Breeding

Pair Formation

Most birds appear to arrive on nesting grounds, or on early, snow-free openings near nesting grounds, unpaired. The earliest arrivals tend to be males (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 , 124 Jehl, J. R., Jr. (2004a). Birdlife of the Churchill Region: Status, History, Biology. Trafford, Victoria, BC, Canada. Close ). Pairing takes place as the plovers linger on openings or when a female settles on territory established by a male (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 , 119 Jehl, J. R., Jr., and B. A. Smith (1970). Birds of the Churchill region, Manitoba. Special Publication no. 1. Manitoba Museum of Man and Nature, Winnipeg, Manitoba, Canada. 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 ). Territorial display (Butterfly Display) flights begin within the first few days after arrival (120 Martin, P. D., and C. S. Moiteret (1981). Bird populations and habitat use, Canning River delta, Alaska. U.S. Fish and Wildlife Service, Arctic National Wildlife Refuge, Fairbanks, Alaska, USA. 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 ). 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 three to six days of the earliest arrivals (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 ). Display flights are common throughout the day and night early in the breeding cycle, and gradually decline as clutches are completed. The onset of bad weather on nesting grounds sometimes results in the displacement or disappearance of pairs. Males occasionally remain unpaired throughout all or a portion of the season. The latter situation probably indicates failure of pairing due to delayed snowmelt or loss of a replacement nest (see Demography and Populations: Measures of Breeding Activity), followed by desertion of the female (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 , O. W. Johnson).

Nest Building

The production of multiple nest scrapes by the male is part of pair-bonding (see Behavior: Sexual Behavior). A male completes his final nest within a few days after initiation of pairing behavior.

First/Only Brood

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 (O. W. Johnson), 5 June at Churchill, Manitoba (124 Jehl, J. R., Jr. (2004a). Birdlife of the Churchill Region: Status, History, Biology. Trafford, Victoria, BC, Canada. Close ), 9 June in north-central Alaska (59 Irving, L. (1960). Birds of Anaktuvuk Pass Kobuk and Old Crow: A study in Arctic adaptation. U.S. National Museum Bulletin 217. Close ), 14–15 June on Victoria Island, Nunavut (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 ), and 18–20 June on Bylot Island, Nunavut (171 Drury, W. H., Jr. (1961). The breeding biology of shorebirds on Bylot Island, Northwest Territories, Canada. Auk 78:176–219. Close ).

Median clutch completion dates on Melville Peninsula, Nunavut were 20 June (n = 24) in 1981 and 19 June (n = 3) in 1982 (121 Montgomerie, R. D., R. D. Cartar, R. L. McLaughlin, and B. Lyon (1983). Birds of Sarcpa Lake, Melville Peninsula, Northwest Territories: breeding phenologies, densities and biogeography. Arctic 36:65–75. Close ). Median nest initiation dates on North Slope, Alaska varied from 8–12 June over four consecutive seasons (138 Moiteret, C. S., T. R. Martin, and P. D. Martin (1996). Predevelopment surveys of nesting birds at two sites in the Kuparuk Oilfield, Alaska, 1988–1992. U.S. Fish and Wildlife Service, Northern Alaska Ecological Services, Fairbanks, AK, USA. Close ), and from 1–26 June near Prudhoe Bay, Alaska, over nine seasons (250 Troy, D. M. (1996). Population dynamics of breeding shorebirds in arctic Alaska. International Wader Studies 8:15–27. Close ).

Late clutches, some probably replacement layings (see Demography and Populations: Measures of Breeding Activity), are reported until the end of June. As with egg-laying, hatching schedule varies with seasonal conditions. Approximate periods for various locations are: first and second weeks of July, Churchill (119 Jehl, J. R., Jr., and B. A. Smith (1970). Birds of the Churchill region, Manitoba. Special Publication no. 1. Manitoba Museum of Man and Nature, Winnipeg, Manitoba, Canada. Close , 221 Krijgsveld, K. L., J. W. H. Reneerkens, G. D. McNett, and R. E. Ricklefs (2003). Time budgets and body temperatures of American Golden-Plover chicks in relation to ambient temperature. Condor 105:268–278. Close ); second week of July, Victoria Island, Melville Peninsula, Prince Charles Island (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 , 121 Montgomerie, R. D., R. D. Cartar, R. L. McLaughlin, and B. Lyon (1983). Birds of Sarcpa Lake, Melville Peninsula, Northwest Territories: breeding phenologies, densities and biogeography. Arctic 36:65–75. Close , 173 Johnston, V. H., and S. T. Pepper (2009). The birds of Prince Charles Island and Air Force Island, Foxe Basin, Nunavut. Occasional Paper No. 117, Canadian Wildlife Service, Ottawa, Ontario, Canada. Close ); third week of July, Bylot Island (171 Drury, W. H., Jr. (1961). The breeding biology of shorebirds on Bylot Island, Northwest Territories, Canada. Auk 78:176–219. Close ); fourth week of July, Devon Island (174 Hussell, D. J. T., and G. L. Holroyd (1974). Birds of the Truelove Lowland and adjacent areas of Northeastern Devon Island, N.W.T. Canadian Field-Naturalist 88:197–212. Close ); late June to mid July, Seward Peninsula (O. W. Johnson). If nesting is delayed by late snowmelt or clutches are replaced, hatching may extend to late July or early August (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 ). They produce a maximum of one brood per season.

Selection Process

Males establish territories in which they create several scrapes during pair-bonding, but the selection process for the final nest is unknown. On the Seward Peninsula, Alaska, males return to the previous year's territory (females are seldomly site faithful) and often nest within 100 m of last year's nest (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 , 118 Johnson, O. W., P. L. Bruner, A. E. Bruner, and P. M. Johnson (2007c). American Golden-Plovers on the Seward Peninsula, Alaska: a new longevity record for the species and other breeding ground observations. Wader Study Group Bulletin 114:56–59. Close ).

Microhabitat and Site Characteristics

The American Golden-Plover nests in relatively dry, open tundra, but where sympatric with the Pacific Golden-Plover (Pluvialis fulva) (as on Seward Peninsula), typical conditions at nests differ between 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 ; see Habitat​: Habitat in Breeding Range). The typical nest is located within a mosaic of lichen-covered rocks or ground, covered with vegetation of Dryas spp., crowberry, and other scattered forbs, grasses, and sedges. On the Seward Peninsula, the American Golden-Plover tends to select high, rocky, steep sites with sparse, short vegetation. The Pacific Golden-Plover selects lower sites with more vegetation. At the extremes, the nests of the American Golden-Plover are built in lichen-covered rocky barrens with few flowering plants present, and the nests of the Pacific Golden-Plover 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 the Seward Peninsula, a reversal of the foregoing occurs occasionally with the American Golden-Plover nesting in habitat typical of the Pacific Golden-Plover and vice versa. There was one instance of the same American Golden-Plover male nesting in a moist, grassy tussock the first year, then switching to a nearby dry, barren, rocky site the next season (118 Johnson, O. W., P. L. Bruner, A. E. Bruner, and P. M. Johnson (2007c). American Golden-Plovers on the Seward Peninsula, Alaska: a new longevity record for the species and other breeding ground observations. Wader Study Group Bulletin 114:56–59. Close ).

Construction Process

The male forms a nest cup by scraping with the toes and rubbing with the breast (see Behavior: Sexual Behavior). The process also involves the male standing either in or next to the scrape and tossing bits of lichen alternately over each shoulder (O. W. Johnson).

Structure and Composition

The nest is usually lined with lichens, especially Thamnolia vermicularis, and some nests contain dry grasses, leaves of willow (Salix) and Dryas, and occasionally a few pebbles and/or small sticks. Some nests are very simple, with an incomplete lining over the surface of the ground (O. W. Johnson; also see 248 Murie, A. (1946). Observation on the birds of Mount McKinley National Park, Alaska. Condor 48:253–261. Close ).

Nests and eggs are impressively camouflaged. Nests are usually challenging to locate, as aptly described by Murdoch (224 Murdoch, J. (1885). Birds. In Report of the International Polar Expedition to Point Barrow, Alaska. U.S. Government Printing Office, Washington, DC, USA. pp. 104–128. Close ): “ . . . 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

The nest is a shallow circular depression. On Seward Peninsula, Alaska, nests ranged from 110–125 mm in diameter x 32–59 mm deep (n = 9). The diameters included the rim of plant material which sometimes extended slightly beyond the cup itself and the depth was to the surface of the lining at the center of the cup (O. W. Johnson). At Cape Henrietta Maria, Ontario, dimensions of nests were 100–125 mm (n = 5) x 40–50 mm deep (n = 3) (251 Peck, G. K., and R. D. James (1993). Breeding Birds of Ontario: Nidiology and Distribution Volume 1: Nonpasserines (First revision–Part B: Vultures to Phalaropes). Ontario Birds 11:83–91. Close ).

Microclimate

Insulative properties of nests are relatively inefficient, contributing to energetically expensive incubation (see 252 Andreev, A. V. (1984). Experimental study of energetic parameters of incubation in some arctic birds. Doklady Biological Sciences 278:559–563. Close ). Among shorebirds nesting on the Taimyr Peninsula, Far North Russia, Tulp and coworkers (216 Tulp, I. (2007). The arctic pulse. Timing of breeding in long-distance migrant shorebirds. Ph.D. dissertation, University of Groningen, The Netherlands. Close ) found that larger birds (including the closely related Pacific Golden-Plover (Pluvialis fulva)) 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.

Maintenance or Reuse of Nests

Nest cups are known to be reused, how often is uncertain because the original “owner/builder” of the cup is 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 the cup was first found. Nests are readily available for reuse, since cup and lining often persist for many seasons (118 Johnson, O. W., P. L. Bruner, A. E. Bruner, and P. M. Johnson (2007c). American Golden-Plovers on the Seward Peninsula, Alaska: a new longevity record for the species and other breeding ground observations. Wader Study Group Bulletin 114:56–59. Close ; O. W. Johnson 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 the Pacific Golden-Plover nesting in a previous cup of an 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 the initial male captured there, and once by an unmarked conspecific (253 Johnson, O. W., R. H. Goodwill, R. S. Gold, and P. M. Johnson (2008e). Clutch replacement and between-clutch mate fidelity of Pacific Golden-Plovers Pluvialis fulva breeding near Nome, Alaska. Wader Study Group Bulletin 115:157–160. Close ). 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) ( 138 Moiteret, C. S., T. R. Martin, and P. D. Martin (1996). Predevelopment surveys of nesting birds at two sites in the Kuparuk Oilfield, Alaska, 1988–1992. U.S. Fish and Wildlife Service, Northern Alaska Ecological Services, Fairbanks, AK, USA. Close ). At Utqiagvik, Alaska, the American Golden-Plover had higher rates of reuse (10%, apparently all conspecific) than other shorebirds (254 Herzog, P., S. T. Saalfeld, H-H. Kaatz, and R. B. Lanctot (2018). Nest reuse in arctic‐breeding shorebirds: an analysis of potential benefits and factors affecting the occurrence of this rare behavior. Journal of Avian Biology e01737. Close ).

Nonbreeding Nests

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

Shape

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

Size

Similar in both the American Golden-Plover and the Pacific Golden-Plover (Pluvialis fulva). Representative size measurements are shown in Table 1. Mean egg volume is 24.2 ml ± 1.5 SD, n = 88 eggs (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 ).

Mass

Egg mass is about 25–29 g at laying (255 Parmelee, D. F., and J. M. Parmelee (1995). Additional bird nesting records and egg masses for the Seward Peninsula, Alaska. Northwestern Naturalist 76:134–136. Close , O. W. Johnson), declining to about 18–23 g near hatching (O. W. Johnson; 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.

Color

Similar in both the American Golden-Plover and the Pacific Golden-Plover; probably not distinguishable. Ground colors vary from whitish to buff, cinnamon buff, creamy buff, greenish buff, and ivory yellow. Eggs are heavily marked (especially near the large end) with irregular splotches and spots of dark brown and black, also a few underlying spots of gray (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 , 227 Sauer, E. G. F. (1962). Ethology and ecology of golden plovers on St. Lawrence Island, Bering Sea. Psychologische Forschung 26:399–470. Close , 256 Baicich, P. J., and C. Harrison (1997). A Guide to the Nests, Eggs, and Nestlings of North American Birds. 2nd edition. Academic Press, San Diego, California, USA. Close , 257 Bowman, T. (2004). Field Guide to Bird Nests and Eggs of Alaska's Coastal Tundra. Alaska Sea Grant College Program, University of Alaska, Fairbanks, AK, USA. Close ). On Seward Peninsula, ground color is typically greenish buff, predominant markings are dark brown (O. W. Johnson). Whether ground colors of the Pacific Golden-Plover “average paler” than those of the American Golden-Plover (203 Bent, A. C. (1929). Life histories of North American shorebirds, Part 2. United States National Museum Bulletin 146. Close ) is uncertain. Intensity and pattern of markings are variable both within and between clutches (O. W. Johnson).

Surface Texture

Smooth and slightly to moderately glossy (256 Baicich, P. J., and C. Harrison (1997). A Guide to the Nests, Eggs, and Nestlings of North American Birds. 2nd edition. Academic Press, San Diego, California, USA. Close , O. W. Johnson).

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. There was 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 the American Golden-Plover(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 , 119 Jehl, J. R., Jr., and B. A. Smith (1970). Birds of the Churchill region, Manitoba. Special Publication no. 1. Manitoba Museum of Man and Nature, Winnipeg, Manitoba, Canada. Close , 138 Moiteret, C. S., T. R. Martin, and P. D. Martin (1996). Predevelopment surveys of nesting birds at two sites in the Kuparuk Oilfield, Alaska, 1988–1992. U.S. Fish and Wildlife Service, Northern Alaska Ecological Services, Fairbanks, AK, USA. Close , O. W. Johnson). 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, 138 Moiteret, C. S., T. R. Martin, and P. D. Martin (1996). Predevelopment surveys of nesting birds at two sites in the Kuparuk Oilfield, Alaska, 1988–1992. U.S. Fish and Wildlife Service, Northern Alaska Ecological Services, Fairbanks, AK, USA. Close ]), 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 (123 Byrkjedal, I. (1989b). Nest habitat and nesting success of Lesser Golden-Plovers. Wilson Bulletin 101:93–96. Close ), likely involved egg-dumping. Replacement laying after nest loss together with between-clutch mate fidelity has been documented in both the American Golden-Plover and the Pacific Golden-Plover(118 Johnson, O. W., P. L. Bruner, A. E. Bruner, and P. M. Johnson (2007c). American Golden-Plovers on the Seward Peninsula, Alaska: a new longevity record for the species and other breeding ground observations. Wader Study Group Bulletin 114:56–59. Close , 253 Johnson, O. W., R. H. Goodwill, R. S. Gold, and P. M. Johnson (2008e). Clutch replacement and between-clutch mate fidelity of Pacific Golden-Plovers Pluvialis fulva breeding near Nome, Alaska. Wader Study Group Bulletin 115:157–160. Close ).

Egg Laying

Laying presumably starts with completion of the nest, but there are no precise observations. One female laid eggs at 2 day intervals (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 ), other findings suggest approximately a 1.5 day interval (172 Kessel, B. (1989). Birds of the Seward Peninsula, Alaska: Their Biogeography, Seasonality and Natural History. University of Alaska Press, Fairbanks, AK, USA. Close , O. W. Johnson). It is reasonable to assume a 6 to 7 day typical interval for laying of a complete 4-egg clutch, though a shorter period (approximately 4.5 to 5.5 d) was estimated by 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 . There is no information concerning the time of day when laying occurs. Replacement of individual eggs is not reported; for replacement of clutches, and also egg dumping, see Demography and Populations: Measures of Breeding Activity. Broken eggs are removed from the nest promptly. In one instance where eggs were probably trampled by reindeer, a pair of birds carried off the remains of the entire clutch, then deserted nest (O. W. Johnson).

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

Onset of Broodiness and Incubation in Relation to Laying

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

Incubation Patches

Both the male and female have 2 oblong patches, 1 on either side of the mid belly.

Incubation Period

About 25 days, no doubt varies somewhat with ambient temperatures and frequency of disturbance by investigators. Measurements: 26–27 days (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 ), 25 days (258 Montgomerie, R. D., R. V. Cartar, B. Lyon, and R. McLaughlin (1982). Shorebird studies at Sarcpa Lake, Melville Peninsula, Northwest Territories. Wader Study Group Bulletin 34:30–32. Close ), 25–27 days (138 Moiteret, C. S., T. R. Martin, and P. D. Martin (1996). Predevelopment surveys of nesting birds at two sites in the Kuparuk Oilfield, Alaska, 1988–1992. U.S. Fish and Wildlife Service, Northern Alaska Ecological Services, Fairbanks, AK, USA. Close ), and 23–24 days (124 Jehl, J. R., Jr. (2004a). Birdlife of the Churchill Region: Status, History, Biology. Trafford, Victoria, BC, Canada. Close ). Liebezeit et al. (259 Liebezeit, J. R., P. A. Smith, R. B. Lanctot, H. Schekkerman, I. Tulp, S. J. Kendall, D. M. Tracy, R. J. Rodrigues, H. Meltofte, J. A. Robinson, C. Gratto-Trevor, B. J. McCaffery, J. Morse, and S. W. Zack (2007). Assessing the development of shorebird eggs using the flotation method: Species-specific and generalized regression models. Condor 109:32–47. Close ) give egg flotation measurements for estimating the stage of incubation. Rates of daily energy expenditure (DEE) are very high among incubating shorebirds (260 Piersma, T., A. Lindstrom, R. H. Drent, I. Tulp, J. Jukema, R. I. G. Morrison, J. Reneerkens, H. Schekkerman, and G. H. Visser (2003). High daily energy expenditure of incubating shorebirds on High Arctic tundra: A circumpolar study. Functional Ecology 17:356–362. Close , 261 Tulp, I., H. Schekkerman, L. W. Bruinzeel, J. Jukema, G. H. Visser, and T. Piersma (2009a). Energetic demands during incubation and chick rearing in a uniparental and a biparental shorebird breeding in the High Arctic. Auk 126:155–164. Close ). There are no precise measurements in this species, however, based on Piersma et al. (260 Piersma, T., A. Lindstrom, R. H. Drent, I. Tulp, J. Jukema, R. I. G. Morrison, J. Reneerkens, H. Schekkerman, and G. H. Visser (2003). High daily energy expenditure of incubating shorebirds on High Arctic tundra: A circumpolar study. Functional Ecology 17:356–362. Close ), DEE during incubation is likely around 375 kJ/day.

Parental Behavior

The male usually incubates during the day (about 0800–2000 h) and the female incubates at night (about 2000–0800 h), but there is considerable individual variability (244 Byrkjedal, I. (1989a). Nest defense behavior of Lesser Golden-Plovers. Wilson Bulletin 101:579–590. Close , O. W. Johnson). When not incubating, the male generally forages on territory or at least within earshot of territory, and remains alert to predators that might threaten the incubating mate, and to intrusion by intraspecific or interspecific competitors. It also advertises territory with Butterfly Display flights (see Behavior: Spacing). The male may assist the disturbed female with distraction displays and aggressive behaviors, but the off-duty male frequently performs these actions with less intensity than the on-duty female (see Behavior: Predation). The non-incubating female usually forages at some distance from territory, generally beyond earshot from the mate.

Change-over is performed rapidly (O. W. Johnson): 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). The incubating birds carry shed feathers from the nest and discard them ≥ 75 m away (O. W. Johnson).

Hardiness of Eggs Against Temperature Stress

No experimental data, but Spindler (262 Spindler, M. A. (1978). Bird populations and habitat use in the Okpilak River delta area, Arctic National Wildlife Range, Alaska, 1978. Unpublished report, Arctic National Wildlife Refuge, U.S. Fish and Wildlife Service, Fairbanks, AK, USA. Close ) described an incident suggesting great hardiness, at least during early incubation. An American Golden-Plover nest about 5 days past completion of egg-laying was buried by a late snow storm on the Arctic Slope of Alaska, 21–23 June 1978. The nest was first located 23 June from behavior of the adults and plover footprints on a snowdrift. It was 15–20 cm under the snow, and the 4 eggs were cold. With the nest uncovered, the adult plovers resumed incubating, and eggs hatched between 12–14 July.

Preliminary Events and Vocalizations

During pipping, the Pacific Golden-Plover chick calls with high-pitched pfib (227 Sauer, E. G. F. (1962). Ethology and ecology of golden plovers on St. Lawrence Island, Bering Sea. Psychologische Forschung 26:399–470. Close ). A similar call is likely in the American Golden-Plover.

Shell Breaking and Emergence

The following details are based on hatching records from 8 nests on the Seward Peninsula, Alaska (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 ): 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 per chick from pipping until dry (n = 32) at Churchill, Manitoba (244 Byrkjedal, I. (1989a). Nest defense behavior of Lesser Golden-Plovers. Wilson Bulletin 101:579–590. Close ), approximately 4 d pipping to complete hatch of a clutch on Bylot Island (171 Drury, W. H., Jr. (1961). The breeding biology of shorebirds on Bylot Island, Northwest Territories, Canada. Auk 78:176–219. Close ).

Parental Assistance and Disposal of Eggshells

No assistance is reported during hatching. Parents remove eggshells soon after emergence of chicks, flying away to deposit fragments remote from nest site (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 , O. W. Johnson). Both parents are typically at the nest during the hatching process, with the female possibly doing the most incubation and brooding through this period (227 Sauer, E. G. F. (1962). Ethology and ecology of golden plovers on St. Lawrence Island, Bering Sea. Psychologische Forschung 26:399–470. Close ; O. W. Johnson; P. Bruner and A. Bruner, personal communication).

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 (263 Ricklefs, R. E. (1984a). Egg dimensions and neonatal mass of shorebirds. Condor 86:7–11. Close ); mass 19–21 g, culmen 10–1 mm, tarsus “about” 30 mm, n = 5 (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 ); mass 17.2 g and 19.5 g, n = 2 (39 Van Tyne, J., and W. H. Drury Jr. (1959). The birds of southern Bylot Island, 1954. Occasional Papers of the Museum of Zoology University of Michigan 615:1–37. Close ); mass 19.7 g, n = 2 (220 Visser, G. H., and R. E. Ricklefs (1993b). Temperature regulation in neonates of shorebirds. Auk 110:445–457. Close ); 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 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 ). For descriptions of plumage and bare parts, see Appearance: Plumages.

After hatching, the Pacific Golden-Plover (Pluvialis fulva) young use the same high-pitched pfib call mentioned earlier (see Hatching); also pfeb, pfiib, pfilib, and pfeeberee (227 Sauer, E. G. F. (1962). Ethology and ecology of golden plovers on St. Lawrence Island, Bering Sea. Psychologische Forschung 26:399–470. Close ). Calls of the young American Golden-Plover are likely similar, but have not been described. Early-hatched chicks (both species) frequently forage near the nest, while the 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 (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 ). Mass of chicks increased from approximately 20 g to 75 g in 15 days (222 Williams, J. B., B. I. Tieleman, G. H. Visser, and R. E. Ricklefs (2007). Does growth rate determine the rate of metabolism in shorebird chicks living in the arctic? Physiological and Biochemical Zoology 80:500–513. Close ). Fledging is estimated at 22–23 days (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 , 121 Montgomerie, R. D., R. D. Cartar, R. L. McLaughlin, and B. Lyon (1983). Birds of Sarcpa Lake, Melville Peninsula, Northwest Territories: breeding phenologies, densities and biogeography. Arctic 36:65–75. Close ). Compared to other shorebird neonates, the 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 (220 Visser, G. H., and R. E. Ricklefs (1993b). Temperature regulation in neonates of shorebirds. Auk 110:445–457. Close ). According to Williams et al. (222 Williams, J. B., B. I. Tieleman, G. H. Visser, and R. E. Ricklefs (2007). Does growth rate determine the rate of metabolism in shorebird chicks living in the arctic? Physiological and Biochemical Zoology 80:500–513. Close ), young birds can maintain body temperature under freezing ambient conditions when they reach 48% of adult mass.

Brooding

Chicks are brooded in the nest for variable period of hours after hatching is complete, and desert the nest thereafter. Both parents tend foraging chicks and frequently brood them. Visser and Ricklefs (220 Visser, G. H., and R. E. Ricklefs (1993b). Temperature regulation in neonates of shorebirds. Auk 110:445–457. Close ) studying metabolic characteristics of American Golden-Plover neonates in a laboratory chamber suggested that chicks might experience body temperature fluctuation between 38 and 30°C under field conditions. Krijgsveld et al. (221 Krijgsveld, K. L., J. W. H. Reneerkens, G. D. McNett, and R. E. Ricklefs (2003). Time budgets and body temperatures of American Golden-Plover chicks in relation to ambient temperature. Condor 105:268–278. Close ) conducted similar work in both laboratory and field, and among their 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 the nest, disturbance or threat causes chicks to flatten and remain motionless. The excellent camouflage of downy plumage makes the chicks 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. The chicks are not fed by parents. They forage independently by pecking at prey on vegetation or on the ground. Diet is apparently insects (Diptera and Coleoptera likely predominate, see 264 Machin, P., J. Fernández-Elipe, H. Flinks, M. Laso, J. I. Aguirre, and R. H. G. Klaassen (2017). Habitat selection, diet and food availability of European Golden Plover Pluvialis apricaria chicks in Swedish Lapland. Ibis 159:657–672. Close ) and spiders. Both adults tend the young during at least the first 2 weeks, leading them to foraging areas and protecting them from predators.

Initial foraging takes place on territory, but the family soon moves to more densely vegetated tundra in search of food and, perhaps most importantly, concealment (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 , 120 Martin, P. D., and C. S. Moiteret (1981). Bird populations and habitat use, Canning River delta, Alaska. U.S. Fish and Wildlife Service, Arctic National Wildlife Refuge, Fairbanks, Alaska, USA. Close , 172 Kessel, B. (1989). Birds of the Seward Peninsula, Alaska: Their Biogeography, Seasonality and Natural History. University of Alaska Press, Fairbanks, AK, USA. Close , 265 Pattie, D. L. (1990). A 16-year record of summer birds on Truelove Lowland, Devon Island, Northwest Territories, Canada. Arctic 43:275–283. Close , 264 Machin, P., J. Fernández-Elipe, H. Flinks, M. Laso, J. I. Aguirre, and R. H. G. Klaassen (2017). Habitat selection, diet and food availability of European Golden Plover Pluvialis apricaria chicks in Swedish Lapland. Ibis 159:657–672. Close , O. W. Johnson, P. G. Connors). A pair with young chicks moved 0.8 km from the nest within 3 days of hatching (174 Hussell, D. J. T., and G. L. Holroyd (1974). Birds of the Truelove Lowland and adjacent areas of Northeastern Devon Island, N.W.T. Canadian Field-Naturalist 88:197–212. Close ); another family, within 4 days 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 (O. W. Johnson).

Never observed.

Never observed.

Based on relatively few observations, both parents remain with offspring through most of the 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.

With departure of adults on fall migration, flocks of fledged and independent young appear on the tundra and especially along the coast (172 Kessel, B. (1989). Birds of the Seward Peninsula, Alaska: Their Biogeography, Seasonality and Natural History. University of Alaska Press, Fairbanks, AK, USA. Close , P. G. Connors). The last individuals remain until late August to early October, typically juveniles.