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Dunlin

The Dunlin (Calidris alpina) is a small migratory shorebird in the family Scolopacidae, renowned for its long, slightly down-curved bill adapted for probing mudflats. It measures 16–22 cm in length, has a wingspan of 36–38 cm, and weighs 49–76 g, making it one of the more compact species among calidrid sandpipers. In breeding , adults display a vivid rusty back with black edges and a distinctive black belly patch, while nonbreeding and juvenile birds adopt a more subdued gray-brown ("dun") coloration overall. This variation aids in on breeding grounds and coastal wintering sites, respectively. Dunlins breed across high-arctic and subarctic regions worldwide, including northern , , , , the , , and extending to and the , favoring moist , wet sedge marshes, and coastal grasslands. They undertake extensive migrations, with populations from wintering along the Atlantic, Pacific, and Gulf coasts from southern to , while Eurasian birds head to , northwest Africa, the , and and , often gathering in massive flocks on estuarine mudflats, lagoons, and sandy beaches. During migration, they pass over much of and , utilizing a variety of habitats including flooded fields and lake shores. These birds forage in flocks by rapidly running across exposed mudflats, using their bills to extract such as , crustaceans, and worms from just below the surface, with feeding rates increasing dramatically in groups for efficiency. Nesting occurs in shallow scrapes on the ground, typically producing four eggs per in late spring or early summer. Globally abundant with an estimated 3–7 million mature individuals, the dunlin faces declining populations due to habitat loss from coastal development, on breeding sites, and hunting pressures, leading to its classification as Near Threatened on the .

Taxonomy

Etymology and Classification

The common name "dunlin" derives from the Old English term "dunling," a compound of "dun" (referring to a dull brownish-gray color) and the diminutive suffix "-ling," meaning "little dun-colored one," which aptly describes the bird's subdued winter plumage. This name was first recorded in 1531–1532 and has been in use since to reflect the species' overall brownish tones across seasons. The genus name Calidris originates from the Ancient Greek kalidris or skalidris, a term employed by Aristotle to denote a gray-colored waterside bird, aligning with the genus's members as small, grayish shorebirds. The dunlin was originally described by in 1758 as Tringa alpina in his , placing it among the larger sandpipers in the genus . It was later reclassified into the genus within the family Scolopacidae, a change formalized in the early to better group small "stint-like" sandpipers based on morphology, with the current binomial Calidris alpina widely adopted by the mid-20th century. Historically, the dunlin was sometimes segregated with other small calidrines into the genus Erolia (e.g., as Erolia alpina in early 20th-century checklists), reflecting debates over whether such species formed a distinct lineage from larger shanks. Molecular studies in the 2010s have confirmed the dunlin's phylogenetic position within the monophyletic genus , closely related to other small sandpipers such as the ( alba), based on multi-gene analyses resolving relationships across the Scolopacidae subfamily. For instance, (mtDNA) sequencing of and control regions from global samples revealed four major clades corresponding to groups, with significant genetic differentiation (ΦST = 0.773) supporting the species' placement amid between populations. These findings resolved earlier taxonomic uncertainties, such as the separation of stint-like , leading to updates in authoritative lists like the IOC World List through version 14.1 in 2024, which maintains Calidris alpina as a single polytypic with up to 10 .

Subspecies

Ten subspecies of the dunlin are currently recognized, varying in and wintering ranges, , and . These reflect adaptations to different and environments, with genetic studies identifying five major s based on mtDNA analyses. The are as follows:

Description

Physical Characteristics

The Dunlin (Calidris alpina) is a small to medium-sized shorebird, measuring 16–22 cm in length, with a typically 32–40 cm and a body weight ranging from 45–80 g, though this can vary by , sex, and season, with individuals sometimes reaching up to 77 g during pre-migratory fattening. Measurements and bill length vary among ; for example, the Pacific subspecies C. a. pacifica has a longer bill (up to 4.5 cm) and larger body size compared to the smaller Arctic C. a. arctica. The bill is notably long and slightly drooping, typically 2.5–4 cm in length (longer in some ), adapted for probing into soft substrates. Its legs are short and black, suited for wading in shallow water, while the overall build is stocky with a short and pointed wings. Plumage varies markedly across seasons and life stages. Breeding adults exhibit striking rufous-brown upperparts with black-centered feathers on the back and scapulars, a prominent black belly patch, and white underparts often stippled with dark spots; white wing bars are visible in flight. In non-breeding , the appears more subdued, with gray-brown upperparts, a pale gray head and breast, and clean white underparts lacking the belly patch. Juveniles display a scaled pattern on the upperparts with buff-fringed black and rusty-brown feathers, pale underparts, and subtle buffy scaling on the breast, transitioning to adult-like after the post-juvenile molt. Sexual dimorphism is minimal in plumage, with both sexes sharing similar coloration patterns, though females are slightly larger than males in body size, bill length, and , a common trait among scolopacid shorebirds. The Dunlin undergoes a complete prebasic molt after that produces the winter plumage, often completed on or near the breeding grounds, and an incomplete prealternate molt prior to the next season that restores much of the vibrant summer feathers, sometimes suspended during migration. Key anatomical features enhance its foraging efficiency. The long, flexible contains specialized Herbst corpuscles at the tip, enabling tactile detection of buried prey such as in or without relying solely on , which supports nocturnal and low-light feeding. The short legs facilitate stable movement in shallow intertidal zones, preventing sinking in soft sediments while allowing rapid stitching motions during probe .

Vocalizations and Displays

The Dunlin produces a variety of vocalizations for communication, including and contact calls that alert flock members to potential threats. The primary is a high-pitched, scratchy "krree" or "cheerp," often given during flight or in response to disturbances, serving as a signal to initiate evasive maneuvers. Contact and flight calls typically consist of softer, repetitive notes such as a rapid "trr-trr" or chattering sequence, used to maintain cohesion in flocks or during . During the breeding season, male Dunlins perform elaborate displays to attract mates and defend territories, combining vocalizations with visual elements. Aerial displays involve steep ascents followed by slow, circling flights with fluttering wingbeats resembling a butterfly pattern, accompanied by a harsh, trilling of drawn-out burry notes that descend into a twittering . On the ground, males engage in territorial posturing, such as advancing with wings raised or performing bowing motions that expose the black belly patch, while emitting aggressive or short, melodic calls upon landing. These displays often incorporate wing-clapping or rapid beats to emphasize the performance. Vocalizations vary by context and location; for instance, calls in non-breeding flocks are softer and more subdued for coordination, whereas trills become louder and more aggressive during disputes. Subspecies differences are minimal. Audio analyses from the Cornell Lab of confirm variation in Dunlin calls and songs, aiding detection over or coastal habitats.

Distribution and Habitat

Breeding Distribution

The Dunlin (Calidris alpina) exhibits a circumpolar breeding distribution across high and tundra regions of the Holarctic, spanning from eastward through to , and from and across and to the . In , breeding occurs primarily on coastal from southwestern and the northward and eastward to in . Southern breeding limits extend to the for the subspecies C. a. schinzii, which also nests in southeastern , , the [Faroe Islands](/page/Faroe Islands), and . Breeding ranges vary by subspecies, with C. a. pacifica occupying lowlands along the Alaskan Pacific coast and associated , while C. a. alpina is found along the edges of Eurasian and wetlands. These distributions align with the short summer window, where brief periods of daylight and abundance support nesting and chick-rearing. Preferred nesting habitats consist of moist characterized by mosses, sedges, grasses, and shallow ponds or boggy areas interspersed with surface water, such as tussock tundra, hummocks, low ridges, and wet meadows. Dunlins avoid dense vegetation, favoring open, boggy ground that provides and access to invertebrate prey. Most breeding occurs at lower elevations near coasts. Breeding is closely tied to the ephemeral conditions of summers, but recent warming has prompted northward shifts in distribution for -breeding shorebirds, including the Dunlin, as encroachment reduces suitable open habitat at southern edges. Studies from 2020–2024 indicate these shifts are driven by advancing treelines and altered vegetation patterns, potentially contracting available breeding areas in zones. A 2025 study further shows that Dunlin and other shorebirds are advancing nesting by 0 to 0.4 days per day of warming, demonstrating adaptive responses to . Recent reports as of February 2025 highlight steep declines in Dunlin populations over the past 12 years, underscoring ongoing challenges to southern breeding ranges.

Non-breeding Distribution and Migration

The Dunlin (Calidris alpina) exhibits a broad non-breeding distribution, primarily occupying coastal mudflats, estuaries, and intertidal zones in temperate and subtropical regions worldwide during the boreal winter. Key wintering areas include the in , where large numbers of the subspecies C. a. alpina and C. a. schinzii congregate; the region in , serving as a vital site for C. a. arcticola, C. a. sakhalina, and C. a. kistchinski; and the Atlantic and Gulf Coasts of , frequented by C. a. hudsonia and C. a. pacifica. In Africa, northwest coastal sites such as Banc d'Arguin in host birds from European breeding populations, while some Asian breeders overwinter in southern , , the , or the , including for C. a. actites. Migration from and breeding grounds involves long-distance travel, with annual round-trip distances reaching up to 15,000 km for some populations, though leg distances typically range from 4,000 to 7,000 km depending on and route. North American C. a. hudsonia follows either or Mississippi Flyway southward, using staging areas like , , and for refueling, while C. a. pacifica and C. a. arcticola from may take direct overwater flights across the or head to . European C. a. schinzii migrates rapidly along coast via the to northwest , stopping at the and French coasts, whereas East Asian populations utilize the East Asian-Australasian Flyway, with the acting as a central hub for multiple . Some shorter-distance movements occur, such as Alaskan breeders wintering along the of . Southward migration typically begins post-breeding in August to October, with arrival at wintering sites by late October to December; for instance, C. a. hudsonia takes about 69 days on average, while East Asian C. a. arcticola requires around 65 days with multiple stops. Return northward occurs from April to June, often more direct and faster—lasting about 26 days for C. a. hudsonia—with departures synchronized around mid-May for East Asian subspecies. Flocks can number in the tens of thousands during passage, particularly at key stopovers like or the mudflats. Sex-specific patterns influence timing, with males departing breeding areas earlier in fall and using more direct spring routes than females in North American populations. Geolocator tracking studies from 2010 to 2019, involving over 84 individuals on the and others in , reveal route variations by subspecies, such as eastern paths for C. a. arcticola versus western for C. a. sakhalina, and highlight the 's role in connecting breeding origins to diverse wintering grounds. Challenges include the degradation and loss of stopover sites, such as intertidal mudflats in the , which threaten refueling opportunities for long-distance migrants. Ring recovery data further confirm these patterns, showing distinct connectivity for European and Asian populations.

Behavior and Ecology

Foraging and Diet

The Dunlin (Calidris alpina) employs a variety of techniques adapted to its intertidal and habitats, primarily involving rapid pecking at surface prey and deeper probing into mud or soil for buried . A distinctive "stitching" method, where the bill is inserted shallowly and moved side-to-side in a sewing-like motion, allows detection of subsurface prey through tactile cues, particularly during low when feeding grounds are exposed. These birds are predominantly diurnal , though they may feed nocturnally in certain conditions, with activity peaking in soft sediments where penetrability facilitates efficient probing. During the breeding season on Arctic tundra, the Dunlin's diet shifts to terrestrial invertebrates, dominated by insects such as midges (Chironomidae), beetles (Coleoptera), and craneflies (Tipulidae), supplemented by spiders (Araneae). This high-protein intake supports the energetic demands of reproduction, with adults consuming thousands of small items daily to achieve sufficient caloric intake for egg production and territory defense. In contrast, non-breeding diets in coastal estuaries and mudflats consist mainly of marine invertebrates, including polychaete worms (Hediste diversicolor), mollusks like the gastropod Hydrobia ulvae and bivalve Scrobicularia plana, and crustaceans such as shrimp (Crangon crangon); during severe winters, birds may opportunistically consume seeds and plant matter when invertebrate availability declines. Key adaptations enhance efficiency, notably the high density of sensory pits in tip—up to several hundred Herbst corpuscles per square millimeter—that detect from hidden prey, enabling tactile foraging even in turbid or dark conditions. Intake rates vary seasonally, averaging around 200–350 J/min of energy in estuarine habitats. Foraging observation studies from the 2010s have revealed dietary shifts, showing that migrants achieve 65% higher energy intake in compared to winter through prey selection for more profitable surface-active items.

Breeding Biology

The Dunlin exhibits a primarily monogamous , with pairs forming briefly for the breeding season on and tundra habitats. Males typically arrive at breeding grounds first in late May or early , establishing territories through aerial display flights involving short glides with stiff arched wings interrupted by rapid, shallow wing flutters. Pair bonds are territorial and last only until the female initiates a second or departs, with extra-pair copulations rarely observed. Nesting occurs in shallow ground scrapes, often on dry upland near wetlands, created by the and lined by the with lichens, , grass, or leaves for and insulation. Clutches consist of 4 olive-buff to pale green eggs, blotched with brown, measuring approximately 35–37 mm in length and 24 mm in width, laid at intervals of about one per day from late May to early depending on . begins with the completion of the clutch and lasts 21–22 days, shared by both sexes, though the often handles the initial half before departing to allow for potential renesting or sequential . Upon hatching, which peaks in early to mid-July, are precocial, , and mobile within hours, following the male parent—who assumes primary brooding and protection duties—to nearby wet areas for foraging, as the female typically deserts soon after hatch. Males lead and defend for about 21 days until , after which family units disband. at around 19–25 days old, capable of short flights but remaining dependent on the male for guidance. In some cases, females exhibit facultative , laying a second with a new mate after deserting the first, potentially increasing their reproductive output to two broods per season. Clutch success varies by predation pressure and weather, with hatching rates typically ranging from 30% to 50% in low-disturbance areas; primary losses stem from predation by arctic foxes (Vulpes lagopus) and red foxes (Vulpes vulpes), which target nests during lemming low cycles, alongside avian predators like jaegers and . Renesting is common, with 75% of females attempting a replacement after early-season failure, though success declines later due to time constraints. Overall, annual productivity is influenced by these factors, with males investing heavily in single-brood care to maximize chick survival.

Social Behavior and Movements

Dunlins exhibit highly gregarious behavior outside the season, forming flocks that enhance efficiency and predator detection. On coastal feeding grounds, groups typically range from 10 to 500 individuals, allowing coordinated probing of intertidal mudflats for . At high tide roosts, flock sizes can swell to thousands or even hundreds of thousands, with extreme concentrations reaching over a million at major estuaries like those along the . These large aggregations often engage in synchronized flight displays, characterized by rapid, twisting maneuvers that confuse aerial predators such as merlins (Falco columbarius), reducing individual capture risk through collective evasion tactics. During the breeding season, Dunlins shift to territorial behavior, with males primarily defending nesting areas against intruders to secure resources and mates. Territories vary by habitat density but typically encompass 100–200 m² around the nest site in high-density arctic tundra populations, where males patrol boundaries through aerial chases and ground displays. Following successful breeding, adults undergo post-breeding dispersal to molting sites, often moving tens to hundreds of kilometers from nesting grounds to coastal or inland wetlands rich in food resources; for instance, Alaskan breeders may relocate within a 130 km radius during this period of feather replacement. Interspecific and intraspecific interactions among Dunlins involve displays of , primarily through high-speed chases to displace rivals from prime or roosting spots, with rates varying from 0.005 to 5.08 interactions per minute depending on habitat crowding. Hybridization occurs rarely with closely related calidrids, such as the ( maritima), with apparent cases reported in . Daily movements of non-breeding Dunlins are closely tied to cycles along coastal habitats, where birds follow receding to expose areas and retreat to elevated roosts during inundation, covering distances of several kilometers in a single cycle. Seasonal shifts include broader dispersals to wintering ranges, but local vigilance remains key to survival; in flocks, individuals alternate scanning for threats, functioning as informal sentinels that alert the group to predators via alarm calls, thereby diluting risk through shared monitoring.

Conservation

The global population of the Dunlin (Calidris alpina) is estimated at 3,000,000–7,000,000 mature individuals, with the overall trend decreasing at a moderate rate of 20–29% over the past three generations (20 years; 2007–2027). Populations remain stable in the core breeding areas, but significant declines have occurred at southern edges of the range, particularly in temperate zones. In , breeding populations total approximately 1.5 million individuals across subspecies (C. a. arcticola, pacifica, and hudsonia), with trends generally steady but showing localized declines in some regions. The southern subspecies C. a. schinzii has experienced severe declines in Europe, with an estimated 90% reduction since the early 20th century. For example, in Denmark, breeding pairs numbered around 50,000 circa 1900 but had fallen to approximately 150 pairs as of recent estimates (circa 2023), reflecting broader patterns across the Baltic and western Europe where the total schinzii population in the Baltic region is now approximately 500 breeding pairs as of 2021. These subspecies-specific trends align with genetic structuring observed in long-distance migrant populations. Monitoring efforts include aerial surveys such as the Program for Regional and International Shorebird Monitoring () in and coordinated counts by through the International Waterbird Census (IWC), which track non-breeding aggregations exceeding 3 million individuals in some regions between 2020 and 2024. Data from 2015–2024 indicate annual declines of 1–5% in key southern breeding sites, contributing to the species' moderate global reduction. Climate-driven habitat shifts are influencing , with some northern populations potentially increasing due to expanding suitable as southern habitats warm and fragment. This northward range expansion may offset losses in peripheral areas, though fragmentation risks persist for isolated southern groups.

Threats and Management

The Dunlin (Calidris alpina) faces multiple threats that contribute to its ongoing population decline, primarily driven by habitat degradation and loss across its breeding, , and wintering grounds. Coastal development, including for , , and , has severely impacted intertidal mudflats, with over 65% of tidal flats in the lost since the 1950s, a critical stopover site for the along the East Asian-Australasian . exacerbates these pressures by altering breeding habitats through thaw, increased storm frequency, and shifts in , which reduce nesting success and availability. Additionally, illegal or unregulated persists in parts of the East Asian-Australasian and the , where thousands of shorebirds are harvested annually, posing a direct mortality risk during . Secondary threats include pollution from oil spills and industrial runoff, which contaminate foraging areas and affect prey populations, as well as human disturbance at key stopover sites that disrupts feeding and increases energy expenditure. Predation pressure has intensified in breeding areas due to fluctuations in (Vulpes lagopus) populations, driven by climate-induced changes in cycles that lead to predator booms and higher nest predation rates on shorebirds. The International Union for Conservation of Nature (IUCN) assesses the Dunlin as Near Threatened globally, a status upgraded in the assessment due to a moderate rapid decline of 20–29% over three generations, reflecting cumulative threats across flyways. Certain , such as C. a. schinzii in the , are regionally endangered, with populations reduced by over 50% in recent decades owing to habitat loss and low breeding success. Conservation management focuses on habitat protection and restoration through international cooperation. Key protected areas include the , a and Ramsar spanning , , and the , which supports over 10 million migratory shorebirds annually, including significant Dunlin numbers, and safeguards essential wintering and stopover habitat. The species is covered by agreements such as the Agreement on the Conservation of African-Eurasian Migratory Waterbirds (AEWA), which promotes flyway-wide protections, and the Convention on Migratory Species (), facilitating coordinated actions across 120+ range states. Restoration initiatives, such as China's national program to eradicate 90% of invasive smooth cordgrass (Spartina alterniflora) from mudflats by 2025, aim to reclaim degraded foraging areas and enhance tidal flat functionality. Ongoing research, including geolocator tagging along the East Asian-Australasian Flyway, informs targeted conservation by mapping migration routes and identifying priority sites for intervention.

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