Behavior

Walking, Running, Hopping

This species commonly walks and runs. The birds often stand on one leg while loafing or roosting. If disturbed, they may hop on that leg for a considerable distance before changing gait. Except with certain topographic features of tundra landscape (see Sounds: Vocalizations), locomotion does not involve elevated perches.

Flight

The American Golden-Plover is capable of swift and extended flight. These plovers are considered "the high speed champions among shorebirds" (7 Paulson, D. R. (1993). Shorebirds of the Pacific Northwest. University of Washington Press, Seattle, Washington, USA. Close ). Numerous records of the Pacific Golden-Plover (Pluvialis fulva) equipped with geolocators indicate an average ground speed of around 50 kph during long trans-oceanic flights, with considerable variation (38 to > 100 kph) presumably influenced by the direction of winds (231 Johnson, O. W., L. Fielding, J. W. Fox, R. S. Gold, R. H. Goodwill, and P. M. Johnson (2011). Tracking the migrations of Pacific Golden-Plovers (Pluvialis fulva) between Hawaii and Alaska: New insight on flight performance, breeding ground destinations, and nesting from birds carrying light level geolocators. Wader Study Group Bulletin 118:26–31. Close , 232 Johnson, O. W., L. Fielding, J. P. Fisher, R. S. Gold, R. H. Goodwill, A. E. Bruner, J. F. Furey, P. A. Brusseau, N. H. Brusseau, P. M. Johnson, J. Jukema, L. L. Prince, M. J. Tenney, and J. W. Fox (2012). New insight concerning transoceanic migratory pathways of Pacific Golden-Plovers (Pluvialis fulva): The Japan stopover and other linkages as revealed by geolocators. Wader Study Group Bulletin 119:1–8. Close , 199 Johnson, O. W., R. R. Porter, L. Fielding, M. F. Weber, R. S. Gold, R. H. Goodwill, P. M. Johnson, A. E. Bruner, P. A. Brusseau, N. H. Brusseau, K. Hurwitz, and J. W. Fox (2015). Tracking Pacific Golden-Plovers Pluvialis fulva: Transoceanic migrations between non-breeding grounds in Kwajalein, Japan and Hawaii and breeding grounds in Alaska and Chukotka. Wader Study 122:4–11. Close , 233 Johnson, O. W., P. S. Tomkovich, R. R. Porter, E. Y. Loktionov, and R. H. Goodwill (2017). Migratory linkages of Pacific Golden-Plovers Pluvialis fulva breeding in Chukotka, Russian Far East. Wader Study 124:33–39. Close ). Presumably, flight speeds of the American Golden-Plover during intercontinental migrations are similar. According to radar measurements in various arctic regions, ground speeds of migrating shorebird flocks (some were probably American Golden-Plovers) were often > 72 kph, ranging as high as 136 kph (189 Richardson, W. J. (1979). Southeastward shorebird migration over Nova Scotia and New Brunswick in autumn: A radar study. Canadian Journal of Zoology 57:107–125. Close , 185 Alerstam, T., and G. A. Gudmundsson (1999a). Bird orientation at high latitudes: flight routes between Siberia and North America across the Arctic Ocean. Procedings of the Royal Society of London B 226:2499–2505. Close , 186 Alerstam, T., and G. A. Gudmundsson (1999b). Migration patterns of tundra birds: tracking radar observations along the Northeast passage. Arctic 52:346–371. Close , 143 Gudmundsson, G. A., T. Alerstam, M. Green, and A. Hedenström (2002). Radar observations of Arctic bird migration at the Northwest Passage, Canada. Arctic 55:21–43. Close , 187 Alerstam, T., J. Bäckman, G. A. Gudmundsson, A. Hedenström, S. S. Henningsson, H. Karlsson, M. Rosén, and R. Strandberg (2007). A polar system of intercontinental bird migration. Proceedings of the Royal Society B 274(1625):2523–2530. Close , 188 Hedenström, A., T. Alerstam, J. Bäckman, G. A. Gudmundsson, S. Henningsson, H. Karlsson, M. Rosén, and R. Strandberg (2009). Radar observations of arctic bird migration in the Beringia region. Arctic 62:25–37. Close ). The mean ground speed of a single American Golden-Plover tracked between Illinois and Iowa was 56 kph (62 Stodola, K. W., B. J. O'Neal, M. G. Alessi, J. L. Deppe, T. R. Dallas, T. A. Beveroth, T. J. Benson, and M. P. Ward (2014). Stopover ecology of American Golden-Plovers (Pluvialis dominica) in Midwestern agricultural fields. Condor 116:162–172. Close ).

Swimming and Diving

This is not an aquatic species. The sighting of a plover on the ocean surface during migratory periods would probably indicate a bird forced to rest on water by dense fog (see 234 Piersma, T., B. Spaans, and A. Dekinga (2002). Are shorebirds sometimes forced to roost on water in thick fog? Wader Study Group Bulletin 97:42–44. Close ), fatigue, or injury.

Preening, Head-Scratching, Stretching, Bathing, Anting

Preening and stretching usually accompany periods of loafing. Head-scratching is indirect, with the foot passed over a lowered wing (O. W. Johnson). Migrant juvenile American Golden-Plovers were observed bathing in Iowa wetlands during fall 1988 and 2000 (M. Kenne, personal communication). Bathing occurs on the overwintering grounds (147 Myers, J. P., and L. P. Myers (1979). Shorebirds of coastal Buenos Aires Province, Argentina. Ibis 121:186–200. Close ). There are no reports of bathing on nesting grounds and no records of anting.

Sleeping, Roosting

Overwintering Pacific Golden-Plovers in Hawaii sleep intermittently during daylight hours, squatting or standing on one leg, either with their head resting on their shoulders and bill forward, or with their head turned and bill tucked beneath the scapulars. Presumably, the American Golden-Plover is similar, but there are no reports. Brief intervals of sleep likely occur on breeding grounds, but no information is available.

The American Golden-Plover roosts in flocks during the nonbreeding season. In Argentina, it joins other shorebirds on multispecific nocturnal roosts "at the edge of lagoons and swamps" (235 Myers, J. P. (1984). Spacing behavior of non-breeding shorebirds. In Shorebirds: Migration and Foraging Behavior (J. Burger and B. L. Olla, Editors). Plenum Press, New York, NY, USA. pp. 271–321. Close ). Rooftop roosting by Pacific Golden-Plovers occurs in Hawaii and elsewhere on the overwintering range (230 Johnson, O. W., and R. M. Nakamura (1981). The use of roofs by American Golden-Plovers Pluvialis dominica fulva wintering on Oahu, Hawaiian Islands. Wader Study Group Bulletin 31:45–46. Close ), attesting to the adaptability of this species in urban environments. Notably, this behavior also has been observed in the European Golden-Plover (Pluvialis apricaria) during the nonbreeding season at several sites in England (236 Calbrade, N., C. A. Entwistle, A. J. Smith, and K. G. Spencer (2001). Roof assemblies of lapwings and plovers in Britain. British Birds 94:35–38. Close ). In contrast, the American Golden-Plover has not adapted to urban environments on the overwintering grounds, and there are no reports of this species using rooftops.

Daily Time Budget

No detailed information.

Territorial Behavior

Breeding Grounds

Territories are established and defended with ground and aerial displays, vocalizations, chases, and fights. Their Complex Whistle (see Sounds: Vocalizations) is a frequent warning from the ground as intruders fly by. Males advertise with wide-ranging Butterfly Display flights (171 Drury, W. H., Jr. (1961). The breeding biology of shorebirds on Bylot Island, Northwest Territories, Canada. Auk 78:176–219. Close ) while vocalizing their Repetitive Call (see Sounds: Vocalizations). American Golden-Plover flight uses slow, measured wingbeats, with wings almost vertical at the top of the stroke, performed at heights of 10–100 m above the ground (J. T. Nichols in 203 Bent, A. C. (1929). Life histories of North American shorebirds, Part 2. United States National Museum Bulletin 146. Close , 171 Drury, W. H., Jr. (1961). The breeding biology of shorebirds on Bylot Island, Northwest Territories, Canada. Auk 78:176–219. Close , 46 Parmelee, D. F., H. A. Stephens, and R. H. Schmidt (1967). The birds of Southeastern Victoria Island and adjacent small islands. National Museum of Canada Bulletin 222. Close , 37 Connors, P. G., B. J. McCaffery, and J. L. Maron (1993). Speciation in golden-plovers, Pluvialis dominica and P. fulva: evidence from the breeding grounds. Auk 110:9–20. Close , 31 Byrkjedal, I., and D. B. A. Thompson (1998). Tundra Plovers: The Eurasian, Pacific and American Golden Plovers and Grey Plover. T & AD Poyser Ltd, London, United Kingdom. Close ). Usually Butterfly Display flight commences with normal wingbeat as a plover departs from the ground silently. When the bird reaches display altitude, the wingbeat changes, and calling begins. According to Byrkjedal and Thompson (31 Byrkjedal, I., and D. B. A. Thompson (1998). Tundra Plovers: The Eurasian, Pacific and American Golden Plovers and Grey Plover. T & AD Poyser Ltd, London, United Kingdom. Close ), the Pacific Golden-Plover butterfly wingstroke is deeper (“wings sometimes seem almost to meet above the back and below the belly”) than that of the American Golden-Plover. Butterfly Display flights may cross territories of other males, who often respond with similar flight. Flights end in rapid, nearly vertical descent with wings held in a 'V' above the back; Complex Whistles are frequent as the bird nears the ground.

Neighboring males perform coordinated parallel marches and often vocalize with Complex Whistles. Whether these behaviors define territorial boundaries or occur in zones near potential nest sites is unknown (O. W. Johnson, P. G. Connors). More vigorous ground actions involve running toward and chasing intruders - the pursuer with his head down, back horizontal, back feathers ruffled, and breast and side feathers fluffed out. Accompanying vocalizations typically include the Complex Whistle (see Sounds: Vocalizations).

Both sexes, but especially the male, conduct aerial chases. Male chases may be brief and direct, or long (2+ min) and meandering, with birds 1–3 m apart (occasionally making physical contact) and 1–4 m above the ground (P. G. Connors), though sometimes at much greater heights (171 Drury, W. H., Jr. (1961). The breeding biology of shorebirds on Bylot Island, Northwest Territories, Canada. Auk 78:176–219. Close ). Their flight is rapid, with quick turns and other erratic maneuvers. Chase flights are almost always accompanied by specific aggressive vocalization (see Sounds: Vocalizations). The pursuer in an aerial chase is frequently an incubating bird disturbed by an intruder either landing or flying nearby. Aerial chases are often followed by male combatant Butterfly Display flights.

The foregoing aggressive behaviors occasionally escalate to contact fighting, with birds trying to peck at the feathers, head, or legs of the opponent. These attempts occur on the ground, while jumping and fluttering with or above an opponent, or in short flights directed at an adversary on the ground (O. W. Johnson, P. G. Connors) and accompanied by frequent Complex Whistles. During the on-ground phases of these skirmishes, both wings are often held vertically over the back. Where they are sympatric on the Seward Peninsula, Alaska, American Golden-Plovers and Pacific Golden-Plovers appear to be equally aggressive toward neighbors or intruding individuals, no matter the species (37 Connors, P. G., B. J. McCaffery, and J. L. Maron (1993). Speciation in golden-plovers, Pluvialis dominica and P. fulva: evidence from the breeding grounds. Auk 110:9–20. Close ). Prolonged disputes sometimes occur: an interspecific bout between two males lasted 105 min and involved parallel marching, aerial chases, contact fights, and repeated displays and vocalizations (37 Connors, P. G., B. J. McCaffery, and J. L. Maron (1993). Speciation in golden-plovers, Pluvialis dominica and P. fulva: evidence from the breeding grounds. Auk 110:9–20. Close ). Lengthy interactions are probably most frequent in early spring when territories are being established, and these may be a more common occurrence than presently recognized.

Breeding territories are defended against other bird species. A partial list includes aggression toward the Dunlin (Calidris alpina), Rock Sandpiper (Calidris ptilocnemis), Wilson's Snipe (Gallinago delicata), Black-bellied Plover (Pluvialis squatarola), Western Sandpiper (Calidris mauri), and Lapland Longspur (Calcarius lapponicus) (37 Connors, P. G., B. J. McCaffery, and J. L. Maron (1993). Speciation in golden-plovers, Pluvialis dominica and P. fulva: evidence from the breeding grounds. Auk 110:9–20. Close , O. W. Johnson, P. G. Connors; P. Bruner, personal communication). For additional comments about intraspecific and interspecific agonistic behaviors, see Byrkjedal and Thompson (31 Byrkjedal, I., and D. B. A. Thompson (1998). Tundra Plovers: The Eurasian, Pacific and American Golden Plovers and Grey Plover. T & AD Poyser Ltd, London, United Kingdom. Close ).

Nonbreeding Grounds

Territorial defense is best known in the Pacific Golden-Plover (137 Johnson, O. W., P. G. Connors, and P. Pyle (2018). Pacific Golden-Plover (Pluvialis fulva), version 3.0. In The Birds of North America (P. G. Rodewald, Editor). Cornell Lab of Ornithology, Ithaca, New York, USA. Close ), but is likely similar in the American Golden-Plover. Among overwintering Pacific Golden-Plovers, territorial individuals are intolerant of each other and often highly aggressive; nonterritorial birds forage communally, but maintain spacing through low-intensity aggression. Agonistic interactions are frequent at nighttime roosts. Presumably, similar behaviors occur among overwintering American Golden-Plovers, but details are lacking. In Argentina, the American Golden-Plover is territorial on grasslands and inland wetlands. Birds occupy their territories most of the day, except for mid afternoon, when territorial and nonterritorial birds merge into large flocks for drinking and bathing at local water sources (147 Myers, J. P., and L. P. Myers (1979). Shorebirds of coastal Buenos Aires Province, Argentina. Ibis 121:186–200. Close ). Territories of the Buff-breasted Sandpiper (Calidris subruficollis) overwintering in the same region are superimposed on those of the American Golden-Plover, with neither species showing interspecific territoriality (237 Myers, J. P. (1980b). Territoriality and flocking by Buff-breasted Sandpipers: variations in non-breeding dispersion. Condor 82:241–250. Close ). By contrast, Pacific Golden-Plovers in Hawaii do not readily share territorial space and frequently attack other bird species (O. W. Johnson). For reviews of nonbreeding season behaviors in plovers and other shorebirds see 238 Myers, J. P., P. G. Conners, and F. A. Pitelka (1979a). Territoriality in non-breeding shorebirds. Studies in Avian Biology 2:231–246. Close , 239 Colwell, M. A. (2000). A review of territoriality in non-breeding shorebirds (Charadrii). Wader Study Group Bulletin 93:58–66. Close .

Mating System

The American Golden-Plover is socially monogamous, with pairs remaining together for the breeding season and for replacement nesting (see Demography and Populations: Measures of Breeding Activity). Both sexes defend territory (the male with greater vigor), incubate, and tend to the young prior to fledging (see Breeding: Parental Care). Studies of banded birds on Seward Peninsula indicate that re-mating in subsequent years is much less likely than new pairing, despite the high site fidelity of males (134 Johnson, O. W., P. M. Johnson, P. L. Bruner, A. E. Bruner, R. J. Kienholz, and P. A. Brusseau (1997c). Male-biased breeding ground fidelity and longevity in American Golden-Plovers. Wilson Bulletin 109:348–351. Close , 135 Johnson, O. W., P. L. Bruner, A. E. Bruner, P. M. Johnson, R. J. Kienholz, and P. A. Brusseau (2001e). Features of breeding biology in Pacific and American Golden-Plovers nesting on the Seward Peninsula, Alaska. Wader Study Group Bulletin 95:59–65. Close ; see Demography and Populations: Range).

Courtship, Copulation, and Pair Bond

The male Butterfly Display flight clearly advertises the territory and may also serve to attract a female (72 Miller, E. H. (1996a). Acoustic differentiation and speciation in shorebirds. In Ecology and Evolution of Acoustic Communication in Birds (D. E. Kroodsma and E. H. Miller, Editors). Cornell University Press, Ithaca, NY, USA. pp. 241–257. Close , 31 Byrkjedal, I., and D. B. A. Thompson (1998). Tundra Plovers: The Eurasian, Pacific and American Golden Plovers and Grey Plover. T & AD Poyser Ltd, London, United Kingdom. Close , OWJ, PGC). Males sometimes perform this flight above or alongside a flying female (PGC). These flights often end with both birds landing, wings held aloft in a V while calling with Complex Whistles.

Several other displays (171 Drury, W. H., Jr. (1961). The breeding biology of shorebirds on Bylot Island, Northwest Territories, Canada. Auk 78:176–219. Close , 46 Parmelee, D. F., H. A. Stephens, and R. H. Schmidt (1967). The birds of Southeastern Victoria Island and adjacent small islands. National Museum of Canada Bulletin 222. Close , O. W. Johnson, P. G. Connors) also appear to function in pair-bonding and/or as prerequisites to copulation. Five are described briefly here: (1) on ground performance by both sexes of Complex Whistles; (2) Scraping Display, where the male uses his breast and feet (and probably bill) to form a nest scrape on tundra or occasionally on snow, often picking up bits of vegetation during display; (3) Wing-Stretch, in which the male stretches his wings vertically, head held low; (4) Torpedo Posture of the male with bill, head, and back horizontal, back feathers usually ruffled, wings sometimes raised; here the male either remains stationary or runs toward the female; (5) Erect Posture, in which the male stands very upright, neck stretched upward, motionless for up to 30 s, the female close and observant. The order and relationships of displays is not understood. Copulation usually lasts only one to four seconds and follows a brief pursuit of the female.

Extra-Pair Mating Behavior

This appears to occur at a relatively high rate in the American Golden-Plover. Of 131 offspring sampled at Utqiagvik, Alaska, eight percent were products of non-monogamous matings (240 Yezerinac, S., R. B. Lanctot, G. K. Sage, and S. L. Talbot (2013). Social and genetic mating system of the American Golden-Plover. Condor 115:808–815. Close ). Presumably, extra-pair copulations are associated with the habit of females to forage at considerable distance from the nest when not incubating (see Breeding: Parental Behavior). A sperm length of 67.2 µm (as index to sperm competition) in the American Golden-Plover is similar to the sperm length of other shorebirds considered as monogamous (241 Johnson, D. D. P., and J. V. Briskie (1999). Sperm competition and sperm length in shorebirds. Condor 101:848–854. Close ).

Flocking behavior is characteristic of pre-migrants and migrants. There is no evidence of either intraspecific or interspecific sociality between the American Golden-Plover and Pacific Golden-Plover during the breeding season, except birds are somewhat gregarious on extraterritorial foraging areas (O. W. Johnson). Breeding American Golden-Plovers in Churchill, Manitoba, were relatively tolerant toward the Short-billed Dowitcher (Limnodromus griseus); association with attentive, vigilant plovers may benefit dowitchers (242 Byrkjedal, I. (1987). Short-billed Dowitchers associate closely with Lesser Golden-Plovers. Wilson Bulletin 99:494–495. Close ).

Kinds of Predators and Manner of Predation

The American Golden-Plover is taken by a variety of avian and mammalian predators on breeding and non-breeding grounds. Most significant losses are probably eggs and young (see Demography and Populations: Causes of Mortality).

Response to Predators

Breeding Grounds

There is no “acceptable information on the escape tactics of plovers” when attacked by raptors (243 Lima, S. L. (1993). Ecological and evolutionary perspectives on escape from predatory attack: A survey of North American birds. Wilson Bulletin 105:1–47. Close ). Various behavioral responses occur at or near the nest (244 Byrkjedal, I. (1989a). Nest defense behavior of Lesser Golden-Plovers. Wilson Bulletin 101:579–590. Close , O. W. Johnson, P. G. Connors). These include: (1) sit tight and sink more deeply into the nest; (2) flatten with head outstretched, body motionless; (3) fly from the nest and stand silently some distance away; (4) sneak away from the nest silently; (5) depart noisily from the nest while the predator is still at a considerable distance; (6) sit tight until the intruder is relatively close; (7) perform specialized distraction displays to lure the predator away; (8) attack/mob the predator with aggressive aerial maneuvers.

In Churchill, Manitoba, the American Golden-Plover reacted to a Herring Gull (Larus argentatus) with behavior 1, to the Parasitic Jaeger (Stercorarius parasiticus), and Northern Shrike (Lanius borealis) with behavior 2, and to the Northern Harrier (Circus hudsonius) with behavior 3 (244 Byrkjedal, I. (1989a). Nest defense behavior of Lesser Golden-Plovers. Wilson Bulletin 101:579–590. Close ). On the Seward Peninsula, incubating plovers reacted to the Long-tailed Jaeger (Stercorarius longicaudus) with behavior 1 (P. Bruner, personal communication).

Nesting plovers apparently respond to predators with mostly non-aggressive behaviors (1–7), but a situation sometimes triggers an attack (behavior 8) (171 Drury, W. H., Jr. (1961). The breeding biology of shorebirds on Bylot Island, Northwest Territories, Canada. Auk 78:176–219. Close , 245 Sordahl, T. A. (1981). Predator-mobbing behaviour in the shorebirds of North America. Wader Study Group Bulletin 31:41–44. Close , 246 McCaffery, B. J. (1982). Geographic variation in predator-mobbing behavior-a hypothesis. Wader Study Group Bulletin 35:30. Close ). McCaffery’s account of an interaction between an American Golden-Plover and Long-tailed Jaeger attests to potential ferocity of these encounters: the Long-tailed Jaeger attempting to rob a plover nest grabbed the attacking plover with its bill and began flying off with it; the plover struggled free and continued its aerial attack on the jaeger; despite the plover's efforts, the jaeger eventually succeeded in stealing an egg. Attack response by plovers is unlikely in places where mobbing by another species wards off predators, as with aggressive Whimbrels (Numenius phaeopus) in the Churchill region (244 Byrkjedal, I. (1989a). Nest defense behavior of Lesser Golden-Plovers. Wilson Bulletin 101:579–590. Close ). Plovers occasionally attack/mob mammals, including red foxes (Vulpes vulpes fulvus; Seward Peninsula, O. W. Johnson) and caribou (Rangifer tarandus, northern Alaska, 122 Miller, P. A., C. S. Moitoret, and M. A. Masteller (1985b). Species accounts of migratory birds at three study areas on the coastal plain of the Arctic National Wildlife Refuge Alaska, 1984. U.S. Fish and Wildlife Service, Fairbanks, AK, USA. Close ).

The response to ground predators often involves conspicuous, noisy actions (behavior 7) to confuse and distract the intruder. Alarm/Distraction Calls (see Sounds: Vocalizations) are given after the bird leaves the nest. Calling is often very agitated. A human generally evokes the same noisy behaviors as ground predators (244 Byrkjedal, I. (1989a). Nest defense behavior of Lesser Golden-Plovers. Wilson Bulletin 101:579–590. Close ), but some birds are extremely wary and leave the nest stealthily (behavior 4), often ≥ 200 m in advance of the observer ([“early surreptitious departure” 247 Gochfeld, M. (1984). Antipredator behavior: aggressive and distraction displays of shorebirds. In Behavior of Marine Animals, Volume 5. Shorebirds: Breeding Behavior and Populations (J. Burger and B. L. Olla, Editors). Plenum Press, New York, NY, USA. pp. 289–377. Close ]; O. W. Johnson). After sneaking well clear of the nest, the bird either walks or flies far enough to disappear over a hill, ridge, etc., or remains in view at a considerable distance, calling and/or foraging, occasionally performing distant distraction displays. Stealthy departure is especially characteristic of the Pacific Golden-Plover. Human/plover interaction progresses only to distraction behavior; an observer at a nest does not provoke attack. However, Sauer (227 Sauer, E. G. F. (1962). Ethology and ecology of golden plovers on St. Lawrence Island, Bering Sea. Psychologische Forschung 26:399–470. Close ) recorded an unusual instance in which Pacific Golden-Plovers did attack him by “diving sharply . . . and flying in low tight circles”. This a-typical situation was probably linked to a territorial dispute between the plovers and Ruddy Turnstone (Arenaria interpres) that was occurring at the same time.

Distraction displays are similar in both the American Golden-Plover and Pacific Golden-Plover (248 Murie, A. (1946). Observation on the birds of Mount McKinley National Park, Alaska. Condor 48:253–261. Close , 171 Drury, W. H., Jr. (1961). The breeding biology of shorebirds on Bylot Island, Northwest Territories, Canada. Auk 78:176–219. Close , 227 Sauer, E. G. F. (1962). Ethology and ecology of golden plovers on St. Lawrence Island, Bering Sea. Psychologische Forschung 26:399–470. Close , 46 Parmelee, D. F., H. A. Stephens, and R. H. Schmidt (1967). The birds of Southeastern Victoria Island and adjacent small islands. National Museum of Canada Bulletin 222. Close , 244 Byrkjedal, I. (1989a). Nest defense behavior of Lesser Golden-Plovers. Wilson Bulletin 101:579–590. Close , O. W. Johnson, P. G. Connors). Repertoires include: (1) Tail-Down Run: head low, tail depressed, plumage not ruffled; (2) Rodent-Run: bird crouched, plumage fluffed, wings partially outstretched, drooped and quivering, tail down usually dragging on ground; (3) Injury Feigning: one or both wings extended and flapping as if unable to fly (performed during a slow run, or while creeping with wings beating on the ground as if “rowing,” or in a stationary position, either standing or prostrate); (4) Spread-Wing Display: bird crouched or prostrate, facing intruder, wings outstretched and motionless, tail fanned and either erect or depressed; and (5) False Brooding: rarely, a disturbed bird will sit near intruder as if on another nest. Distractions are spirited and varied; displays rapidly transition from one to another. Three of these displays are shown in Figure 4.

Plaintive Alarm/Distraction Calls are mostly absent during distraction displays (244 Byrkjedal, I. (1989a). Nest defense behavior of Lesser Golden-Plovers. Wilson Bulletin 101:579–590. Close , O. W. Johnson), but do occur during brief pauses in the latter. Males typically give more vigorous distractions than females. On the Seward Peninsula, males often perform impressively over extended periods and may approach an observer closely to within ≤ 1.0 m. The male Pacific Golden-Plover behaves similarly, but is often more wary, tending to perform at a greater distance and for a shorter period, and sometimes will cease the display to walk or fly away (O. W. Johnson).

When an observer locates a nest and does not follow the displaying bird away from it, the plover often returns and initiates a new display (“re-entrapment” of the intruder; 247 Gochfeld, M. (1984). Antipredator behavior: aggressive and distraction displays of shorebirds. In Behavior of Marine Animals, Volume 5. Shorebirds: Breeding Behavior and Populations (J. Burger and B. L. Olla, Editors). Plenum Press, New York, NY, USA. pp. 289–377. Close ). This may occur several times; thereafter, the tendency is for the bird to remain aloof and alarm-call, sometimes calling alternates with displays performed at a considerable distance from the observer. If the observer now stays motionless at the nest, the bird often ceases displays but continues alarm-calling while either standing or running about, occasionally pausing to quickly snatch up an insect or other item of prey. Movements by the intruder (such as passing one's hand over the nest) frequently will draw the bird back to the nest and trigger another bout of distraction behaviors (O. W. Johnson). Distractions are performed mostly by a disturbed, incubating bird. The off-duty mate (especially the male likely to be within earshot) usually returns to the agitated partner, sometimes enters into displays, but may simply watch and call excitedly. The hypothesis that motivational conflicts between escape, aggression, and incubation govern distraction behavior was rejected in studies of the American Golden-Plover (249 Byrkjedal, I. (1991). The role of drive conflicts as a mechanism for nest-protection behaviour in the shorebird Pluvialis dominica. Ethology 87:149–159. Close ). His findings suggest adaptive choice rather than mechanical response. The responses of nesting plovers to human intruders are further described along with drawings (244 Byrkjedal, I. (1989a). Nest defense behavior of Lesser Golden-Plovers. Wilson Bulletin 101:579–590. Close , 31 Byrkjedal, I., and D. B. A. Thompson (1998). Tundra Plovers: The Eurasian, Pacific and American Golden Plovers and Grey Plover. T & AD Poyser Ltd, London, United Kingdom. Close ).

Overwintering Grounds

There is relatively little information. The presence of raptors caused American Golden-Plovers in Argentina to sound alarm calls (237 Myers, J. P. (1980b). Territoriality and flocking by Buff-breasted Sandpipers: variations in non-breeding dispersion. Condor 82:241–250. Close ).