIDONAX ALNORUM), and migrates significantly earlier in fall (Hussell 1991a, 1991b).
Palustrine Habitat(s): FORESTED WETLAND, Riparian
Terrestrial Habitat(s): Old field, Shrubland/chaparral, Woodland - Hardwood, Woodland - Mixed
Habitat Comments: BREEDING: Strongly tied to brushy areas of willow (SALIX spp.) and similar shrubs. Found in thickets, open second growth with brush, swamps, wetlands, streamsides, and open woodland (AOU 1983). Common in mountain meadows and along streams; also in brushy upland pastures (especially hawthorn) and orchards (NGS 1983). The presence of water (running water, pools, or saturated soils) and willow, alder (ALNUS spp), or other deciduous riparian shrubs are essential habitat elements (Sanders and Flett 1989, USDA Forest Service 1994). Occurs in both mesic and drier upland conditions, but apparently reaches highest densities on wet sites (Sedgwick and Knopf 1992). It is associated with dense riparian deciduous shrub cover separated by open areas, but large contiguous willow thickets without openings are typically avoided; it does not occur in dense tree cover but will use scattered trees for song and foraging perches and gleaning substrate (USDA Forest Service 1994). Habitat preferences may overlap with alder (EMPIDONAX ALNORUM) and least flycatchers (EMPIDONAX MINIMUS), to include deciduous woods and thickets, bottomlands and swamps (Griggs 1997). Foraging habitat may overlap with western flycatcher (EMPIDONAX DIFFICILIS; Frakes and Johnson 1982).

In southwestern Ontario, generally occurs in more xeric upland sites, but in some areas uses boggy alder thickets, overlapping with alder flycatcher (Barlow and McGillivray 1983). In the Sierra Nevada of California, broad, flat meadows with willows and water are essential (Sanders and Flett 1989). In the Northern Rockies, is apparently restricted to riparian areas with adequate shrub cover (Hutto and Young 1999).

In Colorado, males and females were found to select for different habitat attributes: female-selected nest sites typically had dense willows and were similar in patch size and bush height, male-selected song perch sites were characterized by large central shrubs and high variability in shrub size. On an increasing scale, breeding sites were respectively characterized by greater willow density, larger willow patches with smaller gaps, and greater percent willow coverage than non-willow coverage (Sedgwick and Knopf 1992).

Southwestern willow flycatcher (E. T. EXTIMUS) breeds only in dense riparian vegetation near water or saturated soil. Habitat typically contains dense vegetation in the patch interior, often interspersed with small openings, sparser vegetation, or open water that creates a habitat mosaic of variable density. It nests in shrub and tree thickets 4-7 meters tall, with dense foliage 0-4 meters above the ground, and usually a high canopy coverage (USFWS 1995). The dominant plant species, size and shape of habitat patch, canopy structure and other habitat variables vary from monotypic to mixed-species stands and from simple to complex vegetation structures (Sogge et al. 1997). Habitats include dense high-elevation willow; native broadleaf shrubs and trees composed of willow, cottonwood (POPULUS spp.), boxelder (ACER NEGUNDO), ash (FRAXINUS spp.), alder, or buttonbush (CEPHALANTHUS OCCIDENTALIS); monotypic closed-canopy stands of tamarisk (TAMARIX spp.) or Russian olive (ELAEAGNUS ANGUSTIFOLIA); or a mix of native shrubs and exotic species (Sogge et al. 1997). Along the Virgin River, Utah, is restricted to shrub communities with shrub densities ranging from 70 percent to 100 percent (Whitmore 1977).

NEST SITE: Nests primarily near slow streams, standing water or seeps, swampy thickets, especially of willow and buttonbush (AOU 1983, USDA Forest Service 1994), also dogwood (CORNUS spp.), elderberry, hawthorn, rose, tamarisk, and others; in fork or on horizontal limb of shrub, usually 1-3 meters above ground (see Harris 1991). In montane habitats, nests are usually in willows at least 2 meters high with foliage density of 50-70 percent and about 1 meter of cover above the nest (Sanders and Flett 1989). Also see Sedgwick and Knopf (1992) for information on nest sites and song perches in northcentral Colorado.

Historically, southwestern willow flycatcher primarily in willows, buttonbush, and BACCHARIS spp. with a scattered cottonwood overstory. With changes in riparian plant communities, non-native tamarisk and Russian olive provide nesting habitat in some areas (Brown 1988, USFWS 1995). Along the Colorado in the Grand Canyon, for example, the flycatcher nests in tall tamarisk within 30 meters of water (Brown 1988, Sogge et al. 1997); however it is not known if nesting success differs in tamarisk compared to native vegetation (USFWS 1996). Where E. T. EXTIMUS nests in tamarisk, the tamarisk are usually taller (more than 5 meters) and denser (90 percent canopy closure) than in tamarisk-dominated areas where the flycatcher has been extirpated, and broadleaf shrubs may also be an important part of the community (Sogge et al. 1997).

NON-BREEDING: Uses same types of habitats during migration and winter as breeding season (McCabe 1991). Occurs in dense scrub, deciduous broadleaf forest, streamside gallery forest, and freshwater wetlands (Rappole et al. 1995). In western Mexico and Central America, found in humid to semi-arid scrubby fields with hedges, fences woodland and edge, plantations; frequents low to mid-vegetation levels and often comes into open (Howell and Webb 1995).

Adult Food Habits: Invertivore
Immature Food Habits: Invertivore
Food Comments: Eats mainly insects caught in flight; occasionally berries. Bent (1942) states that 96 percent of diet is animal matter, most of which is flying insects. Once fledglings are able to forage for themselves, are less dependent on a localized, concentrated food source (USDA Forest Service 1994).
Adult Phenology: Diurnal
Immature Phenology: Diurnal
Length: 15 centimeters
Weight: 14 grams
Economic Attributes Not yet assessed
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Management Summary
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Stewardship Overview: Found in shrubby deciduous habitats, especially riparian areas and meadows with shrubby patches dominated by willows (SALIX spp.) or alder (ALNUS spp.). Has undergone dramatic population declines in the past 30 years, particularly throughout its western range where its preferred riparian and willow habitats have been lost and degraded on a broad scale. The bulk of recent research and management focus on the species has been in California and the Southwest where populations are critically low. Much further study of habitat requirements and response to land management activities and restoration is needed.
Restoration Potential: Has become so rare in the southwest and California that extirpation from many remaining breeding areas seems likely without dramatic management efforts and habitat restoration. Elsewhere in the West, riparian and meadow restoration is needed to sustain the species and reverse declining trends. Fortunately, riparian and willow habitats are resilient where natural hydrologic dynamics and native vegetation are restored. Where cowbirds are affecting population viability, measures to reduce cowbird presence during the breeding season will be needed. Less is known about the restoration needs in the eastern and northwestern portions of its range, but these populations currently seem somewhat more abundant and stable.
Preserve Selection & Design Considerations: Minimum area requirements and patch dynamics are still largely unknown. Small openings in deciduous shrub habitats or adjacent stream edges increase habitat suitability, and large, contiguous willow patches apparently do not support willow flycatchers at the interior of the patch (USDA Forest Service 1994). In California montane meadows, the smallest documented nesting area is a 0.25 hectare meadow in the Sierra Nevada, but other observers found most nesting territories in meadows greater than 8.0 hectares and none in meadows less than 0.4 hectares (USDA Forest Service 1994). For southwestern willow flycatcher (E. T. EXTIMUS), patches as small as 0.5 hectares have been found to support one to two pairs, and habitat patches range from 0.5 to 1.2 hectares (USFWS 1995).
Management Requirements: HABITAT MANAGEMENT: Will benefit from maintaining communities of deciduous shrubs in riparian areas and meadows, with patches of dense shrubs interspersed with openings, and with open water nearby. In areas where populations have declined, existing breeding areas should be protected from habitat loss. It should be noted that unoccupied sites are not necessarily unsuitable, as populations may be dynamic (Harris et al. 1987).

In the Sierra Nevada, needs riparian areas and wet meadows at least 0.25 hectares in size with openings and large, dense patches of deciduous shrubs, and dense foliage at mid-heights (1-2 meters); habitat areas of at least 8 hectares are optimal. Territories contained 5-80 percent willow (SALIX spp.) cover (average 44 percent), 18-78 percent (average 54 percent) foliage density in the 0-1 meter shrub layer, and 45-96 percent (average 69 percent) foliage density in the 1-2 meter shrub layer (Sanders and Flett 1989).

Management recommendations for montane meadows include: (1) maintain riparian deciduous shrubs at least 1-2 meters high in patches greater than 0.1 hectares (100 square meters); (2) maintain more than 40 percent foliage cover density in lower 2 meters of the deciduous shrub layer; (3) maintain shrub patches interspersed with openings, and opening or paths should be at least 2 meters wide to allow aerial foraging (USDA Forest Service 1994).

Critical habitat areas containing the remaining known populations of southwestern willow flycatcher (E. T. EXTIMUS) were identified by the U.S. Fish and Wildlife Service in Southern California, Arizona, and New Mexico (USFWS 1997a, 1997b). In Arizona, habitat recommendations include: (1) establish a "no net loss" policy; increasing suitable riparian habitat and promoting regeneration of native plants; (2) using buffer zones between riparian habitats and adjacent developments; (3) restore connectivity of natural reaches of habitat by restoring degraded segments; (4) establish areas of slow water and backwater; (4) manage for large contiguous habitat blocks rather than isolated fragments (Latta et al. 1999). Also see Deshler et al. (1997) for an extensive bibliography on this subspecies.

WATER MANAGEMENT: The species requires saturated soils, standing water or flowing water near nesting sites. Natural flooding and channel meandering can promote native riparian vegetation communities. Maintaining wetlands and wet meadows will help sustain willow communities. Arizona recommendations include: managing water diversions and groundwater withdrawal to maintain streamside vegetation and mimic natural stream flow regimes, including periodic floods (Latta et al. 1999).

GRAZING: In Oregon, populations dramatically increased after reducing cattle grazing and ceasing the poisoning and removal of riparian willows (Taylor and Littlefield 1986). Reducing or eliminating grazing during the nestling period would reduce direct damage by livestock (USDA Forest Service 1994). Flycatchers will nest near cattle trails (Sanders and Flett 1989), and in some circumstances cattle might be used to create trails and openings in exceptionally dense willow stands (outside the breeding season) to benefit flycatchers. Livestock management recommendations include: (1) eliminate livestock use during the breeding period from early June to mid-August; (2) manage stocking rates and timing to encourage riparian shrub growth and vigor; (3) use exclosures to protect sensitive areas from browsing and soil compaction (USDA Forest Service 1994).

BROOD PARASITES: Eliminating livestock in breeding habitat during the breeding season can help alleviate cowbird brood parasitism. Cowbird trapping and control is intensive, expensive, and must be ongoing to be effective. However, control efforts on two sites in California in the early 1990s helped conserve isolated populations, stabilizing numbers at one site and increasing nest success at the other (USFWS 1996). Management recommendations to reduce the impacts of cowbirds include: (1) manage livestock to prevent aggregations near breeding habitat during nest building, egg-laying and incubation, late May to late July; (2) ideally, corrals, pack stations, and other facilities that concentrate stock should be 5 to 10 kilometers from important habitat areas, and at least 1 kilometer from nest sites; (3) implement cowbird trapping if necessary in coordination with state fish and wildlife agencies (USDA Forest Service 1994). In Arizona, recommendations include: reducing parasitism rates to less than 20 percent; monitoring nests; implementing trapping programs where parasitism rates exceed 20 percent (Latta et al. 1999).

Monitoring Requirements: Willow flycatcher is very difficult to distinguish from other EMPIDONAX species except by its distinctive "fitz-bew" call. Singing birds can be surveyed by standard point count, line transect, and similar survey methods based on vocalization. The species defends relatively small territories which can facilitate nest searches and monitoring. Males generally sing from highest perch available (USDA Forest Service 1994). Females may also vocalize, producing "pit," "weeoo," and "creet" calls as well as the "fitz-bew" and "fizz-bew" male advertising songs (Seutin 1987).

A number of factors can make surveys somewhat problematical. Habitat may be patchy and limited, decreasing detectability. Spring migrants may sing and be confused with local breeders, populations vary in related behavior such as song-flights and in singing frequency throughout the day, and there are apparently subtle dialectic differences in the song and calls across the species range (e.g., Wedemeyer 1973). In winter, the species rarely sings, although call notes may be used frequently and birds will respond territorially with the "fitz-bew" song and other calls in response to taped playbacks (Gorski 1969, McCabe 1991). A standardized survey protocol for southwestern willow flycatcher is presented by Sogge et al. (1997).

For mist-netting and banding, identifying EMPIDONAX species in hand is based on a suite of subtle morphological differences and requires trained handlers, yet willow and alder flycatchers cannot be reliably distinguished from one another in the hand (McKinney 1988, Seutin 1991). Stein (1963) presented an equation for identification of the two species based on bill length and wing formula. However, others have found that this method is not reliable in areas of sympatry (Seutin 1991) or for immature birds (Hussell 1990).

Management Research Needs: Details of habitat and management needs in the eastern extent of its range are largely unknown. Where populations are depleted in the West, annual population and productivity monitoring is needed as well as surveys to determine presence/absence, distribution, and habitat relationships. Information is needed on landscape relationships, minimum area requirements, patch dynamics; also effect of breeding habitat isolation and connectivity. Further study is needed of habitat preferences throughout the species range, particularly in relation to productivity, use of non-native vegetation, relationship to land management activities, and response of flycatchers to habitat restoration. More information is needed on parasitism rates, productivity of parasitized nests, response to parasitism, and activities and habitat that promote the incidence of cowbirds. Effects of pesticides on the species are unknown. Information on winter habitat use, winter ecology, and threats on the wintering grounds are almost entirely lacking.
Biological Research Needs: Many aspects of the taxonomy, biology and ecology remain unstudied. Diet composition, winter habits and ecology are unknown. Genetic studies of subspecies are needed. Migration patterns and the location of wintering sites for different subspecies remain unknown, largely due to the difficulty of distinguishing the species outside the breeding season. Demographics (productivity and survivorship) need further study, especially in relation to habitat quality and to landscape variables at different scales. Variation in songs and calls across the species range and among subspecies need study.
Population/Occurrence Delineation
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Group Name: Passerines

Use Class: Breeding
Subtype(s): Foraging Area, Nest Site, Nesting Colony
Minimum Criteria for an Occurrence: Evidence of historical breeding, or current and likely recurring breeding, at a given location, minimally a reliable observation of one or more breeding pairs in appropriate habitat. Be cautious about creating EOs for observations that may represent single breeding events outside the normal breeding distribution.

Mapping Guidance: Breeding occurrences include nesting areas as well as foraging areas.

For swallows and other species that have separate nesting and foraging areas, separations are based on nest sites or nesting areas, not to locations of foraging individuals. For example, nesting areas separated by a gap larger than the separation distance are different occurrences, regardless of the foraging locations of individuals from those nesting areas. This separation procedure is appropriate because nesting areas are the critical aspect of swallow breeding occurrences, tend to be relatively stable or at least somwhat predictable in general location, and so are the basis for effective conservation; foraging areas are much more flexible and not necessarily static.

Separation Distance for Unsuitable Habitat: 5 km
Separation Distance for Suitable Habitat: 5 km
Separation Justification: Significant dispersal and associated high potential for gene flow among populations of birds separated by tens of kilometers (e.g., Moore and Dolbeer 1989), and increasing evidence that individuals leave their usual home range to engage in extrapair copulations, as well as long foraging excursions of some species, make it difficult to circumscribe occurrences on the basis of meaningful population units without occurrences becoming too large. Hence, a moderate, standardized separation distance has been adopted for songbirds and flycatchers; it should yield occurrences that are not too spatially expansive while also accounting for the likelihood of gene flow among populations within a few kilometers of each other.

Be careful not to separate a population's nesting areas and foraging areas as different occurrences; include them in the same occurrence even if they are more than 5 km apart. Mean foraging radius (from nesting area) of Brown-headed Cowbird females was 4.0 kilometers in California, 1.2 kilometers in Illinois-Missouri (Thompson 1994). Yellow-headed Blackbirds, Brewer's Blackbirds, and probably Red-winged Blackbirds all forage up to 1.6 kilometers away from breeding colony (Willson 1966, Horn 1968). In one study, Brewer's Blackbirds were found as far as 10 kilometers from nesting area (Williams 1952), but this may be unusual.

For swallows and other parrerines with similar behavioral ecology, separation distance pertains to nest sites or nesting colonies, not to locations of foraging individuals. For example, nesting areas separated by a gap of more than 5 km are different occurrences, regardless of the foraging locations of individuals from those nesting areas. This separation procedure is appropriate because nesting areas are the critical aspect of swallow breeding occurrences, tend to be relatively stable or at least somwhat predictable in general location, and so are the basis for effective conservation; foraging areas are much more flexible and not necessarily static.

Be cautious about creating EOs for observations that may represent single breeding events outside the normal breeding distribution.

Unsuitable habitat: Habitat not normally used for breeding/feeding by a particular species. For example, unsuitable habitat for grassland and shrubland birds includes forest/woodland, urban/suburban, and aquatic habitats. Most habitats would be suitable for birds with versatile foraging habits (e.g., most corvids).

Date: 10Sep2004
Author: Hammerson, G.

Use Class: Migratory stopover
Subtype(s): Foraging Area, Roost Site
Minimum Criteria for an Occurrence: For most passerines: Evidence of recurring presence of migrating individuals (including historical) and potential recurring presence at a given location; minimally a reliable observation of 25 birds in appropriate habitat.

For swallows: Evidence of recurring presence of migrating flocks (including historical) and potential recurring presence at a given location; minimally a reliable observation of 100 birds in appropriate habitat (e.g., traditional roost sites).

Occurrences should be locations where the species is resident for some time during the appropriate season; it is preferable to have observations documenting presence over at least 7 days annually.

EOs should not be described for species that are nomadic during nonbreeding season: e.g., Lark Bunting.

Be cautious about creating EOs for observations that may represent single events.

Separation Distance for Unsuitable Habitat: 5 km
Separation Distance for Suitable Habitat: 5 km
Separation Justification: Separation distance somewhat arbitrary but intended to define occurrences of managable size for conservation purposes. Occurrences defined primarily on the basis of areas supporting concentrations of birds, rather than on the basis of distinct populations.

For swallows and other species with similar behavioral ecology, the separation distance pertains to communal roost sites rather than to foraging areas; the former tend to be more stable and specific over time than the latter.

Date: 03Sep2004
Author: Hammerson, G., and S. Cannings

Use Class: Nonbreeding
Subtype(s): Foraging Area, Roost Site
Minimum Criteria for an Occurrence: Any area used traditionally in the nonbreeding season (used for populations that are not resident in a location year-round). Minimally, reliable observations of 10 or more individuals in appropriate habitat for 20 or more days at a time. For G1-G3 species, observations of fewer individuals could constitute an occurrence of conservation value. Sites used during migration should be documented under the 'migratory stopover' location use class.

Separation Distance for Unsuitable Habitat: 5 km
Separation Distance for Suitable Habitat: 5 km
Separation Justification: Separation distance is necessarily arbitrary but attempts to balance the high mobility of birds with the need for occurrences of reasonable spatial scope. Note that a population's roost sites and foraging areas are parts of the same occurrence, even if they are more than 5 km apart.

For swallows and other species with similar behavioral ecology, the separation distance pertains to communal roost sites rather than to foraging areas; the former tend to be more stable and specific over time than the latter.

Date: 03Sep2004
Author: Hammerson, G.

Use Class: Nonmigratory
Minimum Criteria for an Occurrence: Occurrences are based on evidence of historical presence, or current and likely recurring presence, at a particular location. Such evidence minimally includes collection or reliable observation and documentation of one or more individuals in or near appropriate habitat.

These occurrence specifications are used for nonmigratory populations of passerine birds.

Separation Barriers: None.
Separation Distance for Unsuitable Habitat: 5 km
Separation Distance for Suitable Habitat: 5 km
Separation Justification: Significant dispersal and associated high potential for gene flow among populations of birds separated by tens of kilometers (e.g., Moore and Dolbeer 1989), and increasing evidence that individuals leave their usual home range to engage in extrapair copulations, as well as long foraging excursions of some species, make it difficult to circumscribe occurrences on the basis of meaningful population units without occurrences becoming too large. Hence, a moderate, standardized separation distance has been adopted for songbirds and flycatchers; it should yield occurrences that are not too spatially expansive while also accounting for the likelihood of gene flow among populations within a few kilometers of each other.

Be careful not to separate a population's nesting areas and breeding-season foraging areas as different occurrences; include them in the same occurrence even if they are more than 5 km apart. Blue jays have small summer home ranges but fly up to 4 kilometers to harvest mast (Tarvin and Woolfenden 1999). Flocks of pinyon jays range over 21-29 square kilometers (Ligon 1971, Balda and Bateman 1971); nesting and foraging areas may be widely separated. Tricolored blackbirds forage in flocks that range widely to more than 15 kilometers from the nesting colony (Beedy and Hamilton 1999).

Unsuitable habitat: Habitat not normally used for breeding/feeding by a particular species. For example, unsuitable habitat for grassland and shrubland birds includes forest/woodland, urban/suburban, and aquatic habitats. Most habitats would be suitable for birds with versatile foraging habits (e.g., most corvids).

Date: 10Sep2004
Author: Hammerson, G.
Notes: These specs pertain to nonmigratory species.
Population/Occurrence Viability
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U.S. Invasive Species Impact Rank (I-Rank) Not yet assessed
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Authors/Contributors
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NatureServe Conservation Status Factors Edition Date: 28Jan2005
Management Information Edition Date: 16Sep1999
Management Information Edition Author: PAIGE, C.; REVISIONS BY M. KOENEN AND D.W. MEHLMAN
Management Information Acknowledgments: Support for the preparation of this abstract was provided by the National Fish and Wildlife Foundation's Neotropical Migratory Bird Conservation Initiative, through challenge grant number 97-270 to The Nature Conservancy, Wings of the Americas Program. Matching funds for this grant were donated by Canon U.S.A., Inc.
Element Ecology & Life History Edition Date: 28Mar1995
Element Ecology & Life History Author(s): HAMMERSON, G., MINOR REVISIONS BY S. CANNINGS

Zoological data developed by NatureServe and its network of natural heritage programs (see Local Programs) and other contributors and cooperators (see Sources).

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