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Nature Of Migration In The Species
These plovers make some of the longest migrations in the world, often with extensive nonstop flights over water. Annual travels extend from tundra breeding grounds to winter ranges on grasslands, coastal wetlands, and other habitats in temperate (Southern Hemisphere) and tropical regions.
Timing And Routes Of Migration
Migrations follow coastal, transoceanic, and transcontinental routes. There is a lengthy period of fall migration since movements vary with breeding success and juveniles depart considerably later than adults. During this time, species are widely dispersed as first arrivals on wintering grounds occur well before last departures from tundra. Spring egress from winter range extends over several weeks, perhaps related to responses of particular populations to timing of snowmelt on various breeding grounds.
Adults begin leaving breeding grounds late Jun to mid-Jul (failed breeders); most depart Aug. Peak migration of adults in Atlantic Provinces of Canada 26 Jul–30 Aug (Morrison et al. 1994). Juveniles linger in northerly regions until late Aug–early Oct. Reports of very late Nov–Dec transients in e. North America (especially Florida and coastal states from Carolinas to Texas) probably all juveniles (see Paulson and Lee 1992 for summary of records).
Well known for an elliptical migratory pattern—offshore nonstop Atlantic route in fall, midcontinental flyway in spring. Bad weather may disrupt normal fall passage, forcing some birds to travel along coastline or move inland. In fall, large numbers of birds (particularly adults) drift southeastward, making 1 or more stops (Hudson and James bays in central Canada, se. Canada, New England coast) before transatlantic flight to South America (Williams and Williams 1990, Wilson and McRae 1993). Southeastern coast of Beaufort Sea may be an important flyway for plovers departing Alaska breeding grounds (see Gudmundsson et al. 2002). Latter source also documents eastward and southward migratory movements of shorebird flocks (many likely represent American Golden-Plovers) at numerous sites in the Northwest Passage region. Additional fall routes: many birds (especially juveniles) travel southward through Missouri, Mississippi, and Ohio river valleys, some through e. U.S. and along Atlantic seaboard, thence to South America via trans-Caribbean flights. Small numbers (adults and juveniles) follow Pacific Coast along a flyway presumably extending to South America; these may be from westernmost breeding grounds (see Paulson 1993). A few travel through the intermountain and mountainous West. Records along Pacific Coast ambiguous since both species occur here and reports usually do not distinguish between them; also, species often uncertain farther inland.
Fall passage through n. South America, Trinidad, and Tobago not well understood. Reported as uncommon to fairly common transient, some birds apparently lingering for several weeks before continuing southward migration; about 9,000 birds seen in coastal Guyana in early Sep 1991 (C. Faanes in litt.). Based on dates of fall sightings, juveniles more likely than adults to island-hop or stop short of South American continent. Many birds probably travel well inland before alighting, and some may fly directly to upper Amazon in Brazil. Only limited knowledge of subsequent movements: transcontinental routes follow north-south river valleys where floodplains (water levels are low at this season), pastures, etc., provide necessary resources; passage of juveniles is farther west (consistent with midcontinental pathway through North America) than that of adults; extensive wetlands in Pantanal region at headwaters of Paraguay River in Brazil likely an important stopover for both adults and juveniles (Antas 1983, Stotz et al. 1992). Rate of passage through interior South America unknown.
Birds arrive on wintering grounds from late Aug to Dec; spring migration begins late Jan, with major exodus in Feb, stragglers until the end of Apr (Dabbene 1920, Wetmore 1927, Olrog 1959, Myers and Myers 1979, Hayes et al. 1990, Sick 1993, Isacch and Martínez 2003a). Northward movements coincide with flooding cycle in Brazil; river-valley habitats used in fall generally unavailable. Antas (1983) suggests that spring route lies mostly west of fall route, passing through upper reaches of Amazonian rivers in Bolivia, Peru, and Colombia. Mass staging unknown; flocks of varying sizes apparently depart nw. South America, with most probably flying nonstop across Caribbean and Gulf of Mexico, others following Central America and Atlantic slope of Mexico. Howell and Webb (1995) report American Golden-Plover as widely distributed spring transient in Mexico, from interior to Atlantic slope (“major concentrations in c. Vera Cruz”) and Pacific slope (“from Colima south”). Unknown whether latter birds continue north along Pacific Coast or cross to continental routes. American Golden-Plovers were not found during 2004-2006 coastal shorebird surveys of northern Yucatan Peninsula, Mexico (Correa-Sandoval and Contreras-Balderas 2008). Records insufficient to clarify spring passage through U.S. Pacific Coast states (see Mlodinow 1993, Paulson 1993, Small 1994).
Migration through U.S. and s. Canada as compiled from numerous sources (partial list – Roberts 1936, Lowery 1974, Oberholser 1974, Dinsmore et al. 1984, Mumford and Keller 1984, Godfrey 1986, Tufts 1986, Janssen 1987, Bohlen 1989, Peterjohn 1989, Thompson and Ely 1989, South Dakota Ornithol. Union 1991, Stevenson and Anderson 1994, Alexander and Gratto-Trevor 1997): first arrivals in Texas, Louisiana, and Florida late Feb–early Mar, movements continue until May; some birds alight on or near coastline, others may continue much farther inland; variable arrival (no doubt influenced by weather) early to late Mar, with major influx during Apr over broad region of the Missouri, Mississippi, and Ohio river valleys, primarily from Kansas to Kentucky northward to the Dakotas and s. Minnesota; also a few birds along Atlantic Coast of U.S. and Canada to at least Nova Scotia and in the far west from Nevada and Utah to n. Idaho and w. Montana (some of last might be from Pacific flyway and could include Pacific Golden-Plovers); passage through upper midwestern U.S. and s. Canada usually peaks early to mid-May.
Arrival on breeding grounds influenced by latitude and annual variations in snowmelt; mostly mid-May to mid-Jun (Irving 1960, Drury 1961, Parmelee et al. 1967, Jehl and Smith 1970, Martin and Moiteret 1981, Kessel 1989, Connors et al. 1993, Johnston and Pepper 2009). Earliest arrivals possibly end of Apr at southern edge of range in Alaska (based on Pacific Golden-Plover, see below); latest around third week of Jun in extreme northern part of range on Devon I., NU (Hussell and Holroyd 1974). Passage along coast of the Beaufort Sea may be a major spring pathway for birds breeding in Alaska (Richardson and Johnson 1981).
Remaining on winter range during boreal summer (over-wintering) is apparently unusual for American Golden-Plover, but a few birds have been reported at various sites in South America including Argentina, Chile, and Uruguay (Blanco and Canevari (1998). The full extent of this behavior is uncertain. Wetmore (1926) noted scattered individuals during the Argentine winter and considered them either “wounded, sterile, or otherwise diseased” or lacking in “physiological incentive” for migration. Studies of Pacific Golden-Plover (in which over-wintering is common, see below) support the last alternative and suggest that stragglers in South America represent some fraction of first-year birds that are not yet sexually mature.
Four long-distance recoveries of banded American Golden-Plovers (Bird Banding Lab [BBL]) consistent with migratory routes described above; records (locations and dates [month/year] where banded and recovered) as follows: southern Quebec (Mauricie Region) 9/1948–ne. Guyana (near Georgetown) 1/1949, northern Illinois (LaSalle County) 10/1955–Lesser Antilles (Grande Terre I.) 9/1958, Victoria I. NU 7/1960–ne. Texas (near town of Pittsburg) 3/1962, ne. Alaska (approx. 80 km south of Deadhorse) 6/1979–Lesser Antilles (exact location uncertain) 10/1979. For additional discussion of spring and fall migration, including routes in both North and South America and counts of birds stopping over in various regions, see Clay et al. (2009).
Small numbers follow the Pacific Coast of North America and winter primarily in California. Large numbers migrate along the Asian coast (East Asian-Australasian flyway; see Lane and Parish 1991) with many diverging southeastward on oceanic pathways to w. Pacific islands (Marianas, Carolines, etc.) and Australia; another major corridor occurs in the mid-Pacific from Alaska and e. Siberia to the Hawaiian I. and beyond (Williams and Williams 1988, 1990; Williams and Ying 1990; Johnson et al. 1997b, 2001a, 2004a). The Asian and mid-Pacific flyways converge in the Bering Sea region; thus each pathway may carry birds from both the New and Old worlds.
Male plover banded at nest on the Seward Peninsula summer 2008 and subsequently photographed in Japan indicates Alaska link with Asian Flyway (OWJ unpubl. record). Furthermore, major east (in fall) west (in spring) bird traffic across Bering Strait region between Alaska and Siberia has been revealed by radar (Flock 1972, Alerstam et al. 2007, Hedenström et al. 2009). These movements almost certainly include Pacific Golden-Plovers and suggest connectivity between Siberian breeding grounds and eastern reaches of the winter range (Hawaii, Samoan I., etc.) via Alaska and the mid-Pacific flyway. Such linkages consistent with lack of variation in DNA samples from various sites across the breeding and non-breeding ranges (R. Gold, pers. comm).
Many Pacific Golden-Plovers winter in countries bordering or near n. Indian Ocean (see Distribution: outside the Americas, above) and migrate via transcontinental routes across central and w. Asia (Kozlova 1961, Dolnik 1990, Gavrilov et al. 1993, Syroechkovski and Lappo 1994, Tomkovich et al. 2000). Perhaps most are from the western portion of the breeding range. Possibly, some birds wintering in China, se. Asia, and western Pacific islands also follow continental flyways (see Johnson et al. 2006). The Taimyr Peninsula appears to be a migratory divide from which routes connecting Siberia to wintering grounds extend westward and eastward (Alerstam and Gudmundsson 1999a), with eastbound passages toward North America apparently crossing large expanses of the Arctic Ocean (Alerstam and Gudmundsson 1999b, Alerstam et al. 2007).
Numerous Pacific Golden-Plovers winter in the Hawaiian I. and studies in 2009 using geolocators indicated a direct north-south flyway to Alaska (OWJ, R. Goodwill, R. Gold, unpubl. data). Previously, birds banded on Oahu and sighted in Alaska (at Homer, Naknek, Bethel, Kivalina, and Nome; OWJ and P. Bruner unpubl. records), together with numerous plovers radio-tagged at wintering sites on Oahu and detected subsequently in Alaska (Johnson et al. 1997b, 2001a, 2004a, OWJ unpubl. findings), substantiate a major migratory link between the two regions. Routes to the south and southwest of Hawaii are less certain. Whether birds passing through Hawaii in fall return via the same flyway in spring is unknown. Perhaps some do, whereas others follow an elliptical route similar to that of Ruddy Turnstones (Arenaria interpres): south from Alaska in fall to central and s. Pacific, returning in spring along Asian coast (Thompson 1973, McClure 1974). Most migration to and from Australia is probably via the northeastern end of continent along East Asian-Australasian flyway. Nonstop flights linking Asia with Australia, New Zealand, and other sw. Pacific wintering grounds seem likely (Williams and Williams 1990, Barter 1992, Minton et al. 2010).
Overall timing of fall and spring migrations (see Fig. 4) similar to that of American Golden-Plover. Some reports suggest regional variation between the species: among adults, Pacific Golden-Plovers arrive 2–3 wk earlier than American Golden-Plovers during fall migration in California (Small 1994); among juveniles, peak fall migration of Pacific Golden-Plovers possibly 1 mo later than of American Golden-Plovers along Pacific Northwest coast (Paulson 1993). Most adult Pacific Golden-Plovers arrive Hawaiian I. in Aug (earliest arrivals usually females), most juveniles in Oct; spring departure relatively abrupt in late Apr–early May (Johnson et al. 1981, 1989, OWJ).
At extreme southern end of range in Australia and New Zealand, periods of fall and spring migrations more prolonged (Marchant and Higgins 1993, Watkins 1993, Alcorn et al. 1994). Late fall arrivals and early spring departures in southern hemisphere indicate lingering en route possibly along Asian flyways or amidst islands of central and w. Pacific. Similar early departure (mid-April) occurs in Samoan I. (Tarburton 2001, Johnson et al. 2008a), suggesting stopovers during northward passage. Migrants peak in Japan late Apr to mid-May (Brazil 1991, Watanabe 1991); passage through se. Kazakhstan and s.-central and central Russia late Apr to late May (Dementiev et al. 1951, Moskvitin 1973, Rogacheva 1992).
Telemetry studies suggest that the Alaska Peninsula is a major corridor for transpacific migrants entering the Alaska region (Johnson et al. 1997b, 2001a, 2004a). Flocks of migrants occur on the peninsula last 2 wk of Apr through early May (R. Gill, Jr., in litt.). Whether reports of spring aggregations at Montague and Middleton islands (Isleib 1979, Mlodinow 1993) indicate regular westward passage via those sites is uncertain. As with American Golden-Plover, arrival on breeding grounds varies regionally according to latitude and seasonally with variations in timing of snowmelt: Alaska from late Apr (Alaska Peninsula, S.Savage in litt; Bethel region, B. McCaffery in litt.) to mid-May (Seward Peninsula; Kessel 1989, Connors et al. 1993); St. Lawrence I. late May to early Jun (Sauer 1962, Johnson 1975); Siberian breeding range late May to 3rd week Jun (Dementiev et al. 1951, Portenko 1972, Kistchinski 1980, Dorogoy 1982, Kondratiev 1982, Danilov et al. 1984, Tomkovich 1988b, Rogacheva 1992, Svirodova 2000, Tulp 2007).
Many records of Pacific Golden-Plovers remaining on winter range during boreal summer (e.g., Johnson and Morton 1976, Hewish 1988, Stinson et al. 1997, Tarburton 2001, Johnson et al. 2004c). Over-wintering (or over-summering if on Northern Hemisphere range, as in Hawaii) most common among first-year birds, occasional in second-year birds (Johnson and Johnson 1983, Johnson 1985, Hewish 1988, 1989). Although some individuals develop full Alternate plumage, ovaries remain inactive and testes show only slight recrudescence (Johnson 1973, 1979). Possibility that the proportion of over-wintering birds might be higher with increasing distance from breeding grounds (Johnson and Johnson 1983) not substantiated in Australia where relatively few plovers fail to migrate northward (Hewish 1988, Skewes 2007, Kearney et al. 2008). However, observations above 30°S in Australia and elsewhere across Pacific suggest that first-year Pacific Golden-Plovers departing extreme southern regions might travel varying distances then arrest migration and over-winter or over-summer depending on location (Hewish 1988, Johnson et al. 2004c).
Four long-distance recoveries of Pacific Golden-Plovers banded during fall on St. George I. (Pribilofs), AK (Woodward 1972, McClure 1974, Wyndham 1977, Johnson et al. 1989, BBL): at Kure Atoll (nw. Hawaiian I.), Hokkaido I. (near Yakumo), Solomon I. (Malaita I.), and east coast of Australia (New South Wales). First and second recoveries indicate birds following mid-Pacific and East Asian-Australasian flyways, respectively; routes of other 2 uncertain. In the order given, recoveries were 26 d, 48 d, 14 mo, and 3 yr 7 mo after banding.
A plover wearing a Russian band was found in 1981 at Kwajalein Atoll, Marshall I. (Schipper 1985). Because of subsequent confusion over the band number, exact banding location was never determined (OWJ), but probably the bird was captured as a chick in the lower Kolyma River region (P. Tomkovich in litt.). Two recoveries of birds banded in Australia, one at Vanuatu, the other in China at southern end of the Yellow Sea (Minton et al. 2006).
Essentially nothing known about behavior of migrants in remote areas such as interior South America (American Golden-Plover), interior Asia and most of the Pacific (Pacific Golden-Plover), and much of Alaska (both species). Lack of information even in more populated regions, as with spring passage of American Golden-Plover through prairies of U.S. and Canada (Morrison and Myers 1987). Movements often involve small flocks feeding opportunistically, but migratory aggregations also occur in both species (e.g., Clapp and Wirtz 1975, Isleib 1979, Pyle 1984, Janssen 1987 , Jorgensen 2004, 2008, Clay et al. 2009). Annual gatherings are frequently in the same region, though less dramatic and predictable than the mass staging of some shorebirds at traditional stopover sites.
Exceptional situations involving large spring build-ups have been reported. For American Golden-Plovers this occurs in nw. Indiana (Benton Co. and White Co.) and adjacent ne. Illinois (Mumford and Keller 1984, Bohlen 1989). In Benton Co., yearly census counts around 21 Apr 1980–1986 ranged from about 4,000 to >25,000 birds (Erickson 1992). In Benton and White Cos., counts throughout the spring of 1998 indicated at least 42,000 birds (possibly as many as 84,000) had stopped over amidst the region’s agricultural fields (Braile 1999).
For Pacific Golden-Plovers, the most significant known stopover sites (involving many thousands of birds in both spring and fall) are in the Torey Depression and Selenga River Delta regions of ne. Mongolia and adjacent Russia (Fefelov 2003, Fefelov and Tupitsyn 2004, Johnson 2003, Bamford et al. 2008, J. Jukema in litt.), and in Japan (Brazil 1991; Fujioka et al. 1998a,b; Bamford et al 2008). According to numerous reports (including Barter et al. 1998, 2000, 2002; Moores 1999; Ge et al. 2006, Yang et al. 2008, Reigen et al. 2009), very few Pacific Golden-Plovers occur amidst the massive aggregations of other shorebirds using mudflats along the Asian coast, evidently this plover prefers to forage at inland sites.
At major landfalls (e.g., Pacific Golden-Plovers arriving in Alaska or American Golden-Plovers reaching South America) birds appear to overfly littoral areas in favor of upland habitats, although a relatively large coastal influx might escape detection if flocks were scattered. Generalization that southbound Pacific Golden-Plovers move through Asia gradually with prolonged stays at feeding sites (Dementiev et al. 1951) remains unsubstantiated without additional studies. Oceanic stopovers on atolls and islands are no doubt basic to transpacific flights of Pacific Golden-Plover. Hawaiian I. are visited by many southbound birds (especially juveniles), also by northbound migrants (Ely and Clapp 1973, Clapp and Wirtz 1975, Pyle 1984, Williams et al. 1986). Some plovers en route to s. Pacific winter range probably overfly the Hawaiian archipelago, stopping instead at the Marshall, Line, and Phoenix islands (see Marks and Redmond 1994).
Shorebird migrations often involve nonstop flights of at least 5,000–7,000 km (exceptionally much farther, see Gill et al. 2008). Thus, some plovers (perhaps experienced adults) possibly fly nonstop from breeding grounds to distant landfalls or directly to wintering grounds. Large areas of American Golden-Plover breeding range are within 6,500–7,000 km of n. South America; Pacific Golden-Plover breeding grounds in w. Alaska are approximately 5,000 km from Hawaiian I., and much of the Siberian Pacific Golden-Plover range is within 5,000–6,000 km of se. Asia and India.
Old reports (Cooke 1910) of American Golden-Plover fall staging grounds in Labrador and Nova Scotia have been widely accepted. However, there is no hard evidence from Labrador (Todd 1963) and no large concentrations have appeared in Nova Scotia during the twentieth century (Godfrey 1986). Either past observations were inaccurate or migratory routes have changed (also see Byrkjedal and Thompson 1998).
Premigratory behavior remains unreported for the American Golden-Plover. In Hawaii, departure of Pacific Golden-Plovers is often preceded by flocking over several days. Flocks seem to depart mostly in the afternoon (OWJ), possibly at night also. Departing flock either circles or makes angled ascent to great height, heads out to sea until lost from view (various reports including personal observations summarized by Johnson et al. 1981). Similar behaviors may accompany departure of Pacific Golden-Plovers from some regions of the Arctic (Sauer 1962).
From extensive radar studies in the Siberian and North American Arctic, altitudes of migrating shorebird flocks (some of which were no doubt composed of American or Pacific golden-plovers, depending on location) were commonly around 1 km, often 2-4 km, occasionally >5 km (Alerstam and Gudmundsson 1999a,b; Gudmundsson et al. 2002; Alerstam et al. 2007; Hedenström et al. 2009). Highest measured altitude of flocks (including probable American Golden-Plovers) were 6 km over the Caribbean (Williams and Williams 1990), 6.65 km at Nova Scotia (Richardson 1979). Transoceanic navigation may be relatively simple: constant compass heading combined with effects of prevailing winds (Williams and Williams 1990). Only known inland altitudes are for American Golden-Plovers migrating during daylight hours in e.-central Alaska, most flocks at about 50 m (Cooper and Ritchie 1995).
Control And Physiology
No information concerning proximate stimuli in either species. Significant studies limited to Pacific Golden-Plover. Sauer (1963), using birds hand-raised from chicks, reported migratory restlessness in May and Sep, plus ability for orientation. Fat cyclicity and flight ranges in Pacific Golden-Plovers wintering on Wake I. examined by Johnston and McFarlane (1967). Similar but more extensive investigation on Oahu, HI, by Johnson et al. (1989) revealed: fat-free weight approximately 105 g; adults arriving in fall averaged 118 g, juveniles 109 g; these weights trended down for several weeks, possibly because of physiological rigors associated with establishing and re-establishing residency combined with molting; major premigratory weight gains began late Mar about 1 mo before migration; departing birds may exceed 200 g, with fat accounting for ≥ 37% total weight; fat-free dry weights lowest in fall when birds return to predictably favorable winter range, highest in spring as migrants apparently maximize protein reserves against unpredictable weather and food supplies on breeding grounds; depending on formulae and flight speed, migrants departing Oahu (assuming 175 g total body weight) have ranges of 5,000–8,000 km; in very fat individuals (> 200 g), ranges perhaps exceed 10,000 km. Additional elements related to migration (energetics, hormonal cycles, cyclic regression and re-growth of digestive organs, etc.) not yet studied in these plovers, but likely similar to other long-distance migrant shorebirds (see O’Reilly and Wingfield 1995; Klaassen 1996; Piersma and Gill 1998; Battley et al. 2000, 2001; Piersma and Jukema 2002). Probable energetic scenario at arrival on breeding grounds or northern landfalls: nutrient reserves used in part for regeneration of digestive system (particularly in Pacific Golden-Plovers following transoceanic routes) and as insurance against lack of food should tundra be snow-covered, energy for egg production likely depends on resources acquired after arrival (see Tulp et al. 2009b).