American Golden-Plover

Pluvialis dominica

Order:
Charadriiformes
Family:
Charadriidae
Sections

Behavior

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Several American Golden-Plovers and a Southern Lapwing (Vanellus chilensis).

During the nonbreeding season, may join other shorebirds on multi-species roosts.

© Jerome Foster , Tunapuna-Piarco , Trinidad and Tobago , 30 September 2017
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Figure 4. Distraction displays of the American Golden-Plover.

Some distraction displays of the American Golden-Plover: (top) Tail-Down Run, (middle) Injury Feigning, and (bottom) Stationary Wing-Spread Display. The same behaviors occur in Pacific Golden-Plovers. Drawings by D. Otte, from photographs by OWJ (included below).

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Alternate (breeding) male American Golden-Plover in Tail-Down Run display.

In this distraction display, the head is held low, the tail depressed, and plumage is not ruffled. Photographer Oscar Johnson.

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Alternate (breeding) male American Golden-Plover in Injury Feigning display.

In this distraction display, one or both wings are extended and flapping as if unable to fly (performed during slow run, or while creeping with wings beating on ground as if “rowing,” or in stationary position either standing or prostrate). Photographer Oscar Johnson.

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Alternate (breeding) male American Golden-Plover giving Stationary Wing-Spread Display.

Stationary Wing-Spread Display is one element in a series of behaviors exhibited when adult is disturbed from nest. In this distraction display, bird is crouched or prostrate, facing intruder, with wings outstretched and motionless, tail fanned and either erect or depressed. Photographer Oscar Johnson.

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Merlin (Falco columbarius) with American Golden-Plover prey.

American Golden-Plover is prey for a variety of avian and mammalian predators.

© Ted Keyel , Minnesota , United States , 15 September 2013

Locomotion

Walking, Running, Hopping

Commonly walk and run. Often stand on one leg while loafing or roosting; if disturbed, may hop on that leg for considerable distance before changing gait. Except with certain topographic features of tundra landscape (see Sounds: Vocalizations), locomotion does not involve elevated perches.

Flight

Capable of swift and extended flight. These plovers are considered "the high speed champions among shorebirds" (6). Numerous records of Pacific Golden-Plovers equipped with geolocators indicate average ground speed around 50 kph during long transoceanic flights, with considerable variation (38 to > 100 kph) presumably influenced by direction of winds (222, 223, 172, 224). Presumably, flight speeds of American Golden-Plovers 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) often > 72 kph ranging as high as 136 kph (162, 158, 159, 116, 160, 161). Mean ground speed of a single American Golden-Plover tracked between Illinois and Iowa was 56 kph (60).

Swimming and Diving

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 225), fatigue, or injury.

Self-Maintenance

Preening, Head-Scratching, Stretching, Bathing, Anting

Preening and stretching usually accompany periods of loafing. Head-scratching is indirect, foot passed over lowered wing (OWJ). 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 (121). No reports of bathing on nesting grounds. 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 head resting on shoulders and bill forward or with head turned and bill tucked beneath scapulars. Presumably, American Golden-Plover is similar, but no reports. Brief intervals of sleep likely on breeding grounds, but no information available.

Roosts in flocks during the nonbreeding season. In Argentina, American Golden-Plover joins other shorebirds on multispecific nocturnal roosts "at the edge of lagoons and swamps" (226). Rooftop roosting by Pacific Golden-Plovers occurs in Hawaii and elsewhere on the overwintering range (221), attesting to the adaptability of this species in urban environments. Notably, this behavior also has been observed in the Eurasian Golden-Plover during the nonbreeding season at several sites in England (227). 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.

Spacing

Breeding Territoriality

Highly territorial on the breeding grounds. Territory sizes estimated at roughly 25 ha on North Slope of Alaska, 10–50 ha on Seward Peninsula (PGC). Most pair activity is focused on the territory, but considerable foraging occurs elsewhere as the non-incubating bird (especially female) is often absent from territory (142, OWJ, PGC; see Breeding: Incubation). Extraterritorial feeding probably of a communal nature on specific sites (OWJ). Interyear fidelity to breeding territory high in males, low in females (see Demography and Populations: Range).

Territories established and defended with ground and aerial displays, vocalizations, chases, fights. Complex Whistle (see Sounds: Vocalizations) a frequent warning from ground as intruders fly by. Males advertise with wide-ranging Butterfly Display flight (142), while vocalizing Repetitive Call (see Sounds: Vocalizations). Flight uses slow, measured wingbeats, with wings almost vertical at top of stroke; performed at heights of 10–100 m above ground (J. T. Nichols in 194, 142, 44, 37, 31). Usually Butterfly Display flight commences with normal wingbeat as plover departs ground silently; when bird reaches display altitude, wingbeat changes and calling begins. According to Byrkjedal and Thompson (31), Pacific Golden-Plover butterfly wing-stroke is deeper (“wings sometimes seem almost to meet above the back and below the belly”) than that of 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 back; Complex Whistles frequent as bird nears ground.

Neighboring males perform coordinated parallel marches, often vocalize with Complex Whistles; whether these behaviors define territorial boundary or occur in zone near potential nest site is unknown (OWJ, PGC). More vigorous ground actions involve running toward and chasing intruder, pursuer with head down, back horizontal, back feathers ruffled, breast and side feathers fluffed out; accompanying vocalizations typically the Complex Whistle (see Sounds: Vocalizations).

Both sexes, but especially 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 ground (PGC); sometimes at much greater heights (142). Flight is rapid with quick turns and other erratic maneuvers. Chase flights almost always with specific aggressive vocalization (see Sounds: Vocalizations). Pursuer in aerial chase is frequently an incubating bird disturbed by intruder either landing or flying nearby. Aerial chases often followed by Butterfly Display flights of male combatants.

The foregoing aggressive behaviors occasionally escalate to contact fighting, with birds trying to peck at feathers, head, or legs of opponent. These attempts occur on the ground, while jumping and fluttering with or above opponent, or in short flights directed at adversary on the ground (OWJ, PGC); accompanied by frequent Complex Whistles; during on-ground phases of these skirmishes, both wings often held vertically over back. Where sympatric on Seward Peninsula, Alaska, American and Pacific golden-plovers appear to be equally aggressive toward neighbors or intruding individuals, no matter the species (37). Prolonged disputes sometimes occur: an interspecific bout between 2 males lasted 105 min and involved parallel marching, aerial chases, contact fights, and repeated displays and vocalizations (37). Lengthy interactions are probably most frequent in early spring when territories are being established, and may be a more common occurrence than presently recognized.

Breeding territories are defended against other bird species. Partial list: aggressive toward Dunlin (Calidris alpina), Rock Sandpiper (Calidris ptilocnemis), Wilson's Snipe (Gallinago delicata), Black-bellied Plover, Western Sandpiper (Calidris mauri), and Lapland Longspur (Calcarius lapponicus) (37, OWJ, PGC; P. Bruner, personal communication). For additional comments about intraspecific and interspecific agonistic behaviors, see Byrkjedal and Thompson (31).

Overwintering Territoriality and Nonterritoriality

Many individuals defend feeding territories on wintering grounds, some are nonterritorial. Territory sizes are much smaller than on breeding grounds, ≤ 0.3 ha in Argentina (121). Territorial defense is best known in Pacific Golden-Plover (16), but is likely similar in 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; and 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 (121). Territories of Buff-breasted Sandpipers (Calidris subruficollis) overwintering in the same region are superimposed on those of American Golden-Plovers, with neither species showing interspecific territoriality (228). By contrast, Pacific Golden-Plovers in Hawaii do not readily share territorial space and frequently attack other bird species (OWJ). For reviews of nonbreeding season behaviors in plovers and other shorebirds see 229, 230.

Sexual Behavior

Mating System

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 (male with greater vigor), incubate, and tend 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 high site fidelity of males (231, 232; see Demography and Populations: Range).

Pair Bond

The male Butterfly Display flight clearly advertises the territory and may also serve to attract a female (70, 31, 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 (142, 44, OWJ, PGC) 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 male uses breast and feet (and probably bill) to form nest scrape on tundra or occasionally on snow, often picking up bits of vegetation during display; (3) Wing-Stretch, in which male stretches wings vertically, head held low; (4) Torpedo Posture of male with bill, head, and back horizontal, back feathers usually ruffled, wings sometimes raised; either remains stationary or runs toward female; (5) Erect Posture in which male stands very upright, neck stretched upward, motionless for up to 30 s, female close and observant. Order and relationships of displays not understood. Copulation usually lasts only 1–4 s, follows brief pursuit of female.

Extra-Pair Copulations

Appears to occur at relatively high rate in American Golden-Plover. Of 131 offspring sampled at Barrow, Alaska, 8% were products of non-monogamous matings (233). Presumably, extra-pair copulations are associated with the habit of females to forage at considerable distance from nest when not incubating (see Breeding: Parental Behavior). Sperm length of 67.2 µm (as index to sperm competition) in American Golden-Plover similar to sperm length of other shorebirds considered as monogamous (234).

Social and Interspecific Behavior

Flocking behavior is characteristic of pre-migrants and migrants. No evidence of either intraspecific or interspecific sociality between American and Pacific golden-plovers during the breeding season, except birds are somewhat gregarious on extraterritorial foraging areas (OWJ). Breeding American Golden-Plovers at Churchill, Manitoba, were relatively tolerant toward Short-billed Dowitchers (Limnodromus griseus); association with attentive, vigilant plovers may benefit dowitchers (235).

Predation

Kinds of Predators and Manner of Predation

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. No “acceptable information on the escape tactics of plovers” when attacked by raptors (236). Various behavioral responses occur at or near nest (237, OWJ, PGC). These include: (1) sit tight and sink more deeply into nest; (2) flatten with head outstretched, body motionless; (3) fly from nest and stand silently some distance away; (4) sneak away from nest silently; (5) depart noisily from nest while predator is still at considerable distance; (6) sit tight until intruder is relatively close; (7) perform specialized distraction displays to lure predator away; (8) attack/mob the predator with aggressive aerial maneuvers.

At Churchill, Manitoba, American Golden-Plovers reacted to Herring Gull (Larus argentatus) with behavior 1, to Parasitic Jaeger (Stercorarius parasiticus) and Northern Shrike (Lanius borealis) with behavior 2, to Northern Harrier (Circus hudsonius) with behavior 3 (237). On the Seward Peninsula, incubating plovers reacted to 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 situation sometimes triggers attack (behavior 8) (142, 238, 239). McCaffery’s account of an interaction between American Golden-Plover and Long-tailed Jaeger attests to potential ferocity of these encounters: jaeger attempting to rob plover nest grabbed attacking plover with its bill and began flying off with it; plover struggled free and continued aerial attack of jaeger; despite plover's efforts, 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 (237). Plovers occasionally attack/mob mammals, including Red Fox (Vulpes vulpes fulvus; Seward Peninsula, OWJ), Caribou (Rangifer tarandus, northern Alaska, 175).

Response to ground predator often involves conspicuous, noisy actions (behavior 7) to confuse and distract the intruder. Alarm/Distraction Calls (see Sounds: Vocalizations) are given after bird leaves the nest. Calling is often very agitated. Human generally evokes same noisy behaviors as ground predators (237), but some birds are extremely wary and leave nest stealthily (behavior 4), often ≥ 200 m in advance of observer ([“early surreptitious departure” 240]; OWJ). After sneaking well clear of nest, bird either walks or flies far enough to disappear over hill, ridge, etc., or remains in view at considerable distance calling and/or foraging, occasionally performing distant distraction displays. Stealthy departure especially characteristic of Pacific Golden-Plover. Human/plover interaction progresses only to distraction behavior; observer at nest does not provoke attack. However, Sauer (218) recorded an unusual instance in which Pacific Golden-Plovers did attack him by “diving sharply . . . and flying in low tight circles”. This atypical situation was probably linked to a territorial dispute between the plovers and Ruddy Turnstones (Arenaria interpres) that was occurring at the same time.

Distraction displays are similar in both American and Pacific golden-plovers (241, 142, 218, 44, 237, OWJ, PGC). Repertoires include: Tail-Down Run—head low, tail depressed, plumage not ruffled; Rodent-Run—bird crouched, plumage fluffed, wings partially outstretched, drooped and quivering, tail down usually dragging on ground; Injury Feigning—one or both wings extended and flapping as if unable to fly (performed during slow run, or while creeping with wings beating on ground as if “rowing,” or in stationary position either standing or prostrate); Spread-Wing Display—bird crouched or prostrate, facing intruder, wings outstretched and motionless, tail fanned and either erect or depressed; 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 (237, OWJ), but do occur during brief pauses in the latter. Males typically give more vigorous distractions than females. On Seward Peninsula, males often perform impressively over extended periods and may approach observer closely to within ≤ 1.0 m. Male Pacific Golden-Plovers behave similarly, but many are more wary, tending to perform at greater distance and for shorter period, sometimes cease display and walk or fly away (OWJ).

When observer locates a nest and does not follow displaying bird away from it, plover often returns and initiates new display (“re-entrapment” of the intruder; 240). This may occur several times, thereafter tendency is for bird to remain aloof and alarm-call, sometimes calling alternates with displays performed at considerable distance from observer. If observer now stays motionless at the nest, 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 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 (OWJ). Distractions performed mostly by disturbed, incubating bird. Off-duty mate (especially male likely to be within earshot) usually returns to 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 American Golden-Plover (242). His findings suggest adaptive choice rather than mechanical response. The responses of nesting plovers to human intruders are further described along with drawings (237, 31).

Overwintering Grounds. Relatively little information. Presence of raptors caused American Golden-Plovers in Argentina to sound alarm calls (228).

Recommended Citation

Johnson, O. W., P. G. Connors, and P. Pyle (2019). American Golden-Plover (Pluvialis dominica), version 3.1. In The Birds of North America (P. G. Rodewald, Editor). Cornell Lab of Ornithology, Ithaca, NY, USA. https://doi.org/10.2173/bna.amgplo.03.1