American Golden-Plover

Pluvialis dominica

Order:
Charadriiformes
Family:
Charadriidae
Sections

Breeding

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Figure 2. Annual cycle of American Golden-Plover.

Figure is based on the annual cycle of Pacific Golden-Plovers overwintering in Hawaii and breeding in Alaska (Seward Peninsula). Most of American Golden-Plover cycle in Alaska (arrival, breeding, initiation of body molt, and departure) is similar; completion of body molt and primary molt need study. On the distant overwintering range of American Golden-Plover in Southern Hemisphere, spring migration begins much earlier than shown here, and fall movements of juveniles may extend later into the fall and early winter months; see Migration for details.

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Alternate (breeding) female American Golden-Plover on nest.

Incubation begins before completion of clutch. Males usually incubate during the day, and females at night, but there is considerable individual variation.

© Andrew Spencer , Alaska , United States , 13 June 2013
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American Golden-Plover nest with eggs.

Nest is a well-camouflaged shallow circular depression, usually lined with lichens. Typical nest is located within a mosaic of lichen-covered rocks or ground, with vegetation of scattered forbs, grasses, and sedges.

© Shawn Billerman , Manitoba , Canada , 7 July 2009
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American Golden-Plover nest with eggs.

Clutch size is almost invariably four. Eggs are buffy, and heavily marked with irregular splotches and spots of dark brown and black.

© Shawn Billerman , Manitoba , Canada , 7 July 2009
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American Golden Plover eggs.

Teller, Alaska. 9 June 1961. Photographer Rene Corado.

Phenology

Pair Formation

Most birds appear to arrive on nesting grounds, or on early snow-free openings near nesting grounds, unpaired; earliest arrivals tend to be males (31, 177); pairing takes place as plovers linger on openings or when female settles on territory established by male (44, 143, 37). Territorial display (Butterfly Display) flights begin within first few days after arrival (144, 37, 31). Time required for Butterfly Display flights and other pairing-related activities (see Behavior: Sexual Behavior) to affect bond is unknown, but pairs are evident within 3–6 d of earliest arrivals (31). Display flights common throughout the day and night early in breeding cycle, gradually decline as clutches are completed. The onset of bad weather on nesting grounds sometimes results in displacement or disappearance of pairs, and males occasionally remain unpaired throughout all or portion of season; latter situation probably indicates failure of pairing due to delayed snowmelt or loss of replacement nest (see Demography and Populations: Measures of Breeding Activity), followed by desertion of female (232, OWJ).

Nest-Building

Production of multiple nest scrapes by male is part of pair-bonding (see Behavior: Sexual Behavior). Male completes final nest within a few days after initiation of pairing behaviors.

First/Only Brood per Season

Egg-laying occurs from mid May to late June (Figure 2), but this varies by region and spring weather. Records of early completed clutches: 1 June on Seward Peninsula, Alaska (OWJ), 5 June at Churchill, Manitoba (177), 9 June in north-central Alaska (57), 14–15 June on Victoria Island, Nunavut (44), 18–20 June on Bylot Island, Nunavut (142).

Median clutch completion dates on Melville Peninsula, Nunavut, 20 June (n = 24) in 1981 and 19 June (n = 3) in 1982 (174). Median nest initiation dates on North Slope, Alaska varied from 8–12 June over 4 consecutive seasons (243), and from 1–26 June near Prudhoe Bay, Alaska, over 9 seasons (244).

Late clutches, some probably replacement laying (see Demography and Populations: Measures of Breeding Activity), reported to end of June. As with egg-laying, hatching schedule varies with seasonal conditions; approximate periods for various locations: first and second weeks of July, Churchill (143, 212); second week of July, Victoria Island, Melville Peninsula, Prince Charles Island (44, 174, 146); third week of July, Bylot Island (142); fourth week of July, Devon Island (147); late June to mid July, Seward Peninsula (OWJ). If nesting delayed by late snowmelt or clutches replaced, hatching may extend to late July or early August (232). Maximum of 1 brood per season.

Nest Site

Selection Process

Males establish territories in which they create several scrapes during pair-bonding, but selection process for final nest unknown. On Seward Peninsula, Alaska, males return to previous year's territory (females seldom site faithful) and often nest within 100 m of last year's nest (231, 232, 173).

Microhabitat and Site Characteristics

Nest in relatively dry, open tundra; but where sympatric with Pacific Golden-Plover (as on Seward Peninsula), average conditions at nests differ between species (37; see Distribution, Migration and Habitat​: Breeding Range). Typical nest is located within a mosaic of lichen-covered rocks or ground, with vegetation of Dryas spp., crowberry, and other scattered forbs, grasses, and sedges. On Seward Peninsula, American Golden-Plovers tend to select high, rocky, steep sites with sparse, short vegetation; Pacific Golden-Plover select lower sites with more vegetation. At the extremes, American Golden-Plover nests are built in lichen-covered rocky barrens with few flowering plants present, and Pacific Golden-Plover nests are built in dwarf shrub heath tundra or in low grasses and sedges. In regions where the species are allopatric, each species is less selective.

On Seward Peninsula, reversal of the foregoing occurs occasionally with American Golden-Plover nesting in habitat typical of Pacific Golden-Plover and vice versa. One instance of the same American Golden-Plover male nesting in moist grassy tussock first year, then switching to nearby dry barren rocky site the next season (173).

Nest

Construction Process; Structure and Composition Matter

Male forms nest cup by scraping with feet and rubbing with breast (see Behavior: Sexual Behavior). The process also involves male standing either in or next to scrape and tossing bits of lichen alternately over each shoulder (OWJ). Nest is usually lined with lichens, especially Thamnolia vermicularis, and some nests contain dry grasses, leaves of willow (Salix) and Dryas, occasionally a few pebbles and/or small sticks. Some nests are very simple with lining incomplete over surface of ground (OWJ; also see 241).

Nests and eggs are impressively camouflaged. Nests are usually challenging to locate, as aptly described by Murdoch (215): “ . . . it is simply time wasted to attempt to find the nest by looking for it, as I know by hard experience. The only way to make sure of the eggs is to withdraw some distance, and sit down patiently and wait for the bird to go back to her eggs, watching her if necessary with field glasses. Having marked her on the nest, one must walk towards it in a straight line, looking neither to the right nor the left and keeping his eyes fixed on the spot she rises from. He is then pretty sure of the eggs. However, the surface of the tundra is so uniform that a careless glance to one side or the other after the bird is flushed may throw the collector wholly off the track, and then he has to go back and wait for the bird to return again.”

Dimensions

Nest is a shallow circular depression. On Seward Peninsula, Alaska, nests ranged from 110–125 mm in diameter ⨯ 32–59 mm deep (n = 9); diameters included rim of plant material which sometimes extended slightly beyond the cup itself and depth was to surface of lining at center of cup (OWJ). At Cape Henrietta Maria, Ontario, dimensions of nest were 100–125 mm (n = 5) ⨯ 40–50 mm deep (n = 3) (245).

Microclimate

Insulative properties of nests relatively inefficient, contributing to energetically expensive incubation (see 246). Among shorebirds nesting on the Taimyr Peninsula, Far North Russia, Tulp and coworkers (207) found that larger birds (including closely related Pacific Golden-Plover) insulated nests less effectively than smaller species and offered several possible explanations, (1) larger mass produces energetically favorable surface to volume ratio of both bird and its eggs, (2) building simpler nest costs less energy, (3) biparental incubation results in eggs being seldom uncovered, (4) nests often located on drier tundra, and (5) excessive insulation may negate camouflage of nest.

Reuse of Nests

Nest cups known to be reused, how often is uncertain because original “owner/builder” of the cup generally unknown and may not be the bird captured/banded on the nest. On Seward Peninsula reuse has occurred up to at least 5 years after cup first found; nests are readily available for reuse since cup and lining often persist for many seasons (173; OWJ and P. Bruner, unpublished data). Reuse typically involves the male that was initially captured on the nest, sometimes reuse is by a conspecific or by an interspecific as with Pacific Golden-Plover nesting in previous cup of American Golden-Plover. In an instance of the latter, a Pacific Golden-Plover was the fourth known occupant of an American Golden-Plover cup nest that had already been used twice by an initial male captured there, and once by an unmarked conspecific (247). On the North Slope of Alaska, a record of interspecific reuse was reported involving an American Golden-Plover that nested in the previous cup of a Stilt Sandpiper (Calidris himantopus) (243). At Barrow, Alaska, American Golden-Plovers had higher rate of reuse (10%, apparently all conspecific) than other shorebirds (248).

Nonbreeding Nests

As noted above, extra scrapes are constructed by male during pair-bonding, but apparently only the final nest is lined.

Eggs

Shape

Ovate pyriform. Narrow ends of 4 eggs fit together snugly in center of nest, creating minimum brooding area for incubating bird.

Size and Mass

Similar in both American and Pacific golden-plovers; representative size measurements shown in Table 1. Egg mass about 25–29 g at laying (249, OWJ), declining to about 18–23 g near hatching (OWJ; P. Bruner and A. Bruner, personal communication). Mass of a freshly laid clutch (100+ g) is about equivalent to the fat-free weight of a female. Mean egg volume is 24.2 ml ± 1.5 SD, n = 88 eggs (31).

Color

Similar in both American and Pacific golden-plovers, probably not distinguishable. Ground colors vary from whitish to buff, cinnamon buff, creamy buff, greenish buff, and ivory yellow; heavily marked (especially near large end) with irregular splotches and spots of dark brown and black, also a few underlying spots of gray (194, 142, 218, 250, 251). On Seward Peninsula, ground color typically greenish buff, predominant markings dark brown (OWJ). Whether ground colors of Pacific Golden-Plover “average paler” than those of American Golden-Plover (194) uncertain. Intensity and pattern of markings variable both within and between clutches (OWJ).

Surface Texture

Smooth and slightly to moderately glossy (250, OWJ).

Eggshell Weight and Thickness

From the Western Foundation of Vertebrate Zoology (WFVZ) collection, pre-1947 eggshell weights and thicknesses averaged 1.45 g (range 1.23–1.79, n = 73 eggs, 20 clutches) and 0.188 mm (range 0.167–0.212, n = 77 eggs, 21 clutches), respectively. No evidence of egg-shell thinning associated with pesticides (see Conservation and Management: Effects of Human Activity).

Clutch Size

Four eggs per clutch is typical in American Pacific Golden-Plover (44, 143, 243, OWJ). However, clutch samples often average less than 4 eggs [e.g., 3.7 eggs (n = 21 clutches, WFVZ collection); 3.8 eggs (n = 26 clutches, 243)], presumably owing to partial depredation, or replacement clutches of less than 4 eggs. An unusual clutch of 8 eggs (gradually “depredated one by one”) was reported at Churchill, Manitoba (176), likely involved egg-dumping. Replacement laying after nest loss together with between-clutch mate fidelity has been documented in both American and Pacific golden-plovers (173, 247).

Laying presumably starts with completion of nest, but no precise observations. One female laid eggs at 2 d intervals (44), other findings suggest approximately a 1.5 d interval (145, OWJ). Reasonable to assume 6 to 7 d typical interval for laying of complete 4-egg clutch, though shorter period (approximately 4.5 to 5.5 d) estimated by 31). No information concerning time of day when laying occurs. Replacement of individual eggs not reported; for replacement of clutches, and also egg dumping, see Demography and Populations: Measures of Breeding Activity. Broken eggs removed from nest promptly; in one instance where eggs probably trampled by reindeer, pair of birds carried off remains of entire clutch, then deserted nest (OWJ).

Prior to onset of incubation, paired birds often forage together. With incubation, off-duty bird (especially female) usually departs territory and feeds elsewhere. During egg-laying period and early incubation, birds generally flee from human, may remain silent or sound alarm calls; full repertoire of calls and distraction displays more likely as incubation progresses (see Sounds and Vocal Behavior: Vocalizations and Behavior: Predation).

Incubation

Onset of Incubation

Some incubation begins before completion of clutch, between laying of eggs 2 and 3, or between eggs 3 and 4; appears initially to be partial warming only, full incubation begins when clutch is complete (44, 31, OWJ). Warming of incomplete clutches is probably done by male only; energetics of reproduction likely require female to spend her time feeding.

Incubation Patches

Male and female have 2 oblong patches, 1 on either side of mid belly.

Incubation Period

About 25 d, no doubt varies somewhat with ambient temperatures and frequency of disturbance by investigators. Measurements: 26–27 d (44), 25 d (252), 25–27 d (243), 23–24 d (177). Liebezeit et al. (253) give egg flotation measurements for estimating stage of incubation. Rates of daily energy expenditure (DEE) are very high among incubating shorebirds (254, 255). No precise measurements in this species; however, based on Piersma et al. (254), DEE during incubation is likely around 375 kJ/day.

Parental Behavior

Male usually incubates during the day (about 0800–2000 h) and female incubates at night (about 2000–0800 h), but considerable individual variability (237), OWJ). When not incubating, male generally forages on territory or at least within earshot of territory; remains alert to predators that might threaten incubating mate and to intrusion by intraspecific or interspecific competitors; also advertises territory with Butterfly Display flights (see Behavior: Spacing). May assist disturbed female with distraction displays and aggressive behaviors, but off-duty male frequently performs these actions with less intensity than on-duty female (see Behavior: Predation). Non-incubating female usually forages at some distance from territory, generally beyond earshot from mate.

Change-over is performed rapidly (OWJ): oncoming bird flies to vicinity of nest and walks toward it; when the approaching individual is about 5–15 m from nest, mate flies away; replacement bird usually on the nest < 1 min after mate's departure. Prebasic molt begins during incubation (see Appearance: Molts); incubating birds carry shed feathers from the nest and discard them ≥ 75 m away (OWJ).

Hardiness of Eggs

No experimental data, but Spindler (256) described an incident suggesting great hardiness, at least during early incubation. An American Golden-Plover nest about 5 d past completion of egg-laying was buried by a late snow storm on the Arctic Slope of Alaska, 21–23 June 1978. Nest was first located 23 June from behavior of adults and plover footprints on a snowdrift; it was 15–20 cm under the snow, the 4 eggs cold. With the nest uncovered, adult plovers resumed incubating, and eggs hatched between 12–14 July.

Hatching

Preliminary Events and Vocalizations

During pipping, Pacific Golden-Plover chick calls with high-pitched pfib (218); similar call likely in American Golden-Plover.

Emergence

The following details based on hatching records from 8 nests on the Seward Peninsula, Alaska (232): progression from minute cracks in shell to obvious breakage (star-pip, or tiny pip hole) varied from 5–50 h; went from latter pipping states to actual emergence of individuals (which happened at all hours of the day and night) in 9.5–27.5 h, with most chicks emerging in 10–20 h; the interval from fine hairline cracks to 4 dry chicks in and around nest was approximately 2–4 d (42–96 h); from pipped condition to 4 dry chicks required about 1–3 d (21–67 h); emergence of brood members asynchronous (hatching-order probably directly related to laying-order) with intervals between emergence of first and fourth chicks ranging from 18–22 h; generally first chick to hatch was already mobile to around 3 m from nest before the last chick had emerged; from emergence of first chick to abandonment of nest by parents and brood was estimated at 26–30 h. Other measurements: 77 h average chick from pipping until dry (n = 32) at Churchill, Manitoba (237), approximately 4 d pipping to complete hatch of a clutch on Bylot Island (142).

Parental Assistance

No assistance reported during hatching. Parents remove eggshells soon after emergence of chicks, flying away to deposit fragments remote from nest site (31, OWJ). Both parents typically at the nest during the hatching process, with female possibly doing most incubation and brooding through this period (218; OWJ; P. Bruner and A. Bruner, personal communication).

Young Birds

Condition at Hatching

Mass and linear measurements (mean ± SD) of hatchlings include: mass 19.5 g ± 1.3 SD, range 17–21 g, n = 15 (P. Bruner and A. Bruner, personal communication); mass 18.3 g ± 1.7 SD, n = 14 (257); mass 19–21 g, culmen 10–11 mm, tarsus “about” 30 mm, n = 5 (44); mass 17.2 g and 19.5 g, n = 2 (39); mass 19.7 g, n = 2 (211); mass 19.0 g ± 0.8 SD (n = 7), culmen 11.9 mm ± 0.9 SD (n = 9), and tarsus 30.0 mm ± 0.6 SD (n = 9) (31). For descriptions of plumage and bare parts, see Appearance.

After hatching, Pacific Golden-Plover young use same high-pitched pfib call mentioned earlier; also pfeb, pfiib, pfilib, and pfeeberee (218). Calls of young American Golden-Plover are likely similar, but have not been described. Early-hatched chicks (both species) frequently forage near nest while adult continues to incubate late-hatching egg(s).

Growth and Development

Nidifugous (leave the nest shortly after hatching), ptilopaedic (downy) young grow rapidly. One “nearly six-day-old male” had grown to 30 g, its culmen to 15 mm, tarsus to 37.5 mm; bird had no Juvenile feathers except for “barely visible” wing quills (44). Mass of chicks increased from approximately 20 g to 75 g in 15 d (213). Fledging estimated at 22–23 d (44, 174). Compared to other shorebird neonates, American Golden-Plover hatchlings are relatively effective homeotherms since their body mass is sufficient to provide a favorable ratio between heat production and heat loss (211). According to Williams et al. (213), young birds can maintain body temperature under freezing ambient conditions when they reach 48% of adult mass.

Parental Care

Brooding

Chicks brooded in the nest for variable period of hours after hatching is complete, desert nest thereafter. Both parents tend foraging chicks and frequently brood them. Visser and Ricklefs (211) studying metabolic characteristics of American Golden-Plover neonates in laboratory chamber suggested that chicks might experience body temperature fluctuation between 38 and 30°C under field conditions. Krijgsveld et al. (212) conducted similar work in both laboratory and field, among findings: chicks returning to parents for brooding had body temperatures that never fell below 35.5°C; foraging bouts lengthened with increasing ambient temperatures and age, bouts were substantially longer between 0600–1100 h than at other times; brooding bouts averaged 12 min. Frequency and duration of brooding likely depend on weather conditions. Both before and after deserting nest, disturbance or threat causes chicks to flatten and remain motionless; excellent camouflage of downy plumage makes them almost impossible to detect.

Feeding

Coordinated movements and other responses develop rapidly, and within a few hours of hatching precocial young are apparently adept at finding food. Chicks not fed by parents; forage independently by pecking at prey on vegetation or on ground. Diet apparently insects (Diptera and Coleoptera likely predominate, see 258) and spiders. Both adults tend young during at least first 2 wk, leading them to foraging areas and protecting them from predators.

Initial foraging takes place on territory, but family soon moves to more densely vegetated tundra in search of food and, perhaps most importantly, concealment (142, 44, 144, 145, 259, 258, OWJ, PGC). A pair with young chicks moved 0.8 km from the nest within 3 d of hatching (147); another family, within 4 d of hatching, traveled about 0.5 km downward from its nest on a dry, rocky ridge and crossed a road to reach moist, relatively tall grassy vegetation on a lower slope (OWJ).

Cooperative Breeding

Never observed.

Brood Parasitism by Other Species

Never observed.

Fledgling Stage

Based on relatively few observations, both parents remain with offspring through most of chick stage. As young approach or gain flight capability, one or both parents may abandon them and join other adults to begin migration, or adults and young may join foraging flocks.

Immature Stage

With departure of adults on fall migration, flocks of fledged and independent young appear on the tundra and especially along the coast (145, PGC). Last individuals remain until late August to early October, typically juveniles.

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