Pacific Golden-Plover Pluvialis fulva Scientific name definitions
Version: 1.1 — Published April 15, 2021
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Not well understood. Most birds appear to arrive on nesting grounds, or on early snow-free openings near nesting grounds, unpaired; earliest arrivals tend to be males (37); pairing evidently takes place as plovers linger on openings or when female settles on territory established by male (150, 85). However, at least some appear to arrive already paired (47). Whether this is unusual (perhaps because of late season with prolonged stopovers en route), or simply indicates normal pairing at lengthy stopovers as in Japan (see 102, 179) is uncertain. Possible pairing of some Pacific Golden-Plover (closely spaced male and female foraging together with no aggressive interaction) occasionally seen in late April among pre-migrants in Hawaii (OWJ). Female choosing mate before arrival on breeding grounds then following male to his territory could be a factor in low return rates of marked females (see Movements and Migration: Dispersal and Site Fidelity).
Territorial display (Butterfly Display) flights may begin on day of arrival (47, B. McCaffery, personal communication) or within first few days after arrival (85, 37). 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 arrival (B. McCaffery, personal communication; 37). 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 (168, 167, 171; OWJ).
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.
Egg-laying occurs from mid-May to late June (Figure 2), varies by region and spring weather. Complete clutches by: 15 May on Alaska Peninsula (OWJ), 24 May at Bethel, Alaska (PGC), 3 June on Seward Peninsula, Alaska (OWJ), 18 June on St. Lawrence Island, Alaska (47), 17–24 June on Taimyr Peninsula, Far North Russia (302, 303, 61, 170). Median start of incubation: Alaska Peninsula approximately, 17–18 May (OWJ); Seward Peninsula, approximately 6–7 June (OWJ); and Taimyr Peninsula, 24–27 June (303, 60, 61, 304).
Late clutches, most probably replacement laying (see Demography and Populations: Measures of Breeding Activity), reported through much of July (47, 303, 61, 304, 201, 171; P. Tomkovich, personal communication).
As with egg-laying, hatching schedule varies with seasonal conditions and replacement laying. Main hatching periods in various locations approximately: mid-June southern end of breeding grounds on Alaska Peninsula (97); late June to early July, Seward Peninsula (OWJ); first week of July, southeastern Chukotka, Far Eastern Russia (180); and the third week of July, Taimyr Peninsula (302, 60, 170, 201).
Males establish territories in which they create several scrapes during pair-bonding, but selection process for final nest is unknown. Possibly do less scraping or none if a previous nest is reused. On Seward Peninsula, Alaska, males return to previous year's territory (females seldom site faithful) and nest relatively near previous nest (see Movements and Migration: Dispersal and Site Fidelity), sometimes reuse an earlier nest (293, 167). According to Rakhimberdiev et al. (305), the selection of nest sites by Pacific Golden-Plover in southeastern Taimyr Peninsula region started when snow cover had diminished to about 40%.
Microhabitat and Site Characteristics
Typical nest is located on relatively dry, open tundra within a mosaic of lichen-covered rocks or ground, with vegetation of Dryas spp., crowberry, other scattered forbs, grasses, and sedges (see Habitat: Habitat in Breeding Range). Where sympatric with American Golden-Plover (as Seward Peninsula), average conditions at nests tend to differ interspecifically: Pacific Golden-Plover often nest on more vegetated lower slopes while American Golden-Plover select high, rocky, steep sites with sparse, short vegetation (85, OWJ). At the extremes: Pacific Golden-Plover nests in dwarf shrub heath tundra or in low grasses and sedges, American Golden-Plover on lichen-covered rocky barrens with few flowering plants present. In regions where not sympatric, both species are less selective (see Habitat: Habitat in Breeding Range).
On Seward Peninsula, reversal of the foregoing occurs occasionally with Pacific Golden-Plover nesting in habitat typical of American 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 (147). Klaassen et al. (306) reported tendency for Pacific Golden-Plover in the high arctic (Taimyr Peninsula) to locate their nests near “landmarks” such as rocks and terrain disturbed by vehicle tracks. Reason for this behavior uncertain, among possibilities: “landmark” is cue to nest location amidst large expanse of homogeneous habitat. Similar site selection noted at Port Heiden on Alaska Peninsula, where plover nests were often located along edges of airstrip and gravel roads (OWJ).
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).
Structure and Composition
Nest is usually well-lined with lichens, especially Thamnolia vermicularis; 60–250 pieces in nests of Pacific Golden-Plover (47). Some nests contain dry grasses, leaves of willow (Salix) and Dryas, and occasionally a few pebbles and/or small sticks. Although there is considerable individual variation, nests of Pacific Golden-Plover on the Seward Peninsula often have thick linings (sometimes ≥ 25 mm); American Golden-Plover nests are frequently simpler with lining incomplete over surface of ground (OWJ; also see 307). Nests of Pacific Golden-Plover in eastern Siberia (Koryak region) were lined with 20–30 mm of lichens (149).
On Taimyr Peninsula, Far North Russia, Tulp (201) measured nest linings averaging 15 mm (n = 18), and suggested that variations might reflect adaptation to microclimate where nests are situated. Of these nests, lining composition was: 60% Thamnolia lichen, 25% other lichens, 10% willow leaves, and 5% sedge, grass, moss. Based on reuse of nests, thick lining might in some cases indicate remodeling of the previous nest cup.
Nests/eggs impressively camouflaged and usually challenging to find as aptly described by Murdoch (277): “ . . . 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.”
Nest is a shallow circular depression. On Seward Peninsula, Alaska, Pacific Golden-Plover nests ranged from 102–130 mm in diameter x 26–40 mm deep (n = 13); 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). In northeastern Siberia, nests (n = 11) averaged 115 mm in diameter (150); on the Taimyr Peninsula, average depth of nests was 36 mm (201).
Insulative properties of nests relatively inefficient, contributing to energetically expensive incubation (see 308). Among shorebirds nesting on the Taimyr Peninsula, Tulp (201) found that larger species (including Pacific Golden-Plover) insulated nests less effectively than smaller species and offered possible explanations: larger mass produces energetically favorable surface to volume ratio of both bird and its eggs, building simpler nest costs less energy, biparental incubation results in eggs being seldom uncovered, nests often located on drier tundra, excessive insulation may negate camouflage of nest.
Maintenance or Reuse of Nests
Nest cups known to be reused, but 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 10 years after cup first found; nests are readily available for reuse since cup and lining often persist for many seasons (293, 147; P. Bruner, unpublished data, OWJ). 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 that had already been used twice by initial male captured there, and once by an unmarked conspecific (171).
Extra scrapes may be constructed by male during pair-bonding, but apparently only the final nest is lined.
Ovate pyriform. Narrow ends of 4 eggs fit together snugly in center of nest, creating minimum brooding area for incubating bird.
Essentially the same in both Pacific Golden-Plover and American Golden-Plover; representative size measurements shown in Table 1. Size ranges 44.3–53.2 mm x 31–35.7 mm. For Pacific Golden-Plover, mean egg volume is 23.1 ml ± 1.4 SD, n = 36 eggs (37) and 25.7 ml ± 1.5 SD, n = 277 eggs (170; I. Tulp, personal communication).
Egg mass about 25–29 g at laying (309, OWJ), declining to about 18–23 g near hatching (69; 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 (18).
Color and Surface Texture
Similar in both Pacific and American golden-plovers, probably not distinguishable. Smooth and slightly to moderately glossy (310, OWJ). 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 (42, 279, 47, 310, 311). On Seward Peninsula, ground color is typically greenish buff, predominant markings dark brown (OWJ). Whether ground colors of Pacific Golden-Plover “average paler” than those of American Golden-Plover (42) remains uncertain. Intensity and pattern of markings vary both within and among clutches (OWJ).
Eggshell Weight and Thickness
From the Western Foundation of Vertebrate Zoology (WFVZ) collection, pre-1947 eggshell weights and thicknesses from 22 Pacific Golden-Plover clutches averaged 1.37 g (range 1.19–1.72, n = 85), and 0.177 mm (range 0.163–0.196, n = 88), respectively. No evidence of egg-shell thinning associated with pesticides (see Conservation and Management: Effects of Human Activity).
Four eggs per clutch is typical in Pacific Golden-Plover (150, 61, WFVZ collection, OWJ). However, samples often average less than 4 eggs [e.g., 3.7–3.9 eggs (303); 3.6 eggs (59); 3.9 eggs, 170)], likely due to partial depredation or 3-egg replacement clutches (see 171). Two 5-egg clutches found on Seward Peninsula (OWJ) were presumably the result of egg-dumping.
Laying presumably starts with completion of nest, but no precise observations. Relative to snow melt, Schekkerman et al. (170) noted in Pacific Golden-Plover an apparent 2–5 d lag between exposure of nest site and the start of egg-laying. Eggs are laid at approximately 1.5 d intervals (198, OWJ). Reasonable to assume 6–7 d typical period for laying complete 4-egg clutch, though shorter period (approximately 4.5 to 5.5 d) estimated by Byrkjedal and Thompson (37). 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 (47, OWJ). Sauer observed male removing predator-damaged egg from vicinity of nest: bird “grasped it with his bill,” flew off, and dropped egg about 100 m away.
Prior to onset of incubation, paired birds often forage together near the nest. With incubation, off-duty bird (especially female) usually departs territory and feeds elsewhere. During egg-laying period and early incubation, birds often 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).
Onset of Broodiness and Incubation in Relation to Laying
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 (37, OWJ). Warming of incomplete clutches appears to be by male only; energetics of reproduction likely require female to spend her time feeding.
Male and female have two oblong patches, one on either side of mid belly.
Incubation about 25 d, no doubt varies somewhat with ambient temperatures and frequency of disturbance by investigators. Measurements: 25 d (61; B. McCaffery, personal communication), 22–24 d from laying to first cracking of eggs (150). Schekkerman et al. (170) provide detailed egg flotation measurements for estimating stage of incubation in Pacific Golden-Plover. Rates of daily energy expenditure (DEE) are very high among incubating shorebirds (312, 313). No precise measurements in this species, however based on Piersma et al. (312) DEE during incubation is likely around 375 kJ/day.
Males usually incubate during the day (about 0800–2000 h), females at night (about 2000–0800 h), but considerable individual variability (47, 301, 293, OWJ). Non-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, often beyond earshot from mate.
Incubation change-overs 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 Against Temperature Stress
No data for Pacific Golden-Plover, but likely similar to American Golden-Plover where Spindler (314) described an incident suggesting great hardiness, at least during early incubation: a nest about 5 d past completion of egg-laying was buried by a late snow storm on the Arctic Slope of Alaska that occurred 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 nest uncovered, adult plovers resumed incubating, and eggs hatched 12–14 July.
Preliminary Events and Vocalizations
During pipping, Pacific Golden-Plover chick calls with high-pitched pfib (47).
Shell Breaking and Emergence
The following details based on hatching records from 8 nests on the Seward Peninsula (167): 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 9.5–25.0 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 2–3 d (47–67 h; from pipped condition to 4 dry chicks required about 1–2 d (26–50 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. Timing for Pacific Golden-Plover nesting on St. Lawrence I. agrees with the foregoing: first appearance of fine cracks to hatching of individual 21–27 h, pipping to hatching of individual 9 to 23 h, pipping to hatch of entire brood 32–44 h (47).
Parental Assistance and Disposal of Eggshells
No assistance reported during hatching. Parents remove eggshells soon after emergence of chicks, flying away to deposit fragments remote from nest (37, OWJ). Both parents typically at the nest during the hatching process, with female possibly doing most incubation and brooding through this period (47, 315, OWJ; P. Bruner and A. Bruner, personal communication).
Condition at Hatching
Mass and linear measurements (mean ± SD) of hatchlings include: mass 17.6 g ± 0.4 SD, n = 4, culmen 11.9 mm ± 0.5 SD, n = 4, total head 33.0 mm ± 0.5 SD, n = 4, tarsus 33.8 mm ± 0.8 SD, n = 4 (59); mass 16.4 g ± 0.3 SD, n = 3, culmen 11.8 mm ± 0.3 SD, n = 3, tarsus 32.9 mm ± 1.7 SD, n = 3 (60); mass 17.9 g ± 0.4 SD, n = 4, culmen 11.2 mm ± 0.7 SD, n = 6, tarsus 33.6 mm ± 1.1 SD, n = 6 (37). For descriptions of plumage and bare parts, see Appearance.
After hatching, young use same high-pitched pfib call mentioned earlier; also pfeb, pfiib, pfilib, and pfeeberee (47). Early-hatched chicks frequently forage near nest while adult continues to warm late-hatching egg(s).
Growth and Development
Nidifugous (leave the nest shortly after hatching), ptilopaedic (downy) young grow rapidly. Growth rate likely similar to American Golden-Plover, where one “nearly six-day-old male” had grown to 30 g, its culmen to 15 mm, and tarsus to 37.5 mm; bird had no Juvenile feathers except for “barely visible” wing quills (56). Mass of American Golden-Plover chicks increased from approximately 20 g to 75 g in 15 d (316). Fledging in Pacific Golden-Plover occurs at 26–28 d (B. McCaffery, personal communication). According to Williams et al. (316), young American Golden-Plover can maintain body temperature under freezing ambient conditions when they reach 48% of adult mass. Young Pacific Golden-Plover are likely the same.
Chicks brooded in the nest for variable period (hours) after hatching is complete, then desert nest thereafter. Both parents tend foraging chicks and frequently brood them. No detailed information concerning metabolic features of Pacific Golden-Plover chicks. Presumably, they are similar to American Golden-Plover. Neonates of the latter were studied in laboratory chamber by Visser and Ricklefs (317), findings suggested that their body temperature might fluctuate between 38 and 30°C under field conditions. Krijgsveld et al. (318) conducted similar work on same species 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 and 1100 h than at other times; brooding bouts averaged 12 min. Frequency and duration of brooding likely depends on weather conditions. Both before and after deserting nest, disturbance or threat causes Pacific Golden-Plover chicks to flatten and remain motionless; excellent camouflage of downy plumage makes them difficult to detect.
Coordinated movements and other responses develop rapidly within a few hours of hatching. Young are extremely precocial and apparently adept at finding food. Chicks not fed by parents, forage independently by pecking at prey on vegetation or on ground. Diet most likely spiders, larval and adult insects. Both parents tend young leading them to foraging areas and protecting them from predators.
Initial foraging takes place on territory, but family soon moves (sometimes to wetter tundra) in search of food and, perhaps most importantly, concealment in dense cover (47, 198, OWJ, PGC). Pacific Golden-Plover chicks foraged an average of 2.3 m/min, or about 3,300 m/d, on substrate with approximately 280 mg insect biomass/m2 (289, 319). Where breeding sympatrically with Bristle-thighed Curlew (Numenius tahitiensis), parents and broods occasionally associate for periods up to 6 d with amalgamated curlew families (320).
Brood Parasitism by Other Species
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. Females are often first to desert, leaving males the last adults to accompany juveniles (315, 187). Fledging usually takes c. 26–28 days from hatching.
With departure of adults on southward migration, flocks of fledged and independent young appear on the tundra and especially along the coast (198, PGC). Last individuals remain until late August to early October, typically juveniles.