Pacific Golden-Plover Pluvialis fulva Scientific name definitions
Version: 1.1 — Published April 15, 2021
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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. Three atypical situations recorded on Oahu, Hawaii: group of Pacific Golden-Plover flushed from pond alighted on dead branches of surrounding trees and remained there for several minutes, 3 birds (separate instances) seen standing on upper rail of chain-link fences (P. Bruner and A. Bruner, personal communication; OWJ); 1 bird observed walking and foraging atop a dense, pruned hedge approximately 0.5 m tall (OWJ), hedge was part of the individual’s overwintering territory.
Capable of swift and extended flight. These plovers are considered "the high speed champions among shorebirds" (13). Numerous records from Pacific Golden-Plover 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 (98, 102, 179, 180). According to radar measurements in various arctic regions, ground speeds of migrating shorebird flocks (some composed of Pacific Golden-Plover or American Golden-Plover) often > 72 kph ranging as high as 136 kph (236, 231, 232, 233, 234, 235). Mean ground speed of migrants (likely including Pacific Golden-Plover) passing Oahu was 63 kph (284, 285), similar measurement at Guam 60.7 kph (173).
Swimming and Diving
Not an aquatic species (47). In the above geolocator studies there was no evidence of seawater contact during migrations, thus Pacific Golden-Plover are able to remain aloft for extended periods. 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 286), fatigue, or injury.
Preening, Head-Scratching, Stretching, Bathing, Anting
Preening and stretching usually accompany periods of loafing. Head-scratching is indirect, foot passed over lowered wing (47, OWJ). Bathing observed on overwintering grounds in Oahu, Hawaii (see Diet and Foraging: Drinking, Pellet-Casting, and Defecation), involves vigorous wing-flapping, body sometimes immersed in shallow water. On Maui, an overwintering plover (probably the same individual) was seen on several occasions bathing in a backyard birdbath (S. Southworth, personal communication), and on Kauai, a plover was seen bathing along shallow edge of a swimming pool (OWJ). No reports of bathing on nesting grounds or at migratory stopover sites. No records of anting.
Overwintering 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 (see 160).
Daytime roosting sites (sometimes in response to high tides) during non-breeding season include beaches, rocky points, spits, levees, tops of low mangroves (12, 287). In Hawaii, birds often aggregate at nighttime roosts on flat rooftops (283), rocky offshore islands (172), hillsides, parking lots (OWJ), and barren lava flows (275, P. Bruner in litt.). Observations in Hawaii (172, 283, OWJ) indicate: first birds appear at nocturnal roosts around sunset, with most arriving after dark; roosting flocks vary from a few to 300+ individuals; aggression common when spacing < about 1.5 m; aggressive interactions occur throughout the night; birds depart roosts before dawn and return to territories or communal foraging areas; nocturnal roosts occur annually on same rooftops; diurnal aggregations of loafing, sleeping, preening birds form on rooftops, large grassy areas, and wetlands, these individuals are usually more tolerant (spacing often ≤ 0.5 m) than birds at nocturnal roosts.
Rooftop roosting by Pacific Golden-Plover has been reported elsewhere on the overwintering range (see 283) attesting to the adaptability of this species in urban environments. Notably, this behavior also has been observed in European Golden-Plover during the non-breeding season at several sites in England (288). The latter birds roost on rooftops during the day, then depart to feeding areas at night (the reverse of the pattern in Hawaii). There are no reports of American Golden-Plover using rooftops.
Daily Time Budget
The only study that examined daily activity patterns involved overwintering Pacific Golden-Plover on Oahu, Hawaii; these birds spent majority (68%) of daylight hours foraging (A. Bruner, personal communication).
Highly territorial on breeding grounds. Territory sizes estimated at roughly: 25–50 ha on St. Lawrence Island, Alaska (47), 50 ha in eastern Siberia (289); 10–50 ha on Seward Peninsula, Alaska (PGC). Large size of territory produces a “gradient of defense” (47) in which distraction displays and defensive behaviors are particularly intense in zone around nest, less frequent peripherally. Most of pair activity focused on territory, but considerable foraging occurs elsewhere as non-incubating bird (especially female) is often absent from territory (47, OWJ, PGC; see Breeding: Incubation). Extraterritorial feeding probably of a communal nature on specific sites (47, OWJ). Densities usually c. 1 pair/km², locally up to 6–7 pairs/km²; can vary considerably between years. Interyear fidelity to breeding territory high in males, very low in females (see Movements and Migration: Dispersal and Site Fidelity).
Overwintering Territoriality and Non-territoriality
Many individuals defend feeding territories on overwintering grounds, while others are non-territorial. Territory sizes are much smaller than on breeding grounds. On Oahu, territories are often around 0.4–0.5 ha (172, 157, 159), or a fraction of this such as residential backyard. For reviews of these non-breeding season behaviors in plovers and other shorebirds, see 290, 291.
Other details concerning Pacific Golden-Plover in Hawaii (172, 18, OWJ): territories are held mostly by adults (some juveniles) of both sexes, males predominating in what appear to be the choicest habitats; locations of territories varied, lawns especially favored (see Habitat: Habitat in Nonbreeding Range); territorial birds defend same territories daily for entire overwintering period, typically retire to roosts at night, occasionally found on territories during night; individuals typically reoccupy the identical territory defended in the previous season (i.e., very high interyear fidelity, see Movements and Migration: Dispersal and Site Fidelity); nonterritorial birds also site faithful (but probably at lower rate than territorial birds) to same communal grounds annually; human activities may temporarily displace territorial plovers, which often join nonterritorial birds, sometimes forming large aggregations on areas safe from disturbance; plovers in Hawaii do not readily share territorial space and frequently attack other bird species (OWJ). Details concerning initial establishment of territory, whether this happens only in first season on overwintering grounds (see 292), are unknown.
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 Behavior). Studies of banded birds on Seward Peninsula indicate that remating in subsequent years is much less likely than new pairing, despite high site fidelity of males (293, 167, OWJ; see Movements and Migration: Dispersal and Site Fidelity).
Courtship, Copulation, and Pair Bond
The male Butterfly Display flight clearly advertises the territory and may also serve to attract a female (82, 37, OWJ, PGC). Males sometimes perform this flight above or alongside a flying female (47, PGC). These flights often end with both birds landing, wings held aloft in a 'V' while calling with Complex Whistles (see Sounds and Vocal Behavior: Vocalizations).
Several other displays similar in both Pacific Golden-Plover and American Golden-Plover (279, 47, 56, 37, 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 and Head-to-tail Rocking; (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; and (5) Tipping, in which male (possibly giving a quiet trill; 47) holds body rigid and straight but tipped forward with bill almost touching ground, tail high exposing undertail to female who sometimes sits on ground or in nest scrape. Order and relationships of displays not understood. Copulation follows brief pursuit of female; alternatively, male faces female, depresses tail, then mounts and copulates. Copulation is brief, usually lasting 1–4 s.
Extra-Pair Mating Behavior
Occurs in Pacific Golden-Plover (OWJ; R. Gold and R. Goodwill, unpublished data), rate uncertain. 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).
Social and Interspecific Behavior
Flocking characteristic of pre-migrants and migrants. No evidence of either intraspecific or interspecific sociality between Pacific Golden-Plover and American Golden-Plover during breeding season, except birds somewhat gregarious on extraterritorial foraging areas (OWJ). Breeding Pacific Golden-Plover relatively tolerant toward Dunlin (Calidris alpina). This interaction is beneficial for Dunlin and the details of this relationship (“paging” of plovers) are discussed by Byrkjedal and Thompson (37).
Among overwintering birds, territorial individuals are intolerant and often highly aggressive; nonterritorial birds forage communally but maintain spacing through low-intensity aggression. Most overwintering birds aggregate at nighttime roosts, where agonistic interactions are frequent.
Kinds of Predators and Manner of Predation
Taken by a variety of avian and mammalian predators on breeding and wintering grounds. Most significant losses 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 (294). Observations related to Sauer (47) by St. Lawrence Island resident suggest that pre-migrant Pacific Golden-Plover under attack from Gyrfalcons (Falco rusticolus) attempt to out-climb the predators.
Various behavioral responses (similar in both Pacific Golden-Plover and American Golden-Plover) of breeding birds at/or near nest have been observed (47, 295, OWJ, PGC) including: (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 a considerable distance; (6) sit tight until intruder is relatively close; (7) perform specialized distraction displays to lure predator away; and (8) attack/mob the predator.
According to Sauer (47), Pacific Golden-Plover on St. Lawrence Island responded to Parasitic Jaeger (Stercorarius parasiticus), Long-tailed Jaeger (Stercorarius longicaudus), Glaucous Gull (Larus hyperboreus), Black-legged Kittiwake (Rissa tridactyla), and Common Raven (Corvus corax) by either “crouching and remaining silent” (presumably equivalent to behaviors 1 and 2), or attacking with “fierce and swift aggressive flights” (behavior 8). Other Alaska observations: on Kuskokwin River Delta and Seward Peninsula male Pacific Golden-Plover were seen attacking Long-tailed Jaeger (296, P. Bruner in litt.); on Seward Peninsula foraging birds off duty and away from nests responded to Parasitic Jaeger and Long-tailed Jaegers with behavior 2 (OWJ).
Nesting plovers apparently respond to predators with mostly non-aggressive behaviors (1-7), but situation sometimes triggers attack (behavior 8). Attacks possibly more frequent in some populations. Possible factors accounting for variable responses among populations include: relative risk (e.g., if Pomarine Jaeger [Stercorarius pomarinus] predominates in the area, attacking or mobbing might be maladaptive), the presence of other mobbing species to ward off predators, and benefits of cryptic strategies associated with behaviors 1, 2, and 3 (279, 297, 298, 295). Pacific Golden-Plover on the Taimyr Peninsula were reported 5 times less likely to attack aerial predators than larger Black-bellied Plover (299).
Responses to ground predator generally involve conspicuous, noisy actions (behavior 7) to confuse and distract the intruder. Alarm/Distraction Calls (see Sounds and Vocal Behavior: Vocalizations) are after bird leaves the nest. Calling is often very agitated, and often punctuated with Head-to-tail Rocking. Human generally evokes same noisy behaviors as ground predators (47, 295), but some birds are extremely wary and leave nest stealthily (behavior 4), often ≥ 200 m in advance of observer ([“early surreptitious departure” 300]; 61, 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. Generally, these birds remain aloof for long periods even if nest found by intruder.
Distraction displays (47, 56, 295, OWJ, PGC) 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); Stationary Wing-Spread 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.
Plaintive Alarm/Distraction Calls are mostly absent during distraction displays (295, OWJ), but do occur during brief pauses in the latter. Males give more vigorous distractions than females. Sometimes bird will display for lengthy period and approach intruder to within ≤ 1.0 m, but many individuals perform at much greater distance and for shorter period; often cease display and walk or fly away.
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; 300). 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) frequently within earshot and returns to agitated partner, sometimes enters into displays, but often simply watches and calls excitedly. The responses of nesting plovers to human intruders are further described along with drawings (301, 37).
Pacific Golden-Plover attacking (behavior 8) arctic fox (Vulpes lagopus) on St. Lawrence Island were observed by Sauer (47). Sometimes Ruddy Turnstone entered into the plover/fox interaction: plovers attacked fox, agitation attracted turnstones who joined in mobbing fox, plovers then ignored fox and attacked turnstones, turnstones battled with both plovers and fox and finally drove fox from plover territory. Human/plover interaction progresses only to distraction behavior; observer at nest does not provoke attack. However, Sauer (47) recorded an unusual instance in which Pacific Golden-Plover 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 Turnstone that was occurring at the same time.
Overwintering Grounds. Relatively little information. On Oahu, Hawaii, a group of Pacific Golden-Plover loafing in low vegetation suddenly flattened to the ground with heads outstretched and remained motionless for at least 30 s (OWJ), response apparently triggered by something perceived as a raptor, probably Great Frigatebird (Fregata minor) that was overhead at the time.