Curlew sandpiper

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Curlew sandpiper
Curlew Sandpiper (Calidris ferruginea) (44855173205).jpg
Adult on autumn migration starting moult out of breeding plumage, Pakistan
Calidris ferruginea, winter adult, Pak Thale.jpg
Non-breeding plumage, Thailand
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Order: Charadriiformes
Family: Scolopacidae
Genus: Calidris
Species:
C. ferruginea
Binomial name
Calidris ferruginea
(Pontoppidan, 1763)
Curlew sandpiper rangemap.svg
Range
  Nonbreeding
  Breeding
  Migration
Synonyms

The curlew sandpiper (Calidris ferruginea) is a small migratory wader. It is a long-distance migrant, breeding in the Siberian Arctic, arriving there from June and staying to August, migrating south throughout Europe and Asia, to winter mainly in Africa but also along the coasts of Australasia and Southeast Asia.

Contents

It has three main plumages that change with maturity and season. The non-breeding plumage shows brown-greyish upperparts, with the areas near the head and breast being brighter. The breeding plumage is much more striking, with the entire front tinted a deep rufous, more intensely so in males. A first-year breeder does not migrate, instead remaining in its overwintering range throughout breeding season. However, it may occasionally still moult into its breeding plumage, which is largely similar to that of adult curlew sandpiper but with white spots on the chest. The juvenile plumage is strikingly different, with a creamy-coloured streaking on the breast and pale fringes on some of the wing feathers, leading to a scaly appearance. Adults superficially resemble the larger but shorter-billed red knot in breeding plumage, and dunlin and stilt sandpiper while not breeding.

The curlew sandpiper's courtship behaviour is complex, with multiple different displays, including a ground display where the male makes a faux nest to present to the female, a graceful aerial chase accompanied with frequent trilling calls, and a ground display sometimes followed by copulation, where the male dances around the female, displaying his tail feathers and rump. The female almost always lays one clutch of eggs, usually around the end of June, which hatch after 20 days, with the chicks having fledged at about 14–20 days. The curlew sandpiper is omnivorous, foraging in wetlands in large flocks for various invertebrates, including crabs and insects.

Taxonomy and etymology

The curlew sandpiper was formally described in 1763 by the Danish ornithologist Erik Pontoppidan under the binomial name Tringa ferrugineus. [2] It was later transferred to the genus Calidris , which had been described in 1804 by the German naturalist Blasius Merrem. [3] [4] The genus name is from Ancient Greek kalidris or skalidris, a term used by Aristotle for some grey waterside birds. The specific epithet ferruginea is from Latin ferrugo, ferruginis, "iron rust", referring to the bird's colour in breeding plumage. [5] The curlew sandpiper is a monotypic taxon, meaning that it has no recognised subspecies. [4] Within the genus Calidris, the curlew sandpiper is most closely related to the stilt sandpiper (C. himantopus). [6] [7] The curlew sandpiper occasionally hybridises with sharp-tailed and pectoral sandpipers, producing the hybrids Cooper's sandpiper and Cox's sandpiper, respectively. [8] [9]

Description

The curlew sandpiper is a small wader 18–23 cm (7–9 in) in length and 44–117 g (1.6–4.1 oz) in weight. [10] It has a wingspan of 38–46 cm (15–18 in), [11] a bill length of 32–44 mm (1.3–1.7 in), and a tarsus length of 26–34 mm (1.0–1.3 in). [12] It has a black bill, dark brown irises, and dark grey to black feet. [13] All plumages have a large white patch on the rump, occasionally barred with red in some adults, and in flight show a distinctive white wing stripe along the tips from the secondary to the primary coverts. [14]

Its plumage varies seasonally. [15] The non-breeding plumage has greyish upperparts, and white underparts with some pale grey streaks; however, as feather wear increases, it slowly becomes more brown. The centres of its upperpart feathers are darker, especially on its crest, causing its upperparts to look mottled. The upper tail coverts as well as the lower rump feathers are white, with broad, pale grey rectrices. Its primaries and secondaries are dusky with white streaks along the shaft and a white tip. It has a distinctively white supercilium ("eyebrow") with grey-brown lores (between the base of the bill and the eyes) and auriculars (the feathers protecting the ear). [16] The pre-breeding moult, taking place from July to November and lasting as late as April the next year for some southern populations, is a complete moult, with all feathers being replaced. [17]

Occurring from January to May, the pre-breeding moult is partial, with only the body feathers being replaced. [18] Returning breeders are sexually dimorphic. In the female, the front side, the entire head, and the upper back are tinted variably reddish, with grey-to-dark streaks. The lower scapulars, along with the upperwing coverts and commonly the tertials, are tipped ruddy-silver with black centres, resulting in a spotty appearance. The upper tail and upper wing coverts, along with the rump and flight feathers, are all nearly identical to the non-breeding appearance, except for the central rectrices and 2 to 3 tertials, which are off-black with white or ruddy fringing. Male plumage is similar to that of the female, except with the rufous tinting being almost completely solid, and conspicuous spots above the eye and around the crown. [19]

The pre-juvenile moult of the chick occurs at the nest site from late June to July. [20] The juvenile looks similar to the adult non-breeding plumage, except for more evenly spread-out wear marks and fringing, resulting in a scaly and more buff-tinged appearance. The crown and nape are both dusky with pale creamy streaks, the nape being slightly paler, and the rectrices tapered and narrower. [21]

The post-juvenile moult occurs from October to December (and can finish as late as April) and is a partial to incomplete moult, replacing all the body feathers and some flight feathers. This moult takes place almost entirely at stopover and overwintering sites, [a] only rarely starting at breeding grounds. [23]

The pre-breeding moult of the yearling bird, which occurs from April to June, is absent or limited; the body feathers are largely or completely replaced, but few if any flight feathers are replaced. [24] The plumage of the first-time breeder is similar to the adult summer plumage, with the primary differences being white spots or tufts on the underparts, which are fully and opaquely rufous, not tinted. Sex differences are also similar to that of returning breeders. [25]

Similar species

While in breeding plumage, the curlew sandpiper can be confused with the red knot, as both are reddish on the belly. Compared to the curlew sandpiper, the red knot is larger, has paler orange-buff underparts rather than rich brick-red; has shorter, greenish legs; and is with a stockier bill, as well as a spotted and greyish rump. [26]

The non-breeding or juvenile plumage looks similar to other species of its genus, Calidris, such as the stilt sandpiper, which lacks the curlew sandpiper's distinctive white wing stripe and has longer, yellow legs and a flatter head, and the dunlin, which has a much less distinct supercilium and a straighter bill, along with a generally stouter appearance, and wingtips that barely, if at all, extend beyond the tip of its tail. The juvenile curlew sandpiper can be distinguished from the juvenile dunlin by the dunlin's distinctive brown or black spots on the sides of the upper belly. [26]

In juvenile plumage, the curlew sandpiper can be confused most easily with the juvenile female ruff, which is similar in colour and build, though it is darker, larger, and has a shorter bill. The dunlin also looks similar, but can be slightly more compact in shape and is usually (but not always) shorter-billed; it can easily be distinguished by its streaked breast and flanks. [27]

Vocalisations

Only the male sings. [28] Each song lasts for 10–15 seconds and consists of several parts, starting with several introductory notes, followed by multiple trilled doublets, a four-part phrase, and several drawn-out whine calls. [29] Both genders vocalize calls. [b] The alarm call is a whik, whik, whik repeated or wik-ki-ki-ki... used on the breeding grounds to warn of potential threats, [31] and the chase call is a shrill, musical twittering given primarily by the male but occasionally by the female during courtship flights. [32] The flight call is a chirrup that drops in the middle, usually uttered during takeoff. [33] The whine call is a loud and ascending whaay, whaay, whaay, audible hundreds of metres away, nearly identical to the stilt sandpiper's whine. [34] It is only given by the male and seems to be used for defending its territory. [34]

Distribution and habitat

Typical breeding habitat in the high arctic Bennett-Insel 3 2014-08-25.jpg
Typical breeding habitat in the high arctic

The curlew sandpiper can occur throughout large swathes of Afro-Eurasia during migration. It is notably missing east of the Verkhoyansk Range in eastern Siberia during migration, along with southern inland Asia. It breeds exclusively in the Siberian extreme high Arctic tundra from the Yamal Peninsula to the Kolyuchin Bay. [35] It breeds in coastal lowlands and offshore islands, where the severe climate with harsh cold winds and short growing period prevent the growth of shrubs, and melting snow overlying the permafrost creates boggy conditions. [36] The nest is typically placed on the margins of wet hollows or pools, or on the gentle slopes of hummocks, often close to the nests of Pluvialis plovers like grey plover or Pacific golden plover. [36]

Its non-breeding range includes coastal regions of west Africa (from Mauritania to Gabon and including Cape Verde) and southern Africa (inland throughout Mozambique to Namibia to South Africa, and north through Uganda to Kenya). It is additionally present, albeit more rarely, inland from Niger to Chad, throughout the Republic of the Congo to Angola, and the Nile from Sudan to Egypt and southeastern Tunisia. It is rarer along the coasts of Australasia, as well as inland Victoria and New South Wales. In New Zealand, it can be found on the southeastern coast of the South Island and the entire coastline of the North Island. It overwinters in coastal South Asia as well as Southeast Asia. It does not overwinter in the rest of Eurasia. The curlew sandpiper is a scarce migratory vagrant in North America, especially along the Atlantic Seaboard. [37]

During the non-breeding season, it is commonly found along sheltered mudflats, as well as salt pans and wetland edges. While inland, it occupies the edges of various bodies of water, especially if they have muddy or sandy margins; it is also found along sewage ponds. [38] It occupies coastal and inland wetlands during migration, [39] and during breeding it mostly remains in lowland tundra near bogs, depressions, or pools, away from scrub and other dense vegetation. [40]

Migration

The curlew sandpiper shows little fidelity to nesting sites, making it hard to predict where a specimen overwinters based on its breeding area. However, the reverse is not as true: adults tend to prefer overwintering in the same regions and resting at stopovers at the same points, and males are more faithful to their sites compared to females. Both sexes have their breeding plumage by the time they arrive at their breeding sites. [41]

Males depart from the breeding grounds when the female starts incubating, and females depart either when breeding fails or when the young successfully fledge. A curlew sandpiper chick migrates south soon after learning to fly; however, it does not migrate back for breeding during their first breeding season, instead remaining in their overwintering range. [42] [41] In Western Europe, post-breeding migration commences with adults arriving in July, and largely departing onward before juveniles pass through in August and September. [36]

Behaviour

A juvenile curlew sandpiper preening Curlew Sandpiper (Calidris ferruginea) (8066906558).jpg
A juvenile curlew sandpiper preening

The curlew sandpiper appears to be largely monogamous; although its breeding habits are not well studied. Its behaviour is similar to that of other waders: walking, running, and wading. [43] Its flight speed during migration has been estimated to be around 70–75 km/h (43–47 mph). [44] It practices preening and bathing, sometimes scratching its head with its claws and dipping its head in the water to clean it. It mainly roosts at high tide in large mixed-species flocks on sandspits, lagoon islets, and occasionally dunes, reefs, or beach wrack. [45]

Breeding

The curlew sandpiper forms pairs during northward migration. [46] Pairs usually arrive at their breeding site in early June and perform courtship in mid-June. The female moves around an area a few hectares in size, feeding, and the male follows her attentively, trailing her 3–6 m (10–20 ft) away, occasionally giving partial or complete songs and conducting various types of courtship behaviour. [47]

During aerial courtship, the male pursues the female in long chases, occasionally catching up. When he catches up, he lifts his wings horizontally and glides toward the female while warbling and occasionally performing his chase call. (The female also, although more rarely, performs the chase call.) The aerial chase closely resembles territorial conflicts, though the latter is more erratic, with sharper turns and gyrations. After the pair lands, the male then begins one of the ground courtship displays. [48] During the pre-nesting period, the male intermittently assumes a "courtship" stance known as the ground courtship attitude (lowered head, ruffled scapulars and back feathers, and bent legs, resulting in a hunched appearance) during feeding. It is often followed by a full ground courtship display and seems to represent increased libido in males. [49]

The nest-cup display, one of two ground courtship displays, is often preceded by courtship attitude behaviour. The male, in short bursts, moves across the tundra, eventually stepping into a small depression. Once in the depression, he settles down as if it were a nest, folding his wings over his back and raising his tail. He proceeds to wag his wings and tail side to side, occasionally crouching and kicking up lichen and moss with his claws. He then stands up, picks various bits of vegetation from around him, and deposits them by his feet in the faux nest. Even if the bit of vegetation slips, he still mimes the motion of tossing it into the nest. This sequence is typically followed by him stamping his claws or repeating the pressing motion from earlier. During the sequence, he occasionally makes whine notes and trill calls. This display is not an act of nest-building, as the male performs it several times, each in different locations. Females are usually a few metres away but do not seem to pay full attention to the display. [50]

The precopulatory display precedes copulation and is described as elaborate, beginning with both birds standing near each other. The male raises the wing closer to the female, reaching an angle of about 60°–70° with the horizontal, and raises the other wing to match it. He also raises his head and lowers and fans his tail. Following this, he gives a short series of trilling calls and sways back and forth, alternating between being behind her and in front of her. He attempts to display his fanned tail and rump to the female by keeping his back always facing her. The female, who has so far ignored him, dashes and thrusts her head toward the male's rump, occasionally overshooting such that her head ends up along the male's flank instead. After around a minute of this display, the female turns away, and the male, with his wings still held up high, approaches to mount and copulate. [51]

A curlew sandpiper egg Calidris ferruginea MHNT.jpg
A curlew sandpiper egg

The breeding grounds are occupied from June until late August. [52] Nesting sites are usually located at the edge of a marsh or pool or on dry patches of tundra, often near nests of other curlew sandpipers. Clutches consist of 3–4 eggs (average 3.76) and are typically laid over the course of three days. The first brood is regularly laid from late June to July, with especially early breeders laying eggs only 5–6 days after arriving and the majority of pairs finishing around the end of June. [53] A second brood may occur for some birds nesting in the far south of their breeding range if the first is unsuccessful. [54] Eggs are slightly pear-shaped and olive-coloured with large dark olive or brown splotches, with more splotches on the broad end, giving them a roughly continuous shade of olive-brown. [55] The female incubates the eggs alone for 20 days. [53]

At birth, the nestlings are precocial (relatively mature and mobile), covered with down, and begin to wander from the nest after drying. On average, the nestling weighs 8.2 g (0.29 oz), with a bill length of 10.8 mm (0.43 in), a tarsus length of 23.2 mm (0.91 in), and a wingspan of 17 mm (0.67 in). The rate of growth is proportional to the temperature, with lower temperatures coinciding with slower growth and vice versa. The female leads the nestlings to good feeding habitats, and they become completely independent at 14–20 days of age. By the time they fledge, the nestlings weigh on average 49.6 g (1.75 oz), with a bill length of 27.8 mm (1.09 in), tarsus length of 30.8 mm (1.21 in), and a wingspan of 94.5 mm (3.72 in). [56]

Territorial behaviour

The male curlew sandpiper is territorial, chasing off other birds, including dunlin, Baird's sandpipers, and pectoral sandpipers. [57] Territories are 1.6–4.0 hectares (4.0–9.9 acres) in size for pairs and smaller for individuals. [58] A male would perform aerial displays and whine notes to announce territory, chasing intruders up to the edge of his territory, in flight or on the ground, similar to the aerial courtship display except more erratic, jagged, and rapid. [59] He would also restrict his activity to his own territory, which also meant stopping his chase at the boundaries of their territory, suggesting awareness of his neighbour's territory. Upon concluding a chase, he would often perform a flight song while returning. [60] The aggressive ground display has only been observed once in a 1964 study, and consisted of the male lowering his head and neck parallel to the ground and pulled back towards his body, and fanning his tail before running towards the invading bird. [57]

Feeding and diet

The curlew sandpiper, like other waders, forages in large flocks of up to thousands on tidal flats, shallows, marshes, salt flats, and sewage lagoons, picking up food by sight, [61] sometimes swimming while feeding. [62] It is active throughout the entire day and night, although the frequency at which it forages at night decreases as the time for northward migration approaches. It eats insects and other small invertebrates such as crabs, molluscs, and worms, supplemented with various seeds. [63]

Survival

A juvenile curlew sandpiper Curlew Sandpiper ( Calidris ferruginea ) Juvenile ... Bridlington , Yorkshire (53214618421).jpg
A juvenile curlew sandpiper

The curlew sandpiper has an estimated annual survival rate of 75–80%, with the oldest observed individual being 19 years old. [64] Its lifespan and breeding success are not well researched. [65]

The curlew sandpiper is illegally trapped along the south-east coast of India around Point Calimere and hunted while migrating through the Yellow and East China seas. [66] Waders are also occasionally targeted in the Arabian Peninsula. [67] It is additionally hunted along the Asian–East African Flyway, however the scope of hunting is not well known. [66]

The reproductive success of this species is strongly related to the population of lemmings, specifically the West Siberian, East Siberian and Arctic lemmings. In years with fewer lemmings, predatory species such as the Arctic fox hunt Arctic-breeding waders, including the curlew sandpiper, instead of their mammalian prey. [10] [68] [69]

The curlew sandpiper is subject to infestations from parasitic worms, with a total of 154 worms collected from five curlew sandpipers as part of a migratory bird study in Tasmania. Nadejdolepis paranitidulans was the most common parasite, present in 87% of all birds collected. A total of five species of parasitic worms were observed, with four cestodes and one trematode. [70] The curlew sandpiper is susceptible to avian influenza and botulism. [66]

Conservation

The curlew sandpiper has a vast extent of occurrence at around 3,000,000 square kilometres (1,200,000 sq mi) [71] with an estimated population of 420,000 to 960,000 mature individuals. [72] BirdLife International, which provides the IUCN conservation status for birds, considers the curlew sandpiper to be a vulnerable species, [1] as over the span of 15 years the population has declined by an estimated 30 to 49%. [66] The species is protected by the Agreement on the Conservation of African-Eurasian Migratory Waterbirds. [73]

The curlew sandpiper has experienced habitat loss throughout its wintering and migratory ranges. [66] On the coast of the Yellow Sea, 65% of intertidal habitats in the Yellow Sea have been lost over the past 50 years, [74] and the amount of reclaimed land exceeds the extent of the remaining intertidal mudflats, which may have been a major cause in the population decline. The rate of reclamation has slowed since 2013 and has essentially ceased in China as of 2018, however reclamation has continued around the Korean Peninsula. Another major factor in habitat loss may also be the invasive smooth cordgrass, which may make occupied mudflats unsuitable habitats. [66]

Notes

  1. Overwintering sites are the areas where a species usually stays outside of migrating and breeding. [22]
  2. Calls are less complex and briefer than songs, and are usually uttered by both genders. [30]

References

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  17. Mlodinow & Medrano 2023, Plumages, Moults, and Structure § Second and Definitive Prebasic Molts.
  18. Mlodinow & Medrano 2023, Plumages, Molts, and Structure § Definitive Prealternate Molt.
  19. Mlodinow & Medrano 2023, Plumages, Molts, and Structure § Definitive Alternate Plumage.
  20. Mlodinow & Medrano 2023, Plumages, Molts, and Structure § Prejuvenile (First Prebasic) Molt.
  21. Mlodinow & Medrano 2023, Plumages, Molts, and Structure § Juvenile (First Basic) Plumage.
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  32. Holmes & Pitelka 1964, Vocalizations § Chase note.
  33. Holmes & Pitelka 1964, Vocalizations § Call notes.
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  37. Mlodinow & Medrano 2023, Distribution § Nonbreeding Range.
  38. Mlodinow & Medrano 2023, Habitat § Habitat in Overwintering Range.
  39. Mlodinow & Medrano 2023, Habitat § Habitat in Migration.
  40. Mlodinow & Medrano 2023, Habitat § Habitat in Breeding Range.
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  43. Mlodinow & Medrano 2023, Behavior § Locomotion.
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Bibliography