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Black-bellied Plover
Pluvialis squatarola
Order
CHARADRIIFORMES
– Family
CHARADRIIDAE
Authors: Paulson, Dennis R.

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Behavior

Adult male Black-bellied Plover performing a nest distraction display, Seward Peninsula, AK, June
Black-bellied Plover, flock in non-breeding plumage; California, January

Locomotion

Walking, Hopping, Climbing, Etc

Runs rather than walks; speed not determined but rapid. Typical stride 7.5–10 cm, extreme 15 cm (Bent 1929). Like other shorebirds, stands on 1 foot while roosting and hops on 1 foot readily. Wades in shallow water but does little foraging there.

Flight

One of fastest and most powerful fliers among shorebirds, able to fly with ease into strong winds. Observed flight speed 38–80 km/h (McNeil 1969). Wings in normal flight average 4 beats/s (Bent 1929). Flocks vary from densely packed to more open, even in lines.

Swimming And Diving

Swims short distances readily, floating high at rear like gull (F. M. Weston in Bent 1929).

Self-Maintenance

Preening, Head-Scratching, Stretching, Bathing, Anting, Etc

Squats while bathing, splashing water, fluttering wings, and dipping head; then shakes water out of plumage, flapping wings and wagging tail, then long period of careful preening. Bathes communally at times, with each individual flying up short distance and splashing into water, or hopping through water on 1 leg (F. H. Allen in Bent 1929).

Sleeping, Roosting, Sunbathing

Inactive at high-tide roosts but may not be sleeping; ready for immediate flight if disturbed. Roosts on 1 (typically) or 2 legs or occasionally on belly, typically with bill exposed, facing wind and head tucked in. At times, however, uses bill-in-scapulars position more typical of shorebirds (Paulson 1993).

Daily Time Budget

Other than correlation with tidal cycles, not studied. For time budget while foraging, see Food Habits: feeding.

Agonistic Behavior

Physical Interactions

Rarely observed; displays seem adequate to avoid them.

Communicative Interactions

Threat displays between fall migrants include moving toward one another in stiff walk; one bird bows deeply with head stiffly down in line with body, back ruffled, tail fanned and raised, and breast feathers fluffed. Birds crouch in this position and then, standing stiffly, peck at ground and flick pebbles or bits of weed over shoulder. They edge around each other, crouching and bowing, wrists partly lowered and tail canted and mostly spread; they then stop and flick pebbles again (Drury 1961). More directly, a bird ruffles its neck feathers, crouches, and runs toward another, which withdraws (Michael 1935). Also display at one another with entire body tilted forward, legs bent at ankle, and bills almost touching or touching substrate; near wing partially opened with black primaries conspicuous and tail tilted and fanned with white rump conspicuous to opponent; back feathers only slightly ruffled; may go on >1 min (DRP). Ground displays similar to those of American Golden-Plover (Drury 1961).

Spacing

Territoriality

Breeding territories maintained by flight and ground displays by males fiercely aggressive to other males. Male flies over nesting area at ≥ 25–30 m, with shallow wingbeats that are slow, hesitant, and above body plane. May last several minutes and cover >1 ha, not necessarily confined to area directly above territory. “Butterfly” flight shared by many aerial displayers.

Based on breeding-bird density in plots, average territory in Kuparuk Oilfield in n. Alaska has radius of 412 m (C. Moitoret pers. comm.). Of 40 nests at Hooper Bay, AK, none closer than 0.4 km (Brandt 1943).

On breeding grounds, interspecific territoriality frequent with both American and Pacific golden-plovers, including both aerial chasing and displaying and ritualized border displays; both sexes involved (Connors et al. 1993, DRP). Nest-location maps of sympatric populations (C. Moitoret pers. comm.) show lack of overlap in Black-bellied Plover and American Golden-Plover territories. Also territorial with Greater Golden-Plover (Flint and Kondratiev 1977).

Winter territoriality occurs widely (Myers et al. 1979, Myers and McCaffery 1984). Winter territories with mean length of 125 m (range 80–155 m) along linear habitats on California coast (Michael 1935, Myers et al. 1979). Mean distance between adjacent foraging plovers 96 m in Florida and 269 m in Georgia (Stinson 1977, 1980). Territory size in Europe 200–600 m2(Cramp and Simmons 1983). In spring, both adults and yearlings defend moving territories, foraging within 2–3 m of each other but behaving aggressively as another approaches; or alternatively, feed in fairly dense aggregations, with distances no more than 1–2 m and with no aggression (DRP).

Detailed studies at Teesmouth, U.K. (Townshend et al. 1984, Townshend 1985), showed both long-term (one-third of individuals) and short-term territory holders as well as nonterritorial individuals in same population of adults. Behavior persisted over successive years after first winter. Time spent in aggression between territorial birds higher in autumn than winter, perhaps for early establishment of territory.

Nonbreeding territories in England established soon after arrival, as early as Aug. Juveniles arrive later than adults and move repeatedly to establish territories. Territorial individuals increase during winter as new adults arrive from other areas; some of them evict juveniles from their territories. Larger juveniles more likely to acquire territories and remain for winter (Evans and Pienkowski 1984). In some areas, only territorial individuals, which limit foraging population; in other areas, territoriality absent. Color-marked individuals usually found on same feeding sites at same stage of tidal cycle on different days (Evans 1976). Territories maintained while foraging at night (Turpie and Hockey 1993).

Aggressive encounters against conspecifics 57, American Golden-Plovers 17, Ruddy Turnstones (Arenaria interpres) 4, and Least Sandpipers (Calidris minutilla) 3 times in Manitoba (Wishart et al. 1981). Aggressive in mixed shorebird flocks in New Jersey, intraspecifically and toward Ruddy Turnstones, Red Knots (Calidris canutus), and Sanderlings (C. alba), but not toward Semipalmated Plovers (Charadrius semipalmatus), dowitchers (Limnodromus spp.), and Semipalmated Sandpipers (Calidris pusilla) (Burger et al. 1979). Aggressive behavior consistently against Semipalmated Plovers and once against golden-plovers in Peru (Myers and McCaffery 1984).

Individual Distance

Somewhat evenly dispersed in winter habitats, most dispersed of 8 species studied in Georgia, with mean flock size 1.02 (Stinson 1980). Spacing may be by mutual avoidance or aggressive behavior. In Georgia and at Lindisfarne, U.K., birds avoid one another (Stinson 1977, Townshend et al. 1984), whereas at Teesmouth, U.K., and elsewhere in range, behave aggressively in order to maintain permanent territories. Higher densities of both conspecifics and other shorebirds at site where territorial may be responsible for differences observed in British studies (Townshend et al. 1984).

Sexual Behavior

Mating System And Sex Ratio

Apparently always monogamous. Sex ratio in juvenile specimens Aug–Nov: 56 males: 44 females (n = 492); all ages Dec–Feb: 52:48 (n = 299) (specimen data). Highly skewed to males in spring specimens, probably because more conspicuous.

Pair Bond

Only male displays; see Territoriality for description of display flight. Descends to ground by gliding with wings straight out or somewhat upturned and neck outstretched. May break into zigzag flight, ascend again, then dash abruptly to ground. Usually then displays on ground by running stiffly toward female standing 1–10 m away. Male rushes by or stops abruptly and stands still before her, with tail up and bill down, nearly to ground, or turns away and lowers and fans tail. Copulation occasionally attempted following this rush (Drury 1961, Parmelee et al. 1967, Flint and Kondratiev 1977).

Male typically runs quickly toward mate with head extended or not and wings drooped, straight onto her back, remains about 20–30 s while copulation occurs, then flies to distance of about 10–40 m. No special female posture or display noted (Hussell and Page 1976). Pairs also participate in Joint Run before copulation (Flint and Kondratiev 1977). Copulation noted as late as 17 Jun on Victoria I. and 21–22 Jun on Devon I. (Parmelee et al. 1967, Hussell and Page 1976).

Pair bond maintained throughout nesting period or until one sex (perhaps only the female) leaves area. Remating relatively frequent in consecutive seasons in small, disjunct breeding population in Alaska (O. W. Johnson pers. comm.).

Social And Interspecific Behavior

Degree Of Sociality

Forages singly, flocks to roost and fly, both while resident and during migration. Flying flocks typically <20 but may exceed 100. Roosting flocks typically <100, rarely exceed 1,000; 3,000 in one spring flock in Alberta. Mean 382 in monospecific flocks, 309 in mixed-species flocks in New York study (Burger and Gochfeld 1983). Sociality primarily intraspecific but flocks at times with many other plovers and sandpipers, in size from stints to curlews, godwits, and oystercatchers. Commonly associates with Dunlin (Calidris alpina) throughout winter, also Red Knot and Short-billed Dowitcher (Limnodromus griseus) in migration, in Pacific Northwest (Paulson 1993).

Nonpredatory Interspecific Interactions

Usually nests well away from other species, but one nest within 3 m of Arctic Tern (Sterna paradisaea) nest (Conover 1926). Two female Buff-breasted Sandpipers (Tryngites subruficollis) nested very near pair, probably protected from avian predators by plovers’ vigilance and mobbing behavior (Paulson and Erckmann 1985). Plovers vigorously chased both sexes of sandpiper when encountered near nest, probably behavior directed toward any intruder, as White-rumped Sandpipers (Calidris fuscicollis) and Red Phalaropes (Phalaropus fulicaria) also chased (Mayfield 1973). Golden-plovers chase Black-bellied Plovers (Drury 1961), and Ruddy Turnstones and Black-bellied Plovers fight (DRP); motivation not clear but may be same type of response.

Whimbrels (Numenius phaeopus), Ruddy Turnstones, Bonaparte’s (Larus philadelphia), Common Black-headed (L. ridibundus), Ring-billed (L. delawarensis), Herring (L. argentatus), and Glaucous-winged (L. glaucescens) gulls, and American Crows (Corvus brachyrhynchos) take prey from Black-bellied Plovers (Payne and Howe 1976, Schneider and Harrington 1981, Wishart et al. 1981, Ferns 1989, J. Withgott pers. comm., DRP). In turn, Black-bellied Plovers kleptoparasitize or attempt to kleptoparasitize Nordmann’s Greenshanks (Tringa guttifer), Ruddy Turnstones, and Dunlins (Howes and Lambert 1987, Ferns 1989). In Mauritania, takes prey from Whimbrels and Eurasian Oystercatchers (Haematopus ostralegus); plover approached oystercatcher at night and jumped against it as it picked out a piece of cockle, oystercatcher dropped prey, and plover retrieved it (Swennen 1990). Plovers also cleaned Arca senilis shells opened by oystercatchers (Piersma 1982).

S. F. Rathbun (in Bent 1929) stated that Red Knots use Black-bellied Plovers as sentinels, feeding near them and taking flight when they do. The 2 associate consistently in migration on the New York coast (Burger and Gochfeld 1983). Black-bellied Plover alarm calls elicit predator-evasion behavior in Dunlins (Buchanan 1989).

Predation

Kinds Of Predators And Manner Of Predation

Arctic birds (i.e., jaegers [Stercorarius spp.] and Glaucous Gull [Larus hyperboreus]) and mammals (arctic fox [Alopex lagopus]) take eggs and young; bird-eating hawks and owls take adults. Predators seen capturing or eating adults include Peregrine Falcon (Falco peregrinus) and Snowy Owl (Nyctea scandiaca); one being eaten by a Northern Harrier (Circus cyaneus) may have been captured or taken from another raptor (DRP).

Response To Predators

Adults of both sexes vigorously mob all 3 jaeger species and Herring and Glaucous gulls that overfly their territories (Drury 1961, Flint and Kondratiev 1977, Paulson and Erckmann 1985). A male struck a Long-tailed Jaeger (Stercorarius longicaudus) vigorously in midair (Brandt 1943). At Devon I., adults alarm-called but did not mob overflying Glaucous Gulls and Long-tailed Jaegers but did mob flying Parasitic Jaegers and a Long-tailed Jaeger that landed nearby (Hussell and Page 1976).

Incubating and brooding individuals leave nest well in advance of human approach, often at warning of mate and usually with alarm calls (at distances as much as 200–250 m; Conover 1926, Drury 1961, Hussell and Page 1976, Flint and Kondratiev 1977). Once nest discovered, birds may stay at distances exceeding 100 m (Drury 1961), or 1 or both parents may return and display (Parmelee et al. 1967); probability of this increases with incubation time (Drury 1961). Off-nest bird typically flies to meet intruder with alarm calls, brooding adult then joins mate, and chicks scatter and freeze (Hussell and Page 1976). Male more likely to display and dis-plays more aggressively than female (Drury 1961). Drury (1961) noted close approach only when eggs were about to hatch, but at one nest on about tenth day of incubation, male flushed at 50 m and both sexes came to display, then male returned to incu-bate while observer lying down in view at 10 m (DRP).

Distraction display variable, with intensity varying among individuals. According to Drury (1961), low intensity involves head-up alarm and displacement feeding. This followed by running 10–25 m with head lowered, stretched far forward, wings drooped, tail barely fanned and tilted toward intruder, back feathers ruffled; frequently settles with fast squirming, as if on eggs, but head along ground as if to hide. At higher intensity, prostrate, breast on ground in hollow, bill thrust forward with chin along ground; tail cocked and fanned, wings almost fully spread, beating spasmodically 1 beat/s), primaries brushing ground noisily; feet stamping though obscured; usually faces to one side of intruder, head turned to one side. During pursuit runs off, rolling to side, with wings folded and lowered, tail spread. Höhn (1957), Parmelee et al. (1967), and Flint and Kondratiev (1977) observed similar displays. At 3 nests, male gave typical low-intensity running displays and female typical high-intensity flapping displays several times (Drury 1961, DRP). Sham-incubating and -feeding frequent (Drury 1961).

Incubating male treated dead jaeger in lifelike pose 1 m from nest with alarm (called, flew over it), same bird lying on side with less intensity (no flights), and similar-sized brown parka with little concern (Höhn 1957).

In nonbreeding season, Merlins (Falco columbarius) elicit evasion flights (Buchanan 1989). Black-bellied Plover’s high level of alertness legendary among market hunters; this species thought to have changed stopover points in migration to avoid hunters (Bent 1929).