Pacific Golden-Plover Pluvialis fulva Scientific name definitions
Version: 1.1 — Published April 15, 2021
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Movements and Migration
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Dispersal and Site Fidelity
Natal Philopatry and Dispersal
Fidelity to Breeding Site
Studies of marked birds on the Seward Peninsula, Alaska and on the Taimyr Peninsula, Far North Russia, showed strong gender-bias with males much more site-faithful than females (167, 168, 169; OWJ, unpublished data). Males reoccupy the same territories annually, generally with a new mate each season. It appears that females return to the same breeding locale, but often are neither mate-specific nor site-specific enough to be detected. On the Seward Peninsula, missing females have eventually been found nesting from 1.1 to 5.0 km from the nests where they were originally captured (OWJ unpublished data). On the Taimyr Peninsula, a female Pacific Golden-Plover was reported nesting at 924 m from her previous nest and another female likely nested at 1.5–2 km from a previous nest (168). Thus, reunion of the same pairs, while it does sometimes occur (see 167, 170), is infrequent and unlikely.
The range of interyear distances between nests of marked males varies widely: 0 m (reuse of same nest) to 420 m (mean = 152 m, n = 31) on the Seward Peninsula (167; OWJ, unpublished data); 100–450 m (n = 3) on the northwestern Taimyr Peninsula; 15–530 m (n = 16) on the southeastern Taimyr Peninsula (168); and 8–432 m (n = 12) on the northwestern Taimyr Peninsula (170). Within the same breeding season, the distance between first nests and replacement nests ranged from 74–457 m (n = 4) on the Seward Peninsula (171), and 66–330 m (n = 3) on the northwestern Taimyr Peninsula (170).
Fidelity to Overwintering Site
High return rates of Pacific Golden-Plover are typical on Oahu, Hawaii, overwintering grounds. Territorial birds reoccupy same territories, and non-territorial birds frequent same communal sites annually (172, 18, 155, 156, 159; OWJ and P. Bruner, unpublished data). The same pattern was found at Johnston Island, Saipan Island, and American Samoa (19, 157, 158). No other comparable studies elsewhere on the overwintering range.
This plover makes some of the longest migrations in the world, often with extensive nonstop flights over water. Annual travels extend from tundra breeding grounds to overwintering grounds, primarily in Oceania and Southeast Asia.
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. Non-breeders leave 2–4 weeks before breeders, and young some weeks after adults. Generally departs late August or early September. During this time, species is widely dispersed as first arrivals on overwintering grounds occur well before last departures from tundra. Spring egress from the overwintering range and stopovers extends over several weeks, apparently related to responses of particular populations to timing of snowmelt on various breeding grounds.
Small numbers follow the Pacific Coast of North America and overwinter primarily in California. Large numbers migrate along the Asian coast (East Asian–Australasian flyway) en route to various islands (e.g., Marianas, Philippines); another major corridor occurs in the mid-Pacific from Alaska and eastern Siberia to the Hawaiian Islands and beyond (173, 174, 175, 176, 177, 178, 98, 102, 179, 180). The East Asian–Australasian and mid-Pacific flyways converge in the Bering Sea region; each pathway carries birds from both the New and Old worlds. Such connectivity is consistent with lack of variation in DNA samples from various sites across the breeding and non-breeding ranges (R. Gold, personal communication).
Tracking with geolocators indicates that Pacific Golden-Plovers overwintering in the Hawaiian Islands make direct flights to and from Alaska, whereas individuals overwintering farther south in the Central and South Pacific (Oceania) traverse a much longer elliptical route (179). The latter involves lengthy spring stopovers in Japan and return flights via the mid-Pacific corridor in fall. Furthermore, the plovers on these two pathways do not disperse evenly across the nesting range in Alaska. Rather, birds traveling the direct route from Hawaii arrive earlier and mostly nest farther south than birds arriving via the elliptical route from islands beyond Hawaii. Latitudinal separation between the two groups appears to occur at approximately 61°N on the Yukon–Kuskokwim Delta. The stopovers recorded in Japan (about 3 weeks) likely indicate vital refueling for birds traveling northward from Oceania (181). Migrants arrive in Japan from April to mid-May (182, 183, 102, 179, 180).
Fall and spring flights on the East Asia–Australasian flyway are more or less direct with stopovers (especially spring) in South Korea, Taiwan, China, Japan (see 180). Many Pacific Golden-Plovers migrate via transcontinental routes across central and western Asia (184, 185, 186, 187, 188, 71) and likely breed in corresponding areas of the nesting range in Siberia; passage through southeastern Kazakhstan and south-central and central Russia from late April to late May (22, 189, 141, 71). East Africa and western Asia probably reached by overland migration across western Siberia; birds that winter in India cross Tibet.
Numerous Pacific Golden-Plovers overwinter in Hawaii and the overall timing of their fall and spring migrations is shown in Figure 2. Most adult birds arrive in the Hawaiian Islands in August (earliest arrivals usually females), most juveniles in October; spring departure is relatively abrupt from late April–early May (172, 18, 102, 179, OWJ). On nonbreeding grounds farther south than Hawaii timing is more variable with spring departure often earlier in April and fall arrival of adults later extending into September (12, 190, 191, 153, 192, 158, 179, 180). Such scheduling associated with stopovers (especially spring in Japan), although many birds make nonstop transoceanic flights in fall. Arrival in Fiji in early September, New Zealand in October. Records from central Pacific islands as early as late July and until May (193), with observations in Micronesia and Samoa even in June (194, 195).
Flocks of migrant Pacific Golden-Plovers observed on the Alaska Peninsula last 2 weeks of April through early May (R. Gill Jr., personal communication) likely represented plovers arriving from non-breeding grounds in the Hawaiian Islands (see 179). Whether reports of spring aggregations at Montague and Middleton Island, Alaska (196, 197) indicate regular westward passage via those sites is uncertain. Arrival on breeding grounds varies regionally according to latitude and seasonally with variations in timing of snowmelt (see 179): late April on Alaska Peninsula and in Bethel region (S. Savage and B. McCaffery, personal communications); mid-May on Seward Peninsula (198, 85); late May to early June on St. Lawrence Island (47, 199); late May to the third week June on Siberian breeding ranges (22, 148, 149, 69, 150, 200, 151, 141, 168, 201, 180). Appears in Siberian tundra in early June; Taymyr population migrates over middle taiga along Yenisey River in late May, stopping in parties of 3–7 birds at thawed patches in forest openings and near villages.
Many records of Pacific Golden-Plovers remaining on overwintering range during the boreal summer (e.g., 67, 202, 161, 153, 19). Overwintering (or oversummering if on Northern Hemisphere range, as in Hawaii) most common among first-year birds, occasional in older birds (29, 30, 202, 203). Although some non-migrants develop full Alternate plumage, ovaries remain inactive and testes show only slight recrudescence (204, 205). Possibility that the proportion of overwintering birds might be higher with increasing distance from breeding grounds (29) not substantiated in Australia where relatively few plovers fail to migrate northward (202, 206, 207, 192). However, observations above 30°S in Australia and elsewhere across Pacific suggest that some first-year birds departing extreme southern regions might travel varying distances then arrest migration and overwinter or oversummer depending on location (202, 19).
As of September 2017, there were 52 recoveries/encounters of Pacific Golden-Plovers in files of the U.S. Geological Survey, Bird Banding Laboratory (BBL). Various of these were over long distances such as Pribilof Islands to Australia and Solomon Islands (208, 209, 210). For numerous fixes showing lengthy migratory movements of birds carrying geolocators, see 102, 179, and 180. Two recoveries of birds banded in Australia, one at Vanuatu, the other in China at southern end of the Yellow Sea (211, 212). Sixteen birds variously color-banded at Nome, Alaska, and in Saipan, American Samoa, and Australia were reported at stopovers in Japan, and four others color-banded at Nome, Alaska, were resighted in Queensland, Australia (OWJ, unpublished data); one individual flagged at a nest in southeastern Chukotka was observed at Darwin, Northern Territory, Australia (180).
A Pacific Golden-Plover wearing a Russian band was found in 1981 at Kwajalein Atoll, Marshall Islands (213). 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 of Chukotka, Far Eastern Russia (P. Tomkovich, personal communication). Recent geolocator studies confirmed connectivity between Chukotka and the Marshall Islands of Kwajalein (179) and Majuro (180).
Very little known about behavior of migrants in remote areas such as interior Asia, most of the Pacific, and much of Alaska. Movements may involve birds feeding opportunistically, but migratory aggregations also occur (see 214, 196, 215). Large build-ups (especially in spring) have been reported. The most significant known stopover sites are in the Torey Depression and Selenga River Delta regions of northeastern Mongolia and adjacent Russia (216, 217, 163, 112, 71), and in Japan (182, 218, 219, 112, 179). According to numerous reports (including 220, 221, 222, 223, 224, 225, 226), relatively few Pacific Golden-Plovers occur amidst the massive aggregations of other shorebirds using mudflats along the Asian coast, suggesting this plover prefers to forage at inland sites (180); however, see 227.
At major landfalls (e.g., Pacific Golden-Plovers arriving in Alaska) birds appear to overfly littoral areas in favor of upland habitats, though coastal influx might escape detection. Generalization that southbound Pacific Golden-Plovers move through Asia gradually with prolonged stays at feeding sites (22) seems unlikely, needs further study. Oceanic stopovers on atolls and islands during migrations appear to be infrequent with trans-Pacific flights being nonstop (102, 179, 180), similar to lengthy nonstop flights in other species of shorebirds (see 228). All transoceanic flights were non-stop and typically wind-assisted at a mean minimum migration speed of 49 km/hour, over periods of 3–8 days (229), with other studies having recorded even faster mean flight speeds (230). Some Alaskan breeders fly 4,500 km non-stop across Pacific Ocean to Hawaiian Islands .
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 ascends to great height, heads out to sea until lost from view (various reports including personal observations summarized by 172). Similar behaviors may accompany departure from northern regions in fall (47), and geolocator tracking indicates these birds make pre-migratory stopovers of up to 40 d at sites located south of nesting grounds (98, 102, 180).
From extensive radar studies in the Arctic regions of Siberia and North America, altitudes of migrating shorebird flocks (some of which were likely composed of Pacific Golden-Plover or American Golden-Plover, depending on location) were commonly around 1 km, often 2–4 km, occasionally > 5 km (231, 232, 233, 234, 235). Highest measured altitude of flocks (including probable American Golden-Plovers) were 6 km over the Caribbean (174), 6.65 km at Nova Scotia (236). Capacity for high altitude flight is likely similar in the Pacific Golden-Plover. Transoceanic navigation may be relatively simple: constant compass heading combined with effects of prevailing winds (174).
Control and Physiology of Migration
No information concerning proximate stimuli. Sauer (237), using birds hand-raised from chicks, reported migratory restlessness in May and September, plus ability for orientation. Fat cyclicity and flight ranges in Pacific Golden-Plover overwintering on Wake Island, northeastern Micronesia, were examined by Johnston and McFarlane (65). Similar, but more extensive investigation (18) 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 pre-migratory weight gains began late March about one month 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 overwintering 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, Hawaii (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 (e.g., energetics, hormonal cycles, possible cyclic regression and regrowth of digestive organs) not yet studied in these plovers, but likely similar to other long-distance migrant shorebirds (see 238, 239, 240, 241, 242, 243, 228). Probable energetic scenario at arrival on breeding grounds or northern landfalls after lengthy transoceanic flight: nutrient reserves used in part for regeneration of digestive system and as insurance against lack of food should tundra be snow-covered, energy for egg production likely depends on resources acquired after arrival (see 244).