Northern Pintail

Anas acuta


Distribution, Migration, and Habitat

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Figure 1. Distribution of the Northern Pintail in North America.

This species also breeds in northern Europe and Asia.

eBird range map for Northern Pintail

Generated from eBird observations (Year-Round, 1900-present)

Distribution in the Americas

eBird data provide detailed looks at the range of this species throughout the year: eBird Year-round Range and Point Map for Northern Pintail.

Breeding Range

See Figure 1. Widespread. In w. North America breeds on se. Victoria I. and s. Southampton I., and from n. Alaska, n. Yukon, n. NW Territories and e. Manitoba south to central and sw. California, n. Nevada, central Utah, n. New Mexico, w. Kansas, n. Iowa, and w. Minnesota. Also breeds in Alaska west to Amchitka I., in central Arizona, and on s. Vancouver I., but absent from western portions of British Columbia, Washington, Oregon, and n. California.

In e. North America breeds along the coast of Hudson and James bays, in nw. Quebec, throughout e. Labrador and e. Quebec, south locally to ne. New York, s. Ontario, nw. Ohio, e. Michigan, and central Wisconsin, and on the n. Gaspé Peninsula, Newfoundland I., Anticosti I., Prince Edward I., e. New Brunswick, and w. Nova Scotia. Casual or sporadic in scattered localities in sw. Baffin I., n. New York and Massachusetts, south to Maryland, Virginia, and w. Kentucky (no records in these states 2000-2012), and west to n. Ohio and Indiana, n. Illinois, n. Iowa, Kansas, and s. New Mexico. In summer, casual to Banks, Ellesmere and Baffin I. (Bellrose 1980, Godfrey 1986).

Key portion of range is Alaska and the Prairie Pothole Region (Bellrose 1980); also nests extensively in w. U.S. Great Basin, California, and in n. Canada. Individuals on arctic coastal plain are largely nonbreeders (Derksen and Eldridge 1980).

Strong tendency to pioneer over wide areas, responding opportunistically to available wetlands (Hochbaum and Bossenmaier 1972). Responds initially to wetland conditions in southern parts of the breeding range and proceeds northward if conditions are not suitable, although many migrate directly to far northern habitats, bypassing prairie entirely (Johnson and Grier 1988, Miller et al. 2005). Particularly prone to displacement to northern areas when prairies are dry (Smith 1970d, Derksen and Eldridge 1980).

Recent distributional data for key parts of the breeding range, especially at the periphery, follow:

N. Alaska: common on coastal plain and south side of the Brooks Range ( See eBird data.

British Columbia: breeds throughout the interior of the province, east of the Coast Ranges and locally on the coast of s. Vancouver I., the Fraser Lowlands and Queen Charlotte Lowlands. From sea level to 1070 m (Campbell et al. 1990). See eBird data.

Yukon: widespread breeder, particularly in southwest and on the northern coastal plain (Sinclair et al. 2003). See eBird data.

Nevada: confined primarily to wetland complexes of the north (e.g., Ruby Lakes); occasionally common there (Floyd et al. 2007). See eBird data.

California: local breeder, most numerous on lakes of n.-central CA (Klamanth Basin), with small numbers breeding south to Imperial and San Diego counties (Small 1994). See eBird data.

Arizona: rare and local, primarily in the White Mountains -- central and eastern portions of the state; may be more widespread in the north in years with high levels of rainfall (Piest 2005). See eBird data.

Manitoba: primarily southwest and north ( See eBird data.

Ontario: 2 regions primarily: lake plain areas in the south (Lake Simcoe-Rideau regions) and Hudson Bay lowlands to the north (Gendron 2007). See eBird data.

Ohio: v. rare and local in the north, primarily along the lake plain and associated marshes See eBird data.

Québec: in southern parts of the province; common breeder along the upper St. Lawrence River, between Cornwall and Kamouraska; in the north, abundant in coastal lowlands along Hudson and James bays (McNicoll and Tardiff 1996). See eBird data.

The Maritime Provinces: uncommon and local, primarily e. New Brunswick and PEI (

NY State: uncommon and sporadic breeder, restricted to just a few wetlands in northern (Champlain Valley) and central portions of the state (McGowan 2008; See eBird data.

Winter Range

Along Pacific and Atlantic coasts from se. Alaska and se. Massachusetts south; also in much of the s. U.S. interior through Central America to central Yucatán Peninsula and nw. Costa Rica, and in Bermuda and Cuba (Figure 1).

Winters in distinct areas, especially in Pacific and Gulf Coast states of the U.S. and Mexico, and to a lesser extent along the Atlantic Coast (e.g., central California, Chesapeake Bay, w. Washington, and the Gulf Coast; Hestbeck 1993a). Largest winter concentrations are found in the Fraser River Delta, Puget Sound, and the Lower Columbia River in the Pacific Northwest; Sacramento and San Joaquin valleys, Sacramento-San Joaquin River Delta, San Francisco Bay, and Imperial Valley of California; west coast of Mexico near Los Mochis and Culiacan (state of Sinaloa); interior of Mexico between Guadalajara and Mexico City; east coast of Mexico from Rio Grande to Yucatán Peninsula (Howell and Webb 1995); panhandle, rice prairie, and coast of Texas; rice regions of Arkansas (Grand Prairie) and coastal marshes and central regions (e.g., Catahoula Lake) of Louisiana; and S. Carolina coastal marshes.

Distribution Outside the Americas

Nests in Palearctic from w. Greenland, Iceland, Faroe I., Spitsbergen, and Scandinavia east across arctic areas to Chukotski Peninsula, Kamchatka and Komandorskije Ostrova, and south to British Isles, central Europe, Caspian Sea, Transcaucasia, and Kurile I. (Cramp 1977). Winters in Palearctic from British Isles, s. Scandinavia, s. Russia, Turkestan, and Japan south to n. and e. Africa, Indian Ocean, Borneo, Philippine I., Hawaii, and islands of Micronesia; casual or occasional in Bear I., Madeira, the Azores, and Polynesia (Line, Tokelau, and Cook I.). Some birds that winter in w. U.S. nest in Siberia, especially when prairies are dry (Bellrose 1980).

Nature of Migration

Annual migrations between nesting and wintering areas has been documented by extensive banding records and satellite telemetry. Flight distance is related to latitude of nesting relative to ultimate wintering area. California or other Southwest nesters may not migrate if wintering there; birds nesting in n. Alaska and wintering in Mexico have lengthy, extended migrations. Transoceanic migration over Pacific Ocean (Alaska to California), the Caribbean (Florida to Bermuda and West Indies), and the Gulf of Mexico (Gulf Coast to Yucatán Peninsula) is probably common; also recorded to the South Pacific (e.g., Hawaii; Bellrose 1980).

Inter-regional movements during winter have been documented in California (Fleskes et al. 2002), Louisiana (to Arkansas; Cox and Afton 2000) and Texas (Alford III and Bolen 1977, Moon 2004). Feather and claw isotope signatures can be used to distinguish between Northern Pintails wintering in California and Texas (Clark et al. 2006, 2009).

Timing and Routes of Migration


Details of migration corridors and timing in Bellrose (Bellrose 1980). Major fall migration routes from Alaska and Canadian prairies to central California, Mexico, and the Gulf Coast, especially Texas and Louisiana. Large numbers migrate through the Great Salt Lake, UT, to California and Mexico. Direct route from Alaska to California across the Pacific Ocean, as well as south along the coast and via interior pathways to prairies and ultimately California, Gulf Coast, and Mexico. More than 80% of Alaska-banded birds migrate to Pacific Flyway wintering areas in fall, most to California (Bellrose 1980), but a few to Asia (Nicholai et al. 2005). Two-thirds of Alberta-banded and half of Saskatchewan-banded Northern Pintails migrate to California, the rest to the Gulf Coast and Mexico. Relatively few migrate to the East Coast from the Prairie Pothole Region or Alaska.

One of the first ducks to migrate south in fall. First arrivals, usually adult males, recorded in California the first week Aug; origin unknown (Miller 1985b). In the Atlantic Flyway, females begin southward flights in Aug, with initial arrival on wintering grounds in late Oct (Malecki et al. 2006). Several hundred thousand are already in California by mid-Sep. Heavy migrations in Sep and Oct across the continent. In the Pacific Flyway, fall migration is often direct to wintering regions, most abundant on wintering areas in Dec. Greatest numbers in n. Great Plains Sep–early Oct and s. Great Plains Nov-Dec. In the Great Lakes region and upper Midwest, numbers peak late Oct-early Nov. Most abundant in Gulf Coast wintering regions Dec–Jan. Staging occurs in New Mexico, and likely other wintering areas in U.S., prior to continuing to Mexico (Haukos et al. 2006).

Molt Migration

Summarized by Bellrose (Bellrose 1980). Extensive, usually south from breeding regions, longest of dabbling ducks (Salomonsen 1968). Includes mostly males with some nonbreeding females (Jun–Jul); successful hens remain near brood-rearing areas (Salomonsen 1968). Major molt migrations to marshes of Great Salt Lake, UT (Fuller 1953), se. Idaho (Camas National Wildlife Refuge [NWR]; Oring 1964a); central Saskatchewan (Bellrose 1980); s. Manitoba (Whitewater Lake; Bossenmaier and Marshall 1958); the coast of Hudson Bay, Queen Maude Gulf and the MacKenzie Delta in the Northwest Territories; and the Yukon Delta and North Slope of Alaska.


Based on band recoveries and observations, about 40% of continental spring migrants enter breeding grounds via N. and S. Dakota, the remainder through nw. Montana and Alberta or fly directly to far-northern nesting regions (Johnson and Grier 1988). Most birds probably retrace fall routes (Rienecker 1987a).

Use of satellite telemetry has documented spring migration throughout much of North America. In the Atlantic Flyway, spring migration occurred along two corridors – e. and w. of the Great Lakes (Malecki et al. 2006). In the e. corridor, Northern Pintails staged near Delaware Bay and s. Lake Ontario/St. Lawrence River plain into sw. Quebec, with final destination of northern latitudes in e. Canada (Malecki et al. 2006). The initial destination of birds using the western corridor was the mid-continental Prairie Pothole Region of Dakotas, Manitoba, and Saskatchewan with stopovers noted from north-central Ohio northward to western Lake Erie (Malecki et al. 2006). Arrival of females on mid-continent breeding areas was mid- to late Apr (Malecki et al. 2006).

In the Central Flyway, wintering area determines spring migration route (Haukos et al. 2006). Birds wintering in the Middle Rio Grande Valley, New Mexico, either travel north along the Rio Grande Valley north to s. Colorado, before traveling northeast to the Dakotas and Canada, or initially travel northeast joining birds from Texas in n. Texas or Kansas and settling in s. Saskatchewan (Haukos et al. 2006). Northern Pintails wintering in the Playa Lakes Region of nw. Texas follow three migration routes: (1) NNE to n.-central Kansas or Nebraska Sandhills/western Rainwater Basins; (2) NW through se. and e.-central Colorado; or (3) direct flight to the Dakotas (Haukos et al. 2006). Final destinations for birds wintering in the Playa Lakes Region are the Dakotas/Nebraska, s.-central Saskatchewan, n. Alberta/British Columbia/Nunavut, and n. Manitoba (Haukos et al. 2006). Birds wintering on the Texas Gulf Coast tended to travel northward following the e. sections of the Central Flyway prior to settling in Manitoba, Saskatchewan, or Alberta (Haukos et al. 2006).

In the Pacific Flyway, Northern Pintails wintering in California exhibit five different migration strategies, with four having an extended stopover in s. Oregon-ne. California (SONEC; 80% of birds): over the Pacific Ocean direct to Alaska; along the Pacific Coast with stopovers to Alaska; direct to the Canadian Prairies, using stopovers to reach the Canadian Prairies; and direct to breeding area without stopping at SONEC (Miller et al. 2005a). Alaska is the predominant destination no matter what route is taken for pintails migrating from California.

Northward migration from winter regions begins in late Jan, heaviest in Feb; most leave wintering regions by late Mar and reach nesting regions in Apr and May (Bellrose 1980). Average departure date from wintering grounds in the Central Flyway is mid Mar, earliest in New Mexico and latest from Texas Gulf Coast (Haukos et al. 2006). In the Pacific Flyway, peak initiation of migration is first week of March (Miller et al. 2005a). Stopover duration in the Pacific Flyway (>30 d) is greater than for the Central Flyway (<10 d) (Miller et al. 2005a, 2010; Pearse et al. 2011). Important spring staging areas include the Klamath Basin in SONEC, Rainwater Basin in Nebraska, Cheyenne Bottoms in Kansas, lakes of n.-central Montana, and river courses, deltas, wetlands, and flooded or puddled (snow-melt) agricultural lands across the continent. Staging areas are more consistent for Pacific and Atlantic flyways than the Central Flyway. See also Breeding: phenology.

Migratory Behavior

Gregarious; gathers in large staging flocks. Migrates at night after sunset departure, probably 300–900 m over land (Hochbaum 1955). Clocked at up to 105 km/h by aircraft (Cooke 1937a). Using satellite telemetry, Northern Pintails migrate at an average ground speed of 77 km/h (range 40-122 km/h) (Miller et al. 2005b). Migration usually occurs with a tailwind (Miller et al. 2005a). Longest recorded nonstop flight distance of 2,926 km (Miller et al. 2005a).

Methods Of Orientation

Summarized for waterfowl by Bellrose (Bellrose 1980: 42); for Northern Pintail includes celestial/solar (Hamilton III 1962d), geography (Hochbaum 1955), and probably geomagnetic.

Response To Weather

In N. Dakota and Illinois, little consistent relation between large movements of Northern Pintails and weather variables in fall (fronts, tail wind, temperature, pressure, precipitation, ceiling; Bellrose 1974). Alford and Bolen (Alford III and Bolen 1977) collected circumstantial evidence (increasing proportion of males) of southward migration of females in Texas in relation to cold temperatures. In spring, may reverse migration with a return of winter-like weather (Haukos et al. 2006).

Flight Formations

Spring migration begins with small flocks later coalescing into larger ones (Hochbaum 1955). Fall migration in large flocks of hundreds to thousands, usually in groups of long, wavy, irregular lines with individuals flying abreast.

Control and Physiology of Migration

Migratory Restlessness

Frequent nervous, flighty behavior, including short circling flights by small flocks increasing as spring migration approaches.

Hyperphagia And Fat Deposition

Up to 50% of day spent foraging in California after arrival in fall (Miller 1985b); ducks gain mass, fat, and protein (Miller 1986c), suggesting mass loss occurs during southward migration. Time spent feeding increases in Feb and Mar, coincident with increased consumption of invertebrates and increased body mass (lipid) prior to spring migration from California (Miller Miller 1985b, Miller 1986c). Proportion of day spent feeding increases to nearly 40% in Louisiana rice fields in Feb (Rave and Cordes 1993), but wintering Northern Pintails do not increase body mass before spring migration in Texas (Smith and Sheeley 1993a) or Yucatán, Mexico (Thompson and Baldassarre 1991); these more southerly wintering birds probably accumulate fat en route to nesting regions. Northern Pintails gain body mass in the Rainwater Basin, Nebraska, but amount related to weather conditions (R.R. Cox, unpubl. data).

Response To Photoperiod

No data available for Northern Pintail. Exhibits typical temperate-region photo-refractoriness, in which continued exposure to long day length induces migration and reproduction. Day length mid-Feb to mid-Mar is 11–12 h when radio-tagged Northern Pintails migrate north from Sacramento Valley; wet conditions prolong winter sojourn (MRM).

Roles Of Hormones

No information for this species.

Age/Sex Class Differences

Adult males precede females and young to wintering regions, doing so after wing molt. Sexually segregated flocks noted in fall in Texas, with mixed flocks (70% males) forming in mid-Dec (Smith 1968b); segregated flocks not observed in California (Miller 1985b).

Habitat in Breeding Range

Nesting Habitat

Typically nests in open country with shallow, seasonal, or intermittent wetlands and low vegetation. Nests on islands in shallow basins or, in Alaska, on coastal barrier islands, but most nests are on mainland (Dwernychuk and Boag 1972b, Duncan 1987a, Petrula 1994 Guyn and Clark 2000, Lapointe et al. 2000).

In Alaska, nests on wet sedge (Carex) or grass meadows, slough and river banks, pond shores, and in tidal habitat (Petrula 1994, Grand et al. 1997, C. R. Ely et al. 1997.). Breeding pairs frequent semipermanent wetlands <1 ha characterized by stands of horsetail (Equisetum) and cattail (Typha) (Horn 1991). In boreal forest areas, found in meadows with low sedge and herb growth.

In the Prairie Pothole Region, pairs prefer shallow ephemeral to semipermanent wetlands with emergent vegetation and low upland cover (Podruzny et al. 2002). In N. Dakota, pairs most common on seasonal wetlands (40% of pairs), cultivated wetland basins (24%), and semipermanent wetlands (19%); low use of permanent wetlands, streams, and reservoirs/large impoundments (Stewart and Kantrud 1973). Breeding pair densities positively correlated with area of wetlands per km2 as well as the number of seasonal and semipermanent wetlands (Stewart and Kantrud 1974); densities also associated with interspersion of emergent cover with open water (Kaminski and Prince 1984). Nests are located primarily in upland fields, along roadside allowances or fence rows, and in pastures (Sowls 1955, Keith 1961, Stoudt 1971, Higgins et al. 1992a, Greenwood et al. 1995, McMaster et al. 2005), as well as in spring and fall-seeded cereal crops (Richkus 2002, Devries et al. 2008).

In Arizona, intermittent lakes bordered by spike rush (Eleocharis) preferred in wet years, including wetlands created by irrigation (Brown 1985b).


Adults with broods use shallow seasonal and semipermanent wetlands with abundant emergent cover (e.g., sedge, cattail, bulrush). Interspersion of emergent vegetation with open water provides food resources and escape cover (Mack and Flake 1980, Kaminski and Prince 1984, Peterson 1999).

On the Yukon-Kuskokwim Delta, Alaska, broods use 1- to 5-ha wetlands (0.75–1.5 m deep) with abundant emergent (e.g., sedge) and submersed aquatic vegetation (Potamogeton and Hippurus), and brackish to moderate alkalinity (Grand et al. 1997).

Postbreeding Habitat

Postbreeding males are commonly found on large, shallow marshes with extensive emergent and submersed vegetation that provide abundant cover and food and minimal human disturbance (Salomonsen 1968, Anderson and Sterling 1974). No direct information on habitats used by females following nesting, but large numbers of molting females (and males) are routinely encountered during botulism clean-up and banding operations on large vegetated wetlands in Jul-Aug in prairie Canada (KLG, RGC). On the arctic coastal plain of Alaska, large concentrations of males use large (>85 ha), partially drained basins that by summer have a mosaic of open water and emergent vegetation (Arctophila fulva and Carex aquatilis; Howard 1974c). Following wing molt, also use smaller shallow wetlands where fairy shrimp (Anostraca) and tadpole shrimp (Notostraca) are concentrated (Howard 1974c).

Traditional molting area for males at Camas NWR, ID, is a large (2,000–4,000 ha) shallow marsh; emergent vegetation dominated by hardstem bulrush (Scirpus acutus), common cattail (Typha latifolia), and four-angled water lily (Nymphaea tetragona); submersed vegetation by Potamogeton, naiads (Najas spp.), and hornwort (Ceratophyllum dermersum) (Oring 1964a).

Habitat in Migration

Uses shallow wetlands and flooded agriculture when not frozen, larger lakes and reservoirs, and various estuarine and riverine wetlands. In addition to wetlands, flooded pasture and hay fields in SONEC and flooded cornfields in the Rainwater Basin of Nebraska provide important foraging habitats for spring migrants (Pearse et al. 2011, Fleskes et al. in review). Spring migrants common along James Bay, Ontario, using saltwater habitats such as mud flats, tidal pools, and brackish ponds (Ross 1984b). In the NW Territories, uses extensive wetlands and open bays of the North Arm, Great Slave Lake (Alexander et al. 1991). In Alaska, coastal salt marshes or Cook Inlet and Paristol Bay (R. King pers. comm.). In the Rainwater Basin, Nebraska, complexes of small wetlands with reduced disturbance are often used (Webb et al. 2010).

In fall, large numbers stage at Mills Lake (widening of MacKenzie River, Northwest Territories), an area of emergent and submersed vegetation communities and floating sedge mats (Alexander et al. 1991). Birds arriving in Aug–Sep in central California use moist-soil areas 1–10 cm deep with little emergent cover (MRM), and areas 15–30 cm deep dominated by emergent vegetation (Euliss and Harris 1987). In bottomland lakes of Illinois River Valley, migrants are more abundant and stay longer when favored moist-soil foods are abundant within shallow water levels (Bellrose et al. 1979).

Habitat in the Overwintering Range

A wide variety of shallow inland freshwater and intertidal habitats, most of which include large, shallow wetlands with minimal emergent cover, except some nocturnal use of densely vegetated parts of wetlands. presumably to avoid predation or disturbance (Fleskes et al. 2005a, Moon and Haukos 2008). Uses flooded agricultural habitats (especially rice, corn, wheat, soybeans, and pastures), reservoirs, tidal wetlands, sounds and bays, and estuarine habitats along s. Atlantic, Gulf, and Pacific coastal states but will forage extensively in dry harvested grain fields in some regions during fall and winter (Baldassarre and Bolen 1984, Moon 2004, Anderson and Ballard 2006, MRM).

Along the west coast of mainland Mexico, particularly Ensenada del Pabellon (Sinaloa), favors areas where excess irrigation water flows into salt flats or tidal basins creating fresh to brackish conditions; also mangrove mud flats, irrigation reservoirs, ephemeral ponds, and nearby agricultural habitats (Smith et al. 1989, Migoya et al. 1994). In lagoon-brushland pasture zone of coastal Texas, uses fresh oxbow lakes with some emergent cover and abundant floating and/or submersed vegetation (White and James 1978, Howard and Kantrud 1986). Playa wetlands predominately used both day and night in nw. Texas (Moon and Haukos 2008) but dry field feeding becomes predominant when playa resources are depleted (Moon 2004).

Arkansas' Grand Prairie is an important winter habitat, particularly flooded rice fields (R. R. Cox pers. comm.). In Louisiana, found most commonly in flooded soybean, rice, and pastures in the northeast, in moist-soil habitats of centrally located Catahoula Lake (Smith et al. 1989, Cox and Afton 1997), and in fresh marshes along the Gulf Coast (Howard and Kantrud 1986). Along the S. Carolina coast, uses freshwater wetlands managed in old rice fields, especially in the Santee River estuary; aerial surveys indicate preference for managed units (wigeongrass [Ruppia maritima]/spike rush) and avoidance of natural tidal areas (saltmarsh bulrush) (Smith et al. 1989).

Habitat use in California's Central Valley varies regionally and during the day and night; primarily in wetlands during day and flooded rice fields at night in the Sacramento Valley (Fleskes et al. 2005b), wetlands during the day and flooded corn or wheat fields at night in the Sacramento-San Joaquin River Delta (Fleskes et al. 2005b), wetlands during both day and night in the Suisun Marsh (Coates et al. 2012) and n. San Joaquin Valley (Fleskes et al. 2004, 2005a), and wetlands and a variety of flooded agricultural crop fields during both day and night in the s. San Joaquin Valley (Euliss and Harris 1987, Fleskes et al. 2003).

Historical Changes to the Distribution

No permanent or long-term abandonment or expansion of breeding or wintering ranges for large populations. Shifts in winter (Fleskes et al. 2002, 2005) and possibly breeding (Runge and Boomer 2005) distributions due to long-term habitat changes have been documented; e.g., appears to have expanded breeding range into s. Ontario since the early to mid 1900s (Gendron 2007). Satellite tracking of birds tagged in the Atlantic Flyway (Malecki et al. 2006) suggests a few pintails may breed in ne. Canada, outside the areas shown in Figure 1.

Older breeding distribution summaries exclude e. Canada (Ontario, Quebec, Maritime Provinces); this may reflect recent range expansion or incomplete surveys in an earlier era. First recorded nesting in the Maritime Provinces in 1938; numbers increased to <100 pairs in 1950, declining to <50 in 1980s (Erskine 1992a).

Although thousands reported during the 1940s-1950s, now listed as vagrant or rare winter visitor in Jamaica, Haiti, and the Dominican Republic (Latta et al. 2006).

Fossil History

Fossil remains identified from Pleistocene (Irvingtonian and Rancholabrean North American Land Mammal Age, 2.0–1.8 Megannum to 10,000 years before present) localities in Oregon, California, Florida, Kansas, Nevada, New Mexico, and Texas, and from Nearctic sites in Iowa, Illinois, and Florida (Palmer 1976). Fossils also known from Europe.

Recommended Citation

Clark, R. G., J. P. Fleskes, K. L. Guyn, D. A. Haukos, J. E. Austin, and M. R. Miller (2014). Northern Pintail (Anas acuta), version 2.0. In The Birds of North America (A. F. Poole, Editor). Cornell Lab of Ornithology, Ithaca, NY, USA.