Plovers and lapwings

(Charadriidae)

Class Aves

Order Charadriiformes

Suborder Charadrii

Family Charadriidae

Thumbnail description
Charadriids are small-to-medium, chunky shorebirds with rounded head, usually a "steep" forehead, short neck, and large eyes; typically black, brown, gray, and/or white; often with a bold pattern. Legs are short to medium and bill is typically shorter than the head

Size
5.5–16 in (14–40 cm); 1.25–10.5 oz (35–298g)

Number of genera, species
10 genera, 66 species

Habitat
Shorelines, wetlands, and open areas such as pastures, moors, and tundra

Conservation status
Critical, 1 species; Endangered, 2 species; Vulnerable, 5 species; Near Threatened, 6 species

Distribution
Worldwide except for Antarctica

Evolution and systematics

The family Charadriidae, including lapwings and plovers, is a fairly homogeneous group, although the number of genera has been extensively debated. Conservatively there are as few as six genera containing 56 species. Sibley and Monroe suggest as many as 11 genera containing 67 species, but in 1934, Peters divided the family into 34 genera with 61 species. In 2002, most authorities recognized 10 genera. Early efforts to understand the relationships within this family centered on skull characteristics, back color, and presence of a hind toe. These characteristics are no longer considered taxonomically significant. The two largest genera are Vanellus (lapwings) and Charadrius (ringed plovers and sandplovers).

Lapwings, with one exception, have a broad, black band near the tip of the tail; a terminal white band also is often present. The white-tailed lapwing (Vanellus leucurus) has a solid white tail and is sometimes placed, along with the sociable plover (Vanellus gregarius), in the genus Chettusia. Most lapwings, excepting V. vanellus and V. miles, also have a distinctive black-and-white wing pattern. Typically, primaries are black with a broad, white wing stripe extending diagonally across secondary coverts and secondaries. The blacksmith plover (Vanellus armatus) is sometimes placed in its own genus (Antibyx).

Charadrius plovers typically have a dark, complete or partial breast band and/or a white forehead with a dark line connecting the bill and eye. The shore plover (Thinornis novaeseelandiae) is sometimes included in Charadrius. Sibley and Monroe place the red-kneed dotterel (Charadrius cinctus) in the monotypic genus Erythrogonys, and the black-fronted dotterel (Charadrius melanops) in the monotypic genus Elseyornis. They also place the shore plover in Charadrius.

Smaller genera are Pluvialis (including the black-bellied plover [P. squatarola] and three golden-plovers), Eudromias (Eurasian dotterel E. morinellus), Oreopholus (tawny-throated dotterel O. ruficollis), wrybill (Anarhynchus frontalis), the inland dotterel (Peltohyas australis), Mitchell's plover (Phegornis mitchellii), and the Magellanic plover (Pluvianellus socialis). The common name "dotterel" is also used for some Charadrius species. The wrybill shares many characteristics with members of Charadrius, however, because of an unusual bill that bends to the right at about a 12° angle, it is placed in its own genus. The inland dotterel was formerly included in the courser family, Glareolidae. The Magellanic plover has unusually short legs and a bill that is unusually sharp for a plover. Some scientists consider the Magellanic plover more closely related to turnstones than to plovers; some place it in its own family, Pluvianellidae.

Physical characteristics

Charadriids are chunky, small to medium-sized shore-birds with short to medium-length legs, and a bill that is usually shorter than the head. Rarely colorful, most are

black-and-white with shades of gray and brown, though many are boldly patterned with either dark rings around the neck and strong facial markings or boldly marked black and white wings. Many lapwings have colorful wattles on the head and spurs on the wings. The spurs are used as weapons in territorial disputes and in nest defense. The hind toe is absent or greatly reduced in all species.

Distribution

The family is worldwide in distribution with the exception of Antarctica. The genus Charadrius is found throughout the world with a Holarctic center. Lapwings (Vanellus) are found throughout much of the world but are absent from North America. The center of lapwing distribution is apparently Africa. Five separate dispersals from Africa to Eurasia and two to South America probably account for the current distribution of Vanellus.

Several Charadriids limited to the southern tips of southern continents may be the relic descendants of a now extinct Antarctic species or of migratory Northern Hemisphere species that established breeding populations on their wintering ranges.

North American representatives are American golden-plover (Pluvialis dominica), Pacific golden-plover (P. fulva), black-bellied plover (P. squatarola), mountain plover (Charadrius montanus), semipalmated plover (C. semipalmatus), long-billed plover (C. placidus), Wilson's plover (C. wilsonia), piping plover (C. melodus), killdeer (C. vociferus), and snowy plover (C. alexandrinus).

Habitat

Charadriids are primarily birds of open areas and are found along shorelines of salt and/or freshwater bodies and flooded areas or on moors or tundra. Open grasslands and farmlands are often favored. Many species take advantage of human-altered habitats such as agricultural lands, sewage ponds, golf courses, airports, and even gravel roads and rooftops. Lapwings are primarily freshwater birds, but some are found on arid grasslands. The long-toed lapwing feeds and sometimes nests on floating vegetation. Golden-plovers are also primarily freshwater birds, while the black-bellied plover prefers coastal areas.

Migratory species may summer and winter in very different habitats and visit additional habitats along migration routes.

Behavior

Charadriidae includes migrant and resident species. Most are at least semi-gregarious during migration and on wintering grounds, where they form small-to-large communal foraging and roosting flocks. Mitchell's plover is unusual in that it is rarely seen in groups larger than six. While most are strong and graceful fliers, they are most often seen rapidly running. They are active both during the day and night. Most are quite vocal and their vocalizations have often given rise to local names. The familiar "killdee killdee" given to killdeer inspired both the accepted common name and the specific name vociferus.

Feeding ecology and diet

Most forage by rapidly running, then suddenly stopping, looking, and pecking at prey. Foot trembling or patting of the substrate, presumably to reveal potential prey, is seen in many species. The spur-winged lapwing (Vanellus spinosus) stands on one leg and moves the other back and forth to flush potential prey, including small lizards. The wrybill uses its unusual bill to extract mayfly larvae and fish eggs that cling to the underside of stones. The inland dotterel feeds in groups on plants during the day, but at night is more active, solitary, and feeds on invertebrates. The Magellanic plover feeds by pecking and often actively scratches and digs. It turns over stones in a manner similar to that of a turnstone. The red-kneed dotterel, the most aquatic member of the family, swims well and often feeds by wading, and submerges its head. It rarely feeds on dry ground. The white-tailed plover feeds mostly in shallow water and occasionally submerges its head while searching for food. Berries, especially crowberries, may be particularly important to plovers arriving in frozen tundra habitats where few insects are yet available. Gulls have been reported stealing food from both northern lapwings and golden-plovers.

Reproductive biology

Most charadriids lay their eggs in scrapes on the ground. The shore plover is unusual in nesting at the end of a tunnel in thick vegetation, under loose stones, or even in an abandoned petrel burrow. Shore plovers vigorously defend their nests, but feeding areas are communal. Other plovers also commonly feed away from the breeding territory in communal feeding areas.

Clutch size varies from 2–6, but is most often four. Clutches of six are rare. Incubation requires 18–38 days. Young are precocial and leave the nest shortly after hatching; they fledge at between 21 and 42 days of age. Adults, with one exception, do not feed the young. The Magellanic plover lays two eggs, but only one chick typically survives and is fed by the adults until after fledging. Reportedly some feeding is by regurgitation.

Some species prefer nesting on recently burned ground, a preference that provides some protection from future fires, but which also provides chicks with access to a flush of arthropods that take advantage of new plant growth following a fire.

Most are solitary nesters, but the sociable and white-tailed plovers commonly nest in semi-colonial groups. Sociable plover groups include up to two dozen pairs with nests about 150 ft (46 m) apart. Each pair defends its own small territory. White-tailed plover colonies may contain several hundred pairs, and nests may be only a few yards apart. The little-ringed plover (C. dubius) occasionally nests communally with inter-nest distances of as little as 26 ft (8 m). Killdeer often nest near other killdeer in areas with favorable habitat.

Many plovers are seasonally monogamous and some retain the same mates in subsequent breeding season. Snowy plovers in western North America are facultatively polyandrous and polygynous. The southern lapwing (Vanellus chilensis), while typically monogamous, will engage in cooperative breeding. It and the wrybill are the only members of the family in which two birds of the same sex have definitely been observed caring for a single clutch.

In most species parental responsibilities are shared, although the female may desert the male after laying a second clutch. The Eurasian dotterel female takes little interest in caring for her first clutch and may lay additional clutches for later mates. She rarely assists in caring for the young, but may rejoin the family group when the young are nearly grown. In the mountain plover the female often initiates a second clutch that she incubates, leaving the male to care for the first clutch. Mountain plovers produce a maximum of one brood per adult, although it may re-nest up to four times. Most plovers produce from one to three broods annually, but those nesting in warm climates have the potential to produce more. Parental care includes incubation, brooding, leading the chicks to feeding areas, and protecting them from predators by giving warning calls, performing distraction displays, and, less commonly, attacking potential predators. Distraction displays, including injury feigning and false brooding, are particularly well-developed in this family.

Some plovers, including white-fronted (C. marginatus) and Kittlitz's (C. pecuarius), typically nest in sand and cover the nest with sand when unattended. The white-fronted sand-plover brings water to the nest in its belly feathers. Other plovers such as the killdeer wet their belly feathers to help cool the eggs on extremely hot days. Adults in hot climates often spend much of their time shading eggs as opposed to incubating.

The nesting season may be long in species residing in warm climates. The chestnut-banded sandplover's (C. pallidus) nesting season may last nearly all year. The killdeer in Puerto Rico breeds year-round.

Conservation status

Charadriidae includes one Critically Endangered species, two Endangered species, five that are Vulnerable, and an additional six that are Near Threatened. The only charadriid listed as Critically Endangered is the Javanese lapwing (Vanellus macropterus), which is probably extinct. It has not been seen since 1940. Reportedly always uncommon, it frequented steppe-like marshes and river deltas on the west side of the north coast and the east side of the south coast of Java.

The Endangered St. Helena plover (C. sanctaehelenae) is the only surviving endemic land bird on St. Helena. There are only about 325 individuals of this grassland species remaining. A decline in available habitat, a fluctuating invertebrate food supply, human disturbance, and predation of nests by cats and the common myna (Acridotheres tristis) contribute to its decline.

The Endangered shore plover is restricted to South East Island in the Chatham Islands. Only about 159 birds remain. Recovery efforts include a captive breeding program and translocation. One translocated pair raised two chicks on Motuora Island, but in general translocation has been unsuccessful due to failed dispersals to the mainland and predation by the owl morepork (Ninox novaeseelandiae). Loss of habitat, expansion of fur seal (Arctocephalus forsteri) colonies, storms, and predation by cats, brown rats (Rattus norvegicus), and brown skua (Cartharacta lonnbergi) contribute to endangerment. Current trends may require upgrading to Critically Endangered.

The New Zealand dotterel (C. obscurus), mountain plover (8,000–9,000 birds), piping plover (5,913), wrybill (3,000–5,000), and sociable lapwing (2,500–10,000) are listed as Vulnerable. The New Zealand dotterel has a limited distribution in New Zealand. The northern population nests along the coastline, and the southern population nests on bare hilltops, bogs, and grasslands of Stewart Island. Extensive predator protection is potentially allowing populations to stabilize, although population trends are unclear. The Stewart Island population increased from a low of 62 in 1991–92 to 150 in 1999. The entire population is estimated at around 1,600 birds. The Mountain plover nests in short-grass prairie, dry scrub, and fallow fields and is often associated with prairie dogs (Cynomys sp.). Falling population numbers are primarily attributable to hunting and decline in habitat. Piping plover populations are estimated at 5,913 and are improving apparently due to conservation efforts. Drought, inappropriate water management, dredging, human development, efforts to stabilize beaches, and beach disturbance are listed as primary causes of population decline.

The Magellenic plover, Madagascar plover (C. thoracicus), Malaysian plover (C. peronii), Javan plover (C. javanicus), hooded plover (C. rubricollis), and Mitchell's plover (Phegornis mitchelli) are Near Threatened.

Significance to humans

Black-bellied plovers and golden-plovers were game birds commonly sold in markets in North America in the late 1800s and probably continue to be eaten in some cultures. Conservation of species, such as the snowy and piping plovers that nest on scantily vegetated beaches, is often in conflict with human use of beaches for leisure activities.

Species accounts

List of Species

American golden-plover

Pluvialis dominica

subfamily

Charadriinae

taxonomy

Pluvialis dominica P. L. S. Müller, 1776. Monotypic.

other common names

English: American lesser golden-plover; French: Pluvier doré d' Amérique, Pluvier bronzé; German: Sibirischer, Goldenregenpfeifer; Spanish: Chorlito Dorado Americano.

physical characteristics

9.5–11 in (24–28 cm); weight quite variable, ranging from about 3.5 to 7 oz (99–198 g), with post-migration weights

much lower. During breeding season, upperparts are black; yellowish edgings on the feathers of the upperparts give the birds a golden-spangled appearance. The forehead and sides of the neck are white. Face and underparts are black. In winter it is speckled brown above with pale underparts.

distribution

Across Canada and Alaska in tundra habitats; possibly on Chukoctskiy Peninsula, Wrangel Island, and Herald Island in Russia. Winter in grasslands of South America south to northern Argentina and Uruguay. May winter in Central America and along the Atlantic and Gulf coasts of North America, with most records from Florida. It is difficult to distinguish records of migratory stragglers from winter residents. Rarely individuals remain on the wintering grounds throughout the year.

Often found outside its normal range with reported occurrences along western African coast, the Netherlands, Ireland, Okinawa, New Guinea, and New Zealand.

habitat

Most common in the Arctic and sub-Arctic tundra and favors rocky slopes with scattered low vegetation for breeding. A variegated surface of rocks, gravel, lichen, and vegetation is preferred. Moves young to wetter areas with more shrubs and grasses for cover. During migration, found in a variety of open habitats, including inland and coastal areas. Winter primarily on grasslands of South America and less commonly on the coastal wetlands. Agricultural lands are not used.

behavior

Apparently migrate in small flocks, although large spring buildups are known from northwest Indiana and northeast Illinois with reports of more than 25,000 birds. Rapid fliers capable of long-distance flights. Flight speed may exceed 112 mph (180 kph). Some individuals establish small winter territories. Non-territorial individuals maintain individual spacing through low-level aggression. Wintering birds typically form communal roosts. Occasionally gather in large flocks at freshwater wetlands on wintering grounds to drink and bathe. Solitary nesters, but often forage in communal groups away from the breeding territory.

feeding ecology and diet

Feeds on a variety of terrestrial and aquatic invertebrates. Also some berries, seeds, and vegetation. Runs, stops, and pecks as is typical of foraging in plovers. Also pecks small invertebrates off of leaf surfaces.

reproductive biology

Most breed in the first year. Males, but not females, exhibit strong fidelity to breeding territories, which are large and defended by aerial displays and vocalizations. Formation of new pair bond is more common than retention of a mate from the previous season. Nests are shallow scrapes sparsely lined with lichens and/or dry grasses, pebbles, and leaves. Four eggs are laid. Both sexes incubate eggs and care for young. Males usually incubate during the day and females at night. Distraction displays, including false brooding, are given. Adults may attack some avian and mammalian predators, including foxes. May renest, but raise only a single brood. Incubation is about 26 days. Hatching is fairly synchronous (usually one day) and earlier hatched young forage near nest while attentive adult incubates.

conservation status

In 2002, populations appear stable, but populations declined drastically due to extensive hunting during the nineteenth and early twentieth centuries. For example, 48,000 were shot in a single day in Louisiana in 1821, and birds could be purchased for 25 cents a dozen in Maine in the 1850s. Populations rebounded with enactment of protective laws. Currently serious threats are probably loss of habitat in wintering areas due to agricultural and human encroachment. Pesticide exposure may also be a problem.

significance to humans

Hunted in Guyana, Suriname, French Guiana, and Barbados.

Ringed plover

Charadrius hiaticula

subfamily

Charadriinae

taxonomy

Charadrius hiaticula Linnaeus, 1758. Two subspecies.

other common names

English: Common ringed plover; French: Grand Gravelot; German: Sandregenpfeifer; Spanish: Chorlito de Collar, Chorlitejo Grande.

physical characteristics

7–8 in (18–20 cm). Males in breeding season: 2–2.54 oz (57–72g); females 1.98–2.65 oz (56–75 g). Upperparts, including crown, are sandy brown. Neck has a black ring with a white ring above it. White underparts, forehead, and superciliary line. Dark frontal bar and line running under the eye from the bill. In breeding male, bill is orange with black tip. Legs orange.

distribution

Breeds from northeastern Canada east to northeastern Siberia and winters in Africa, southern Europe, the Persian Gulf, and India.

habitat

Coastal during the breeding season, selecting nesting sites on beaches, near coastal tundra ponds, or on muddy plains. Rarely nests along rivers and occasionally found away from water. Favors

nest sites near high tide mark on sandy or shingled beaches. On migration and during winter found along seashores and coastal marshes where vegetation is low or occasionally in dry open areas. May be found on coral reefs exposed at low tide.

behavior

Outside of the breeding season most often seen in small flocks of up to 50 birds, but occasionally solitary and may also be seen in large flocks of up to 1,500 birds. Aggressive to neighbors on wintering grounds.

feeding ecology and diet

As is typical of plovers, it forages by running, stopping, lunging, and then running on. Foot patting or trembling is common. Forages during the day and at night. Eats a variety of invertebrates.

reproductive biology

Strongly territorial during breeding season. Nesting densities are typically low with fewer than one pair per hectare, but contiguous nesting territories of about 98 ft (30 m) in diameter occur. Birds may mate with the same partner in consecutive breeding seasons. Some birds arrive on breeding territory already paired. Both sexes incubate eggs and care for chicks. A clutch of four eggs is laid in a hollow in the sand. The nest is often lined with small pebbles. Incubation requires about 25 days, and birds fledge at about 23 days. Both sexes engage in injury feigning displays. Commonly double brooded, especially in southern populations. Nest sometimes reused, but new sites as much as 1,640 ft (500 m) from the first may be chosen.

conservation status

Populations are healthy and some populations in England and Scotland are expanding their ranges inland.

significance to humans

None known.

Killdeer

Charadrius vociferus

subfamily

Charadriinae

taxonomy

Charadrius vociferus Linnaeus, 1758. Three subspecies.

other common names

French: Pluvir kildir; German: Keilschwanz-Regenpfeifer; Spanish: Playero Sanero, Chorlito Tildio.

physical characteristics

8–11 in (20–28 cm); 3.3 oz (95 g). Medium-sized and monomorphic, with two black bands across the breast and a dark line extending between the eyes. Underparts are white and upperparts are gray-brown with rufous edging on some feathers. Rump and uppertail-coverts are bright rufous. Wings and tail are long.

distribution

Ranges across all of North America with the exception of most of Alaska and northern Canada. It extends southward throughout Central America, Columbia, and Venezuela, and along the west coast of South America to Peru and Chile. Also found on

Bermuda, throughout the Bahamas, Greater and Lesser Antilles, Cayman Islands, and Virgin Islands. There are accidental records from many places, including Hawaii, Russia, Greenland, Britain, Spain, and France. Breeding occurs across the North American range and south into Central America as well as the West Indies. Also reported breeding in Peru, northwest Chile, and southwest Ecuador. Resident over most of southern and coastal United States as far north as southern Alaska, West Indies, and areas of Central and South America.

habitat

Favors open areas, including sandbars, mudflats, closely mowed or grazed pastures and fields, graveled roads and parking areas, and even paved parking lots. It uses similar habitats during migration, but most likely to be seen in wet areas.

behavior

While they are graceful, rapid flyers, most commonly observed on the ground. Movement involves running a short distance, stopping, bobbing the head, and running again. Their loud calls of "killdee killdee" give the species its common name. They often serve as a sentinel species for mixed shorebird flocks and are frequently found in small to medium flocks in the winter, although individuals appear to protect a small personal space. Both migratory and resident populations are present. Resident pairs may maintain a breeding territory all winter. Migrant flocks that stopover on these territories are largely ignored, but territories are defended from other residents. Killdeer spend considerable time displaying in small groups, especially during fall and late winter to early spring.

feeding ecology and diet

Feeds along water edges, on shorelines, closely mowed pastures, and mudflats. Often forages by running short distances, stopping, peering, and pecking. Birds commonly pat the ground with one foot as they are foraging. Feeds on a variety of adult and larval invertebrates. Small vertebrates, including frogs and minnows, are sometimes eaten, as well as seeds and other plant materials.

reproductive biology

Seasonally monogamous in the United States. Non-migratory individuals commonly mate for life. Migrants have been observed with the same mate on breeding territories in consecutive years. Parental responsibilities are shared by both sexes. Lays four eggs in a scrape on the ground, preferring closely mowed pasture and graveled areas. Flat, gravel-covered rooftops are popular nesting sites. Both sexes participate in scrape formation. The bird crouches in the selected area and digs with the feet, pushing dirt to the rear and using the breast to form the scrape. Pair members take turns scraping. The displaced bird moves away from the scrape, tossing loose materials over the shoulder as it moves away. These loose materials eventually form a simple lining for the scrape. Incubation requires about 25 days, and hatching is usually fairly synchronous. Nests are rarely left unattended, especially when temperatures are high. Belly soaking (wetting of the breast feathers at a nearby pond or stream) is commonly used to cool eggs on extremely hot days. Precocial young are led to a feeding area soon after hatching. Fledging requires approximately 25–30 days, and fledged chicks may remain with the parents after fledging. Performs well-developed distraction displays and engages in false brooding when eggs or chicks are threatened.

conservation status

Populations appear stable over much of their range, although they may be declining in eastern North America. Human activities have increased available habitat for killdeer. Newly created habitats include gravel roadsides, parking lots, rooftops, golf courses, airports, and pastures. Unfortunately, many of these habitats act as ecological sinks when chicks are unable to get off rooftops, cars destroy eggs, or birds are killed through the ingestion of pesticides used to control their prey.

significance to humans

None known.

Kittlitz's plover

Charadrius pecuarius

subfamily

Charadriinae

taxonomy

Charadrius pecuarius Temminck, 1823; Cape of Good Hope. Monotypic.

other common names

English: Kittlitz's sandplover; French: Pluvier patre, Pluvier de Kittlitz; German: Hirtenregenpfeifer; Spanish: Chorlitejo Pecuario.

physical characteristics

5.25–6.5 in (13–16 cm); 0.67–1.7 oz (19–48 g). Characterized by a white forehead delineated by a black frontal bar and black line from bill to eye, which continues around the back and side of the neck. Crown brown with sandy edges on the feathers. Except for frontal bar, crown does not meet black markings, leaving a white ring around sides and back of crown. Upper dark brown with sandy feather edgings. Flight feathers blackish. Bill black, legs black to greenish gray. Sexes similar but black frontal bar less extensive and lighter in females. In juveniles the head color is buff and brown instead of black and white.

distribution

Africa and Madagascar.

habitat

Flat, exposed areas, including sand banks, mud banks, and dry veld. Frequently found far from water, rarely on sandy or rocky seashores.

behavior

Usually in pairs, but in winter found in small flocks (usually around 20 birds), but one flock of 270 reported. Often flocks with wintering Calidris species. Concentration of birds during non-breeding season are partially a result of receding water levels. They are gregarious even in the breeding season. May roost in mixed flocks or separately. Usually resident, but some populations undergo poorly understood seasonal movements.

feeding ecology and diet

Often feeds in groups of two to five birds. As is typical of plovers, it runs, stops, pecks, then runs on. It often pats its foot on foraging surface. Commonly feeds on moonlit nights.

reproductive biology

Somewhat gregarious in breeding season, sometimes nesting in semi-colonial groups (nests as close as 26 ft [8 m] but usually at least 59 ft [18 m] apart). Nest sites include sandy areas, sandy patches in open grassland, dry mud, and even droppings of cattle or horses. During copulation the male reportedly grabs the female's neck with his bill and falls onto his back, pulling the female with him so she ends up on top with her legs in the air. Two eggs are laid, and both sexes incubate and care for the young. When leaving the nest unattended in the middle of the day, or when disturbed, incubating bird quickly kicks sand over the eggs or newly hatched chicks. May stand over eggs and/or use belly soaking to cool eggs. Reportedly moves eggs up to 12 in (30 cm) in response to nest flooding. Incubation requires about 25 days, and chicks fledge at about 30 days, although brooding 42-day-old captive chicks have been observed. Both adults give injury-feigning distraction displays and false brooding when nest or young are threatened. One pair initiated a second nest while caring for previous brood.

conservation status

Common over much of the open habitat of Africa. Flooding is major cause of egg and chick loss, but predation and motor vehicles are also concerns.

significance to humans

None known.

Snowy plover

Charadrius alexandrinus

subfamily

Charadriinae

taxonomy

Charadrius alexandrinus Linnaeus, 1758. At least six subspecies are recognized.

other common names

English: Kentish plover, sandplover; French: Gravelot à Collier interrompu; German: Seeregenpfeifer; Spanish: Chorlitejo Patinegro.

physical characteristics

6–6.75 in (15–17 cm); 1–2 oz (28–57 g). Upperparts pale brown, especially pale in North American population. Breast band restricted to the lateral edges of the breast; white ring around the neck. Crown may be rufous. In breeding males a distinct, dark frontal bar marks the end of a white forehead and a dark line extends from bill to ear. Bill is black, and legs and feet are dark gray to black.

distribution

Breeds along the western U.S. coast from Washington to lower California and intermittently along the Gulf Coast from Marco Island, Florida, to the north coast of the Yucatan Peninsula. Breeds inland in south central Oregon, Salton Sea and eastern California, western and central Nebraska, northwestern Utah, and southern Arizona. Also breeds in the West Indies and on islands off the coast of Venezuela. Winters in

the Gulf drainage south to Yucatan and northern South America and in the Pacific drainage from central California to western Mexico. A resident population (C. a. occidentalis) is found on the coast of Peru and Chile. C. a. alexandrinus breeds in Eurasia from southern Sweden to the northern Sahara to western India and the steppes of central Asia through western China. Winters from the Mediterranean Basin south to tropical Africa, Angola, and Sri Lanka. C. a. seebohmi is resident in Sri Lanka. C. a. dealbatus breeds in eastern China and Japan and winters from Japan to Taiwan and the Philippines and from southeastern China to Indochina and the Greater Sundas.

habitat

Prefers open coastal areas, including sand and shingle beaches, estuaries, lagoons, and mudflats as well as inland along saline lakes. May also be found along rivers and on sparsely vegetated steppes. In North America it favors beaches and both coastal and inland salt flats. Inland it is found along braided river channels. Nests primarily on sandy coastal beaches, dunes, spoil islands, and salt flats or inland near brackish or saline wetlands. Several hundred birds nest and winter at agricultural wastewater ponds in the San Joaquin Valley and at the Salton Sea formed in southern California as a result of accidental flooding in the early 1900s. Mostly coastal outside the breeding season. Rare in freshwater habitats.

behavior

Species includes both resident and migrant populations. Most birds in flocks of up to 300 during winter, although some defend winter territories. Small groups of birds may take off in coordinated flights when disturbed. Both chicks and adults can swim, and adults may lead chicks across ponds and rivers.

While territories are important sites for nesting and feeding, some birds frequently forage communally at undefended feeding areas. Feeding areas as much as 3.7 mi (6 km) from the nest have been documented.

feeding ecology and diet

Runs, stops, looks, and pecks as is common in plovers, but also probes at the bases of plants and runs into dense masses of flies with bill open and snapping. May also charge after solitary insects. Occasionally pats substrate with foot. Mostly gathers food from sand surface both above and below mean high tide mark. Inland, often forages in 0.39–0.79 in (1–2 cm) deep water or on wet surfaces.

reproductive biology

Frequently retain their mate for more than one year, and one pair remained mated for at least six years, apparently as a result of territorial fidelity. Rarely male may mate with two females and alternate incubation at the two resulting nests. Birds often show nest site fidelity. Nest is often located near an item such as piece of driftwood or small clump of vegetation that distinguishes an otherwise uniform landscape. Nests are formed by scraping and lined by both sexes randomly picking up bits of debris and tossing it over their shoulders and into the nest. Lining eventually includes such items as pebbles, fish bones, and arthropod skeletons. During copulation male grasps the female's neck and falls backward, pulling her with him. Clutch size is usually three, but varies from 2–6.

Both sexes incubate, but at least in some localities the female incubates more during the day and the male at night. Incubation requires from 23–32 days. Young are precocial and leave nest soon after hatching.

In western North America females desert broods soon after hatching, mate with a different male, and initiate another nest. This allows for the production of multiple broods. Males also take a new mate and re-nest, in some cases as early as 10 days before the current brood fledges. Birds may move several hundred kilometers to re-nest. Females may mate with original mate to produce a third brood. In some localities only one brood is produced. Parental care includes leading chicks to food, watching and warning of danger, and giving injury-feigning distractions.

Breeding territories may be small. In some localities average distance between nests was 120 ft (36 m). Breeding typically occurs during the first year.

conservation status

North American Pacific and Gulf Coast populations are shrinking, with current populations estimated at 21,000 individuals in the United States. The Pacific Coast population is listed as Threatened by the U.S. Fish and Wildlife Service. Habitat loss and destruction is an important factor contributing to population decline. Chief problems are increased recreational use of beaches, regular raking to keep beaches attractive to humans, and use of beach grass (Ammophila arenaria) to stabilize beaches. Inland habitat has been lost as a result of human-mediated changes such as dam construction and growth of vegetation. Less important factors include pesticides, entanglement in monofilament line, being run over and stepped on, and shooting and trapping.

significance to humans

None known.

Wrybill

Anarhynchus frontalis

subfamily

Charadriinae

taxonomy

Anarhynchus frontalis Quoy and Gaimard, 1830, New Zealand. Monotypic.

other common names

English: Wry-billed plover, crook-bill plover; French: Pluvieranarhyngue; German: Schiefschnabelregenpfeifer; Spanish: Chorlitejo Piquituerto.

physical characteristics

7.75–8 in (19.7–20 cm); weight: males 2.1 oz (59.5 g), females 2.0 oz (56.7 g). Unique in having a bill that bends to the right at about a 12° angle. White forehead extends backwards. Dark gray band extends from bill, continues under eye, gradually lightening toward crown. Crown is nearly black where it meets white forehead. Rest of upperparts are uniform bluish gray, including wing coverts that are edged in white. Flight feathers are brown with outermost flight feathers gray with white edges. A broad, black band covers upper breast. Bill is black and legs are dark gray. Female similar, but breast band paler and narrower and crown is light where it meets white forehead. Breast band absent in juveniles and winter plumage.

distribution

Canterbury and Otago, South Island, New Zealand. Winters mainly on the Firth of Thames at Manukau Harbor, and on the Kaipara River, North Island, New Zealand.

habitat

Nest on large expanses of stones near rivers. During nonbreeding season most often found on silty mudflats near high tide mark on sheltered coasts and estuaries.

behavior

Large pre-migratory flocks at the Firth of Thames perform elaborate mass aerial displays.

feeding ecology and diet

Probes in mud, but also sweeps the bill sideways, capturing tiny crustaceans from water surface. Also pecks and probes between stones and sweeps tiny invertebrates from under stones in riffle areas. Feeds on spiders, insects, crustaceans, small mollusks, small fish, and eggs. Larval mayflies and caddisflies are commonly eaten.

reproductive biology

Have strong fidelity to breeding territories and even nest sites. Birds often pair with same mate in consecutive years, perhaps because of nest site tenacity. Breeding does not occur until second or third year. Typically nest on higher banks and parts of islands in wide areas of shingle with fairly large stones. Male forms scrape, lined by pebbles flicked into the scrape or regurgitated from the crop. Clutch size is two. Parental duties are shared. One male shared incubation of a four-egg clutch with two females. Incubation requires about 31 days, and young fledge at about 29 days. Typically nest again after first clutch fledges.

conservation status

Classified as Vulnerable; population of 3,000–5,000 birds appears to be declining. Breeding habitat is deteriorating due to increasing recreational use of rivers and to invasion of weeds, apparently as a result of hydroelectric plants upriver. Predation by stoat (Mustela erminea), cats, and kelp gulls (Larus dominicanus) is probably significant.

significance to humans

Shot for sport until protected in 1940.

Magellanic plover

Pluvianellus socialis

subfamily

Pluvianellinae

taxonomy

Pluvianellus socialis G. R. Gray, 1846. Sometimes placed in its own family, Pluvianellidae. Monotypic.

other common names

French: Pluvier Magellan; German: Magelanregenpfeifer; Spanish: Chorlito de Magallanes.

physical characteristics

7.75 in (20 cm); weight: males 2.8–3.6 oz (79–102 g), females 2.5–3.1 oz (71–88 g). Upperparts, head, and neck are ashy gray grading to a brownish gray on the upper chest and white on the cheeks, throat, and forehead. Dark area between eye and bill. Underparts are white. In flight a broad, white wing stripe is evident. Bill is black with a small pink spot near base. Turnstone-like in appearance. Iris is black. Legs are short and both legs and feet are coral pink. Females are slightly darker and heavier. In juveniles dorsal surface has golden tint, eye is white with lavender tinge, and legs and feet are orangeish. First winter birds have yellow mark at base of bill. Hind toe is well-developed unlike other plovers.

distribution

Breeds in extreme southern Chile and Argentina, including Tierra del Fuego. Winters northward to Valdez peninsula, south-central Argentina, and occasionally as far north as Buenos Aires province.

habitat

Breeds along shores of shallow fresh water or brackish lakes and lagoons—most of which are glacial—in the steppe-like regions of Tierra del Fuego and southern Patagonia. Winters mostly along the coast in sheltered bays, lagoons, and river mouths.

behavior

Pair members act as a unit in territorial defense. Most spend the winter in sheltered bays and river mouths.

feeding ecology and diet

Feeds by pecking at the water surface and scratching the ground with strong claws. They sometimes dig deeply into the sand, which is uncharacteristic of shorebirds. They also flip shells and bits of debris in the manner of a turnstone. Larvae of chironomid flies are the primary winter food, but a variety of tiny arthropods are eaten during the breeding season.

reproductive biology

Some populations nest up to 3,900 ft (1,200 m). Separate breeding and feeding territories are maintained. Breeding territories tend to be linear (984–1,640 ft [300–500 m]). Nest scrape lined with small pebbles and usually only a few feet from water. Clutch size is usually two. Four-egg clutches occur, probably the result of two females laying in the same nest. Both sexes incubate eggs and care for young. Injury feigning distraction displays are unknown. Hatching is asynchronous and second-hatched nestling often starves. Relative to other plovers, young are weak and remain in nest vicinity for several weeks. Unlike other plovers, parents feed young—often by regurgitation. Fledging occurs around 28–30 days, but adults may feed young until at least 40 days old.

conservation status

Near Threatened. Factors contributing to small population size include hostile environment and the effect of grazing animals and introduced herbivores on natural grasslands.

significance to humans

None known.

Northern lapwing

Vanellus vanellus

subfamily

Vanellinae

taxonomy

Vanellus vanellus Linnaeus, 1758, Sweden. Monotypic.

other common names

English: Eurasian plover, green plover, pewit; French: Vanneau huppé; German: Kiebitz; Spanish: Avefría Europea.

physical characteristics

11.5–13 in (29–33 cm). Spring weights: males 6–9 oz (170–255g), females 7.7–9.9 oz (218–280 g). Adult summer and fall weights average 8.2 oz (232 g). Elongated black crest, thick blackish neck band, and glossy green back are diagnostic. Face is black with a dark line extending under the eye. Tail is white and has a broad sub-terminal black band. Tail coverts are cinnamon rufous. Underparts are white, grading to a light cinnamon on the undertail coverts.

distribution

Breeds in Europe and Asia from the British Isles (occasionally to Iceland) to south Ussuriland and south to northwest Morocco, eastward to north Greece, Iran, and Mongolia. Winter range extends southward to the Mediterranean Basin, northwestern India, and eastern China. Breeding has also been reported from Japan.

habitat

Habitat requirements are broad, and the species can be found nesting from boreal to steppe and even desert habitats. Like other charadriids prefers broad open areas, including grasslands, fields, moors, bogs, and heathlands.

behavior

Migrating flocks are usually small, but huge flocks are also recorded. Winter flocks often contain 100 or more birds, and flocks of over 5,000 have been reported. During winter some individuals feeding in flocks vigorously defend small feeding

territories. They also spend time on communal areas where bathing, resting, and preening occur.

feeding ecology and diet

Foraging behavior is typical charadriid behavior of running, stopping, and pecking. They feed extensively on earthworms, but diet also includes a variety of invertebrates. Reliance on earthworms decreases during unusually dry weather. During unusually cold weather they may eat cattle dung. They are often the victims of kleptoparasitism by gulls—especially black-headed gulls (Larus ridibundus).

reproductive biology

Mature in the first year, lapwings often do not nest until their second or even third year. Solitary nesting is common, but breeding densities are sometimes high (nine pairs in less than a hectare). Fidelity to territory is high, with 70% of lapwings returning to birthplaces in the spring. Male performs elaborate territorial and courtship displays over breeding territory. Flights include a non-vocal humming sound and a three-motif song. At times, the primaries emit loud buzzing noises. Seasonal monogamy is the rule but a weak pair bond and weak territories increase the probability of polygynous mating. Male may fly directly onto the female's back for copulation. A single brood is produced. Four is usual clutch size, occasionally three. Incubation requires about 24–34 days. Parental duties are shared, but one parent—usually the female—deserts before the brood fledges. Fledging occurs in 30–42 days. There are reports of males with two to three females, and simultaneous bigamous matings by males seem to occur often.

conservation status

Not threatened. European breeding area has expanded especially northward.

significance to humans

Eggs were commonly collected for food. There are reports from Holland of a single collector taking over 2,000 eggs in a single season. It remained legal to collect the eggs in Britain until at least the 1970s.

Resources

Books

Barnard, C. J., D. B. A. Thompson. Gulls and Plovers: The Ecology and Behaviour of Mixed-species Feeding Groups. New York: Columbia University Press, 1985.

Birdlife International. Threatened Birds of the World. Barcelona and Cambridge, UK: Lynx Edicions and BirdLife International, 2000.

Byrkjedal, I., and D. Thompson. Tundra Plovers: The Eurasian, Pacific and American Golden Plovers and Grey Plover. London: T & AD Poyser, 1998.

Falla, R. A., R. B. Sibson, and E. G. Turbott. The New Guide to the Birds of New Zealand. Auckland and London: Collins, 1981.

Gosler, A., ed. The Photographic Guide to Birds of the World. New York: Mallard Press, 1991.

Hayman, P., J. Marchant, and T. Prater. Shorebirds: An Identification Guide. Boston: Houghton Mifflin Company, 1986.

Johnsgard, Paul A. The Plovers, Sandpipers, and Snipes of the World. Lincoln: University of Nebraska Press, 1981.

Perrins, C. M., ed. Illustrated Encyclopedia of Birds: The Definitive Reference to Birds of the World. New York: Prentice Hall Editions, 1990.

Urban, E. K., C. H. Fry, and S. Keith, eds. The Birds of Africa, Vol. II. London: Academic Press, 1986.

Vaughan, R. Plovers. Lavenham: Terence Dalton Limited, 1980.

Periodicals

Bock, W. J. "A Generic Review of the Plovers (Charadriinae, Aves)." Bulletin of the Museum of Comparative Zoology, Harvard Coll. 116, no. 2 (1958): 27–97.

Jackson, B. J. S., and J. A. Jackson. "Killdeer Charadrius vociferus." The Birds of North America no. 517 (2000).

Johnson, O. W., and P. G. Connors. "American Golden Plover Pluvialis dominica, Pacific Golden Plover Pluvialis fulva." The Birds of North America no. 210–202 (1996).

Knopf, F. "Mountain Plover Charadrius montanus." The Birds of North America no. 211 (1996).

Page, G. W., J. C. Warriner, and P. W. C. Paton. "Snowy Plover Charadrius alexandrinus." The Birds of North America no. 154 (1995).

Schardien, B. J., and J. A. Jackson. "Belly-soaking as Thermoregulatory Mechanism in Nesting Killdeer." Auk 96 (1979): 604–606.

Bette J. S. Jackson, PhD