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Grey plover

The plover (Pluvialis squatarola), also known as the black-bellied plover in , is a medium-sized shorebird in the family . It has a robust build with a short, stout black bill, long grey legs, and a chunky body, typically measuring 27–30 cm in length, with a of 71–83 cm and a weight of 190–280 g. Adults in breeding plumage feature striking black underparts, face, and foreneck contrasted against a head, rump, and undertail coverts, with spangled upperparts; non-breeding adults are duller, appearing above and whitish below, while juveniles show scaly brown backs. All plumages display diagnostic black axillaries (armpits) visible in flight. This species is a long-distance with a circumpolar breeding distribution across the high of (primarily ) and ( to the Canadian ), where it nests in solitary pairs on dry or wet with low vegetation. It departs breeding grounds from July to September and returns from May to June, undertaking extensive migrations along major flyways to winter in coastal habitats worldwide, including mudflats, estuaries, sandy beaches, and saltmarshes across the , , , , and . During non-breeding periods, it forms large, gregarious flocks—sometimes numbering in the thousands—while foraging diurnally and nocturnally for such as worms, bivalves, and crustaceans using a probing bill technique. Notable for its loud, whistled calls and vigilant sentinel behavior that alerts mixed shorebird flocks to danger, the grey plover's global population is estimated at 1,000,000–2,500,000 mature individuals as of 2024, though it is declining due to habitat degradation, disturbance, and pressures. Consequently, it was uplisted to Vulnerable on the in 2024.

Taxonomy

Etymology and classification

The grey plover bears the scientific name Pluvialis squatarola, first described by the Swedish naturalist in the 10th edition of Systema Naturae in 1758, where it was initially classified under the Tringa as Tringa squatarola based on specimens from regions. The Pluvialis was later established by the French ornithologist in his 1760 work Ornithologie, into which the species was transferred to reflect its distinct characteristics among plovers. This has remained stable since its adoption, with the grey plover placed within the Pluvialis, and three generally recognized, though their status is not universally accepted. The genus name Pluvialis derives from the Latin word pluvia, meaning "rain," stemming from an ancient belief that plovers, including this species, gathered in flocks as harbingers of impending rainfall. The specific epithet squatarola is a Latinized form of "squatarola," a term from dialect referring to a type of , likely alluding to the bird's robust, appearance or local regional naming conventions in . These etymological roots highlight the historical integration of and regional observations into early ornithological naming practices. In common , the is known as the "grey " in the , emphasizing its predominantly grey non-breeding plumage, while in the it is called the "black-bellied " to underscore the striking black underparts visible during the breeding season. This dual naming convention arose in the as European and North American ornithologists documented the ' transcontinental distribution, with early accounts in favoring "grey " based on winter observations and American texts adopting "black-bellied" after descriptions of breeding adults. Alternative historical names, such as "silver ," have occasionally appeared in older texts but are now obsolete. The grey plover is classified within the Charadriidae (plovers) and the order (shorebirds and allies), a placement consistent across major taxonomic authorities since the late . No significant taxonomic revisions affecting its , , or order status have occurred post-2000, reflecting stable phylogenetic analyses based on morphological and molecular data that affirm its position among plovers.

Subspecies

The grey plover (Pluvialis squatarola) is recognized as comprising three subspecies, distinguished primarily by subtle morphological variations and breeding distributions across the Arctic. These include the nominate subspecies P. s. squatarola, P. s. tomkovichi, and P. s. cynosurae, with no taxonomic revisions or genetic reclassifications reported in updates from authoritative sources as of 2025. However, their recognition is not universal, with some sources considering the species monotypic due to clinal variation. The nominate subspecies P. s. squatarola breeds across northern , from eastward through to western and northern , representing the most widespread form. It exhibits typical characteristics for the , with adults showing a black belly and face contrasted against spangled upperparts, though without the heightened contrast seen in other . This is marginally larger on average than cynosurae, with and tarsus lengths reflecting clinal variation increasing eastward within its . P. s. tomkovichi, described from populations on in northeastern , is the largest , with notably longer bills and tarsi compared to western populations of squatarola. Breeding is restricted to this eastern region, though individuals may overlap with squatarola in adjacent Siberian areas; plumage differences are minimal, aligning closely with the nominate form but supporting separation based on size metrics. The North American subspecies P. s. cynosurae breeds in northeastern and , with a more southerly distribution than its Eurasian counterparts. It is marginally smaller overall than squatarola, particularly in breeding males, which display more sharply contrasting black-and-white plumage; juveniles show more extensive white on the underparts relative to other . These distinctions aid field identification, especially in non-breeding ranges where the subspecies mixes.

Description

Plumage variation

The grey plover (Pluvialis squatarola) displays marked seasonal and age-related changes that aid in and identification. In breeding adult , the face, neck, and underparts are predominantly black, creating a strong contrast with the pale grey back and wings, which are spangled with black and white markings. The white rump becomes prominent during flight, enhancing visibility. In non-breeding plumage, adults shift to more subdued tones, with pale upperparts and clean underparts, though the axillaries remain a key feature visible in flight year-round. Juveniles resemble non-breeding adults but exhibit a distinctive scaly appearance on the upperparts due to or yellowish fringes on the feathers, often with yellowish spotting on tertials and coverts. The molt sequence involves a complete prebasic molt primarily in winter quarters to attain non-breeding plumage, followed by a partial prealternate molt in spring on the non-breeding grounds to develop breeding colors. Sexual dimorphism is minimal overall, though males typically show slightly brighter and more intense black on the face and underparts during breeding. The consistent black axillaries serve as a reliable aid across all s and ages, distinguishing the species from similar plovers. Certain , such as P. s. cynosurae in the , exhibit paler overall tones in non-breeding compared to the nominate form.

Size and morphology

The grey plover (Pluvialis squatarola) is a medium-sized shorebird characterized by a robust build that distinguishes it from smaller plovers in the family . Adults typically measure 27–31 cm in length, with a ranging from 71–83 cm and a body weight of 190–280 g; individuals can reach up to 345 g prior to long-distance to fuel endurance flights. This sturdy physique supports its role as one of the largest plovers, enabling efficient across vast distances. Key morphological features include a short, black bill measuring 24–34 mm in length, which is thicker and more robust than in congeners like the golden plovers, a rounded head, and longish black legs adapted for stability on soft tidal substrates. The legs are particularly strong, facilitating movement across uneven intertidal zones, while the feet feature a small hind toe—unique among Pluvialis species but rudimentary compared to more prominent toes in other groups. Sexual dimorphism is minimal, though adult females tend to be marginally larger than males in overall body size. Juveniles exhibit subtle differences, including shorter bills averaging 1.4 mm less than those of adults, and slightly lighter weights around 235 g compared to 243 g in adults; their wing length is also marginally reduced at approximately 195 mm versus 200 mm in adults. These variations in age identification but do not significantly alter the species' overall compact, stocky silhouette.

Distribution and habitat

Breeding grounds

The grey plover (Pluvialis squatarola) breeds exclusively in high regions spanning and . In , the subspecies P. s. cynosura occupies breeding grounds from western eastward through the , including areas around the Melville Peninsula and . In , P. s. squatarola breeds along northern coasts from the Kanin Peninsula to the Chukotskiy Peninsula, while P. s. tomkovichi is found in northeastern , particularly around the Chaunskaya . These populations utilize vast, remote landscapes north of 65°N, with an estimated extent of occurrence exceeding 19 million km² across the species' breeding range. Preferred breeding habitats consist of dry, gravelly or stony with sparse, low vegetation, including sedges, mosses, lichens, grasses, and dwarf birch. Nests are typically shallow scrapes lined with surrounding vegetation, situated on elevated sites such as ridges, riverbanks, raised or beaches, and rocky slopes to ensure visibility and drainage. These sites are often found at low to moderate elevations, up to approximately 1,000 m in some interior areas, favoring open terrains that minimize predation risk. The shows a strong preference for regions influenced by population cycles, as high lemming densities support invertebrate prey abundance and distract predators like arctic foxes from bird nests during peak breeding. Birds arrive on breeding grounds between late May and early June, with nest initiation peaking in mid-June in northern areas. The reproductive period lasts through , with departure for southward occurring from late to early , aligning with the brief summer. poses significant threats to these habitats, with models projecting an 82–87% reduction in suitable breeding area by 2070 due to warming and vegetation shifts.

Non-breeding range

The grey plover exhibits a nearly non-breeding distribution, utilizing coastlines across temperate to subtropical zones worldwide, including North and South America, , , , , and , with occurrences recorded in virtually every country during this period. This extensive winter range spans all major flyways, reflecting the species' adaptability to diverse coastal environments outside its breeding grounds. Key wintering sites include intertidal mudflats and estuaries such as the in northwest , Barr al Hikman in , in , the Yalu Jiang Estuary in , and Río Gallegos in , where significant concentrations gather to and . These locations, along with others like the north-central coast of and , support large portions of the non-breeding population but are generally confined to coastal margins, avoiding deep inland interiors. During the non-breeding season, grey plovers preferentially inhabit intertidal zones featuring soft substrates such as mudflats, sandflats, saltmarshes, and beaches within oceanic coastlines, bays, and estuaries, where they probe for invertebrates as tides recede. At high tide, they roost communally on nearby beaches or elevated areas, often in dense flocks, while some individuals defend small winter territories or roam more widely in search of food. Population distribution in winter shows strong regional biases, with up to 90% of the global population (estimated at 1,000,000–2,500,000 mature individuals) concentrated in the , where Nearctic-breeding birds predominantly overwinter in South American coastal sites. Palearctic populations, in contrast, primarily utilize Afro-Eurasian coasts, with notable concentrations of approximately 200,000 in and , 90,000 in southwest Asia and eastern Africa, 30,000 in , and 80,000 along the East Asian-Australasian .

Migration

Patterns and timing

The grey plover (Pluvialis squatarola) is a long-distance migrant that breeds in the high during the brief summer and winters in coastal regions worldwide across both hemispheres, including coastlines from North and South America, , , , , and . This annual cycle reflects its adaptation to exploit seasonal resources, with breeding timed to the Arctic's short period of abundant invertebrate prey and non-breeding periods aligned with milder conditions in wintering areas. Post-breeding southward typically begins in late for adults, peaking from to as birds depart breeding grounds and arrive at wintering sites, while the northward return occurs from to , with peak passage in mid-May for many populations. Juveniles depart breeding areas later than adults, often starting southward in and peaking in mid-autumn, which extends their overall timeline and increases exposure to environmental risks. Recent 2025 research on Arctic-nesting shorebirds indicates that these migrants exhibit adaptive timing in response to climate-driven advances in green-up, adjusting nesting schedules by up to 0.4 days per day of earlier onset to maintain synchrony with prey availability. Physiological preparations for these long flights involve significant fat accumulation, with birds increasing body mass by about 25%, primarily through protein deposition with some accumulation, at pre-migratory sites to non-stop journeys of thousands of kilometers. This load supports high flight speeds and endurance, essential for crossing ecological barriers like oceans. differences influence these patterns: Nearctic breeders from and often cross the Pacific to East Asian wintering grounds or the Atlantic to South American coasts, while Palearctic populations from primarily follow Eurasian flyways to and western wintering areas.

Routes and stopovers

The grey plover (Pluvialis squatarola) exhibits distinct migration routes depending on its breeding population. Nearctic birds, breeding in Alaska and northern Canada, primarily follow coastal pathways along the Pacific and Atlantic flyways of the Americas, with some utilizing inland corridors across the Great Plains during spring and autumn migrations. Palearctic populations, breeding across Arctic Eurasia from Scandinavia to Siberia, migrate via the East Atlantic Flyway toward West Africa and western Europe or the East Asian-Australasian Flyway to Southeast Asia and Australasia. A 2025 satellite-tracking study of an individual wintering in Singapore documented a northward route along the East Asian-Australasian Flyway to Bolshoy Begichev Island in north-central Siberia, spanning 8,645 km over 46 days, which suggests potential overlap between traditional Yamal-Taymyr and eastern Siberian-Alaskan populations, challenging prior delineations of Russian breeding groups. Migration distances vary by route and season, with total journeys averaging 5,500 km from wintering to grounds, though southward returns can exceed 12,000 km; legs typically range from 2,000 to 3,500 km, enabling efficient progress despite the ' moderate body size. For instance, East birds cover about 4,500 km per direction, with spring flights often faster and more direct than autumn ones. Critical stopover sites serve as essential refueling hubs where grey plovers spend up to 95% of their migration time on intertidal mudflats and gaining body mass for subsequent flights. Key locations include the mudflats in and for East Asian-Australasian migrants, where birds replenish energy reserves amid high prey availability; along the Pacific Americas Flyway, supporting coastal refueling for Nearctic populations; and Banc d'Arguin in on the East , a vital site for Palearctic birds to build fat stores before crossing the . Other notable stops encompass the in northwest and various Chinese coastal provinces like and . Habitat loss at these stopovers poses significant threats, as documented in recent and geolocator tracking data; for example, reclamation of Yellow Sea tidal flats has reduced available area by up to 65% over the past five decades, contributing to broader population declines by limiting refueling opportunities. Similar pressures from development and affect sites like Banc d'Arguin and , underscoring the need for targeted . International efforts, such as conservation programs, aim to protect these sites, though challenges persist as of 2025.

Reproduction

Nesting behavior

The grey plover constructs simple nests consisting of shallow scrapes or depressions in the ground, typically on dry, stony habitats such as raised ridges, hummocks, or gravelly areas that provide good visibility for predator detection. These nests are sparsely lined with local materials including , fragments, pebbles, or bits of plant matter to enhance against the surrounding gravelly , which contributes to breeding success by reducing predation risk. Nests are solitary, with pairs maintaining distances of over 400 m between them, corresponding to breeding densities of 0.3–2.3 pairs per km² in observed populations. Males establish and defend large nesting territories, often encompassing several hectares based on observed densities and display ranges, using aerial chases, vocalizations, and ground displays to deter conspecifics and potential intruders like golden plovers. Pair formation typically occurs on the grounds shortly after arrival, within 1–2 weeks, though some suggests may begin en route during migration or on wintering areas for certain populations. Pairs are monogamous for the season, with grey plovers reaching age at two years old. Reuse of exact nest sites from previous seasons is rare, though pairs may show by selecting locations within a few hundred meters of prior nests, further emphasizing the role of site in overall .

Eggs and parental care

The female grey plover lays a of four eggs, typically completed over 4–5 days with intervals of approximately 36 hours between eggs. The eggs are pear-shaped, measuring on average 51.4 mm in length by 35.3 mm in width (range 47.0–56.5 mm × 32.5–37.5 mm), and colored to olive with dark brown spots and blotches for . Incubation typically begins with or shortly before the final and lasts 23–27 days, during which the loses about 18% of its initial mass of approximately 35 g. Both parents share duties equally, with no consistent sex-specific shifts reported across studies, though males often cover more at night in some populations. survival during the full incubation period averages 54.7%, with hatching success of 97.5% for eggs that survive to pipping; overall nest success varies from 20–53% depending on predation pressure. The chicks are precocial and hatch over 1–2 days, often asynchronously, leaving the nest within 10–12 hours to follow parents while independently on . Both parents provide brooding for the first 2–3 days and continue leading the brood to food sources for about 10–14 days, with the female typically deserting after about 12 days and the male remaining until fledging. Young reach fledging at approximately 23 days, when they achieve flight capability and independence, though parental protection continues briefly thereafter. Fledging success for hatched chicks is generally high (around 70–90% in monitored broods), but overall reproductive output averages 2.1–2.6 fledglings per nesting attempt, limited primarily by predation on eggs and early chicks.

Behavior and ecology

Foraging and diet

The Grey plover (Pluvialis squatarola) primarily on intertidal mudflats and sandy beaches during , employing a visual run-stop-search to locate and capture prey. It runs briefly, halts to scan the surface with its large eyes, and then pecks or probes shallowly with its short, stout bill to extract items from the top few centimeters of . This method contrasts with the deeper probing of longer-billed shorebirds and allows efficient targeting of exposed or near-surface . Unlike more gregarious plovers such as the , the Grey plover tends to forage solitarily or in loosely spaced groups, sometimes displaying territorial aggression toward conspecifics to defend feeding patches. Its diet is predominantly invertebrate-based, consisting of polychaete worms, small bivalve molluscs, crustaceans, and , with occasional berries and seeds during the breeding season in Arctic tundra habitats. On non-breeding grounds, marine prey like polychaetes (Nereis diversicolor) and bivalves dominate, comprising up to 85% of intake in some areas, while terrestrial such as dipterans and lepidopterans become more prominent during migration stopovers. A 2024 DNA metabarcoding study on the southwestern coast revealed a broad spectrum of 332 prey taxa, with limited seasonal shifts: prevailed year-round (e.g., Diptera at 36.7% relative read abundance), but bivalves like mytilids increased in winter (up to 11.9% relative read abundance and 44.7% frequency of occurrence), reflecting adaptations to local prey availability in coastal habitats. Daily food intake typically ranges from 16 to 22 grams of fresh matter, sufficient to meet energetic needs during daylight in mild conditions, though it increases prior to to build reserves for long-distance flights. Nocturnal feeding supplements during winter when daylight hours are short, albeit at reduced rates due to lower . The bill's sturdy structure facilitates rapid extraction of buried prey without deep penetration, enabling the Grey plover to exploit a wide range of microhabitats efficiently. As a key predator in intertidal ecosystems, it helps regulate populations by selectively consuming abundant , thereby influencing benthic dynamics and prey , such as increased burial depths in response to plover presence.

Social structure and vocalizations

During the breeding season, grey plovers (Pluvialis squatarola) maintain a largely solitary or small-group social structure, with males establishing and vigorously defending expansive territories through aerial displays and vocalizations against conspecifics and sympatric species like the golden plover. These territories, often encompassing several hectares in Arctic tundra habitats, support monogamous pairs that share incubation and chick-rearing duties, fostering minimal group interactions beyond the family unit. In contrast, during the non-breeding season on temperate and tropical coasts, individuals shift to more gregarious behavior, foraging alone or in loosely aggregated small groups of up to 20 while roosting in dense flocks numbering hundreds to over 1,000 birds, particularly at high tide refuges on mudflats and beaches. These winter flocks exhibit loose dominance hierarchies, with occasional agonistic encounters such as upright threat postures or bill-fencing escalating in response to resource density, though overt aggression remains infrequent. Grey plovers lack complex songs but rely on a diverse array of calls for communication, which play key roles in territory defense, pair bonding, and group coordination. The most distinctive vocalization is the plaintive three-note flight call, rendered as "pee-wheep," "plu-ee," or "whee-er-eee," typically delivered by isolated birds to signal location, promote , or maintain contact during and , with pitch rising in poor conditions. Alarm calls include a high-pitched, trilling "krr-r-r" or sharp "plee" to alert flockmates and mixed-species shorebird assemblages of predators, positioning the grey plover as an effective worldwide. In contexts, vocalizations become more varied, featuring whistled calls and trills during aerial circuits to advertise ownership, deter intruders, and coordinate with mates, often accompanied by visual signals like slow, deliberate wingbeats and upright postures. These acoustic and postural displays facilitate pair formation and communication, with distraction calls used by parents to lure threats away from nests. Interactions within flocks occasionally involve subtle , such as chasing or pecking, but hybridization with other plovers is rare and poorly documented.

Conservation

Population status

The global population of the grey plover (Pluvialis squatarola) is estimated at 1,000,000–2,500,000 mature individuals, with the majority occurring in the Nearctic breeding populations that winter along the . Overall, populations are declining, with a suspected reduction of 30–49% over three generations (approximately 23 years, from 2005 to 2028), driven by habitat pressures across flyways; however, some regional populations, such as those in the Nearctic, show stability. The species is classified as Vulnerable on the since its 2024 assessment under criteria A2bcd+4bcd, reflecting rapid ongoing declines, with no change to this status in the 2025 Red List update. Regionally, the grey plover is listed as Vulnerable in under the Biodiversity Conservation Act 2016, based on a 2025 determination that estimates a moderate population reduction of 16–44% over three generations (about 22.8 years) for the approximately 11,300 mature individuals using Australian non-breeding sites. Population monitoring relies on the International Waterbird Census, coordinated by Wetlands International, which tracks wintering numbers and trends across key flyways to inform global assessments. In Arctic breeding grounds, densities vary but are generally low, ranging from 0.7 to 2.3 pairs per km², with mean values of 1–1.5 pairs per km² documented in areas like the Lena Delta in Russia. Demographic parameters include annual adult survival rates of 80–90%, with a specific estimate of 86% derived from ringing recoveries in European populations. Recruitment rates vary regionally and annually, influenced by breeding productivity in the Arctic, though precise global figures remain limited by monitoring challenges.

Threats and efforts

The Grey plover faces significant threats from habitat loss and degradation, particularly at key stopover and wintering sites along its migration routes. In the region, land reclamation for development has destroyed vast intertidal mudflats, reducing essential foraging areas by up to 65% since the 1950s and contributing to population declines in the East Asian-Australasian Flyway. Similarly, like alterniflora in coastal wetlands alter sediment dynamics and prey availability, exacerbating . Human disturbance from , activities, and emerging offshore wind developments further disrupts roosting and feeding, forcing birds to expend extra energy and lowering survival rates during migration. Illegal remains a concern across multiple flyways, including the East Asian-Australasian and , with market surveys in detecting Grey plover in 44% of samples. Climate change poses additional risks, particularly to breeding grounds and coastal s. Earlier disrupts breeding cues, creating phenological mismatches between insect hatches and chick rearing; a 2025 study found that snowmelt timing significantly affects and adult recruitment in Arctic shorebirds, with projections indicating 82-87% loss of suitable breeding by 2070 under moderate to high emissions scenarios. On wintering grounds, rising sea levels—projected to reach 70 cm by century's end—threaten tidal flats by inundating low-lying areas and altering nutrient flows, potentially reducing available by 20-50% in vulnerable regions like and . These changes may also lead to timing mismatches, as warmer conditions accelerate prey phenology while birds arrive on fixed schedules, impacting energy reserves. Conservation efforts for the Grey plover emphasize international protections and targeted monitoring to mitigate these threats. The species is safeguarded under the Agreement on the Conservation of African-Eurasian Migratory Waterbirds (AEWA), which promotes habitat safeguards and limits hunting across 255 waterbird species, alongside the Convention on Migratory Species (CMS) and the EU Birds Directive. Numerous Ramsar wetland sites, such as those in the Gulf of Mottama and Ariake Sound, provide critical protected foraging areas, with North Korea's 2018 accession enhancing flyway-wide coverage. Shorebird monitoring programs, including the Australasian Wader Studies Group and the Shorebirds 2020 initiative, track population dynamics and habitat use across flyways, informing adaptive management. Enhanced Spartina control in China aims for 90% eradication by 2025 to restore mudflats. These measures have yielded successes in select protected areas, where populations show signs of stabilization amid broader declines. For instance, in British Columbia's coastal reserves and sites (370 designated for the species), targeted habitat restoration and disturbance controls have maintained or slightly increased local numbers, contrasting with 16-44% declines elsewhere over recent decades.