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Pallid swift

The Pallid swift (Apus pallidus) is a small, migratory species in the swift family Apodidae, renowned for its prolonged periods aloft and insectivorous aerial foraging. Measuring 16–17 cm in length with a of 39–42 cm and a weight of 30–40 g, it exhibits a chunky build with dark brown plumage that appears paler overall than the , including lighter underparts, , and rump, accented by a conspicuous white throat patch and a subtly forked . Native to warm, arid regions, the pallid swift breeds in coastal cliffs, gorges, caves, buildings, and urban structures across (from to ), ( to northwest ), the , , and parts of the extending to , at elevations from to 2,700 m. It constructs a shallow, saucer-shaped nest of , feathers, and vegetable matter bound with , typically laying a clutch of 2–3 eggs from May to , with both parents sharing and chick-rearing duties. The species sustains itself almost entirely in flight, capturing and occasional spiders over diverse open habitats such as farmlands, wetlands, and areas, while drinking and bathing on the wing; its shrill, disyllabic call aids in distinguishing it from similar swifts. As a long-distance migrant, it winters primarily in the of and , with some eastern individuals wintering in coastal areas of the , such as , exhibiting site fidelity to wintering grounds as revealed by GPS tracking. Classified as Least Concern on the IUCN Red List, the pallid swift maintains a stable global population of 800,000–1,499,999 mature individuals (2012 estimate) across its extensive range of over 18 million km², facing no major identified threats despite localized declines in some peripheral breeding areas.

Taxonomy and systematics

Etymology and classification

The specific name pallidus derives from the Latin adjective meaning "pale" or "wan," a reference to the species' comparatively lighter plumage relative to other swifts in the genus Apus. The pallid swift was first formally described in 1870 by British ornithologist George Ernest Shelley, who placed it in the genus Cypselus as Cypselus pallidus, based on specimens from Egypt. It was subsequently reclassified into the genus Apus, reflecting modern taxonomic revisions that consolidate typical Old World swifts under this name. The species belongs to the family Apodidae and the subfamily Apodinae, which encompasses the typical swifts characterized by their aerial lifestyle and reduced legs. Within the genus Apus, the pallid swift is the sister species to the (Apus apus), with genetic analyses confirming their close phylogenetic relationship and a divergence time estimated at approximately 1.9 million years ago (95% HPD: 1.37–2.41) based on mitochondrial and markers. This proximity is further evidenced by ongoing hybridization and asymmetrical between the two species, particularly in urban colonies, as revealed by a 2022 genomic study over 300 individuals across . The pallid swift clusters closely with the in the phylogeny of swifts, showing low genetic differentiation suggestive of recent divergence.

Subspecies and relationships

The pallid swift (Apus pallidus) is classified into three subspecies within the genus Apus, reflecting regional variations in distribution and morphology. The nominate subspecies A. p. pallidus breeds from Banc d'Arguin in eastward to the , including , with migratory populations primarily wintering in the zone of and some individuals recorded in limited numbers during winter along the coastal regions of . The subspecies A. p. brehmorum occupies the southwestern Palearctic, ranging from to (excluding the Dalmatian coast) and from to northwestern , including breeding populations on the and ; it winters mainly in the central . A. p. illyricus has a more restricted breeding range along the Dalmatian coast and possibly the eastern Italian coast, with wintering grounds in the eastern . These subspecies exhibit distinct wintering ranges, contributing to their geographic isolation despite overlapping elements in the breeding areas. Morphologically, the subspecies show minor plumage variations, with A. p. brehmorum being the largest and darkest, featuring a more pronounced black-brown and outer compared to the nominate form; A. p. pallidus is the smallest and palest overall; and A. p. illyricus appears intermediate in size and coloration. Genetic studies indicate subtle differences among the , consistent with their clinal variation across the ' range, though these are not sufficient to warrant further taxonomic splits. The pallid swift maintains close interspecies relationships within the Apus genus, particularly with the (Apus apus), its sister species. A 2022 genomic study revealed widespread cryptic hybridization and between the two species across both sympatric urban colonies and more distant populations, with asymmetrical favoring pallid swifts, which displayed higher proportions of mixed ancestry including late-generation hybrids. This ongoing gene exchange, likely facilitated by shared breeding habitats in the Mediterranean, complicates morphological identification in overlap zones and suggests historical admixture dating back to the .

Physical characteristics

Morphology and identification

The pallid swift ( pallidus) measures 16–17 cm in length, with a of 39–44 cm and a body mass ranging from 30–50 g, averaging around 42 g. It possesses a bulky, stocky build with a broad head, thicker body, and relatively broad, blunt wing tips compared to more slender swifts. The is short and only shallowly forked, with the outermost feathers approximately equal in length to the adjacent ones. The plumage is predominantly dark brown on the upperparts, transitioning to paler underparts that exhibit a scaly appearance, particularly on the belly and flanks due to fine pale fringes on the feathers. A prominent white throat patch stands out against the darker body and head, which features a pale overall tone with a contrasting blackish eye-mask. The wings are long and swept-back, often appearing crescent-shaped in flight, with paler and a subtle pale panel on the greater primary coverts in some individuals. Juveniles differ from adults in having duller with fine whitish edgings on the body, wings, and tail feathers, giving a mottled appearance, along with blunter wings and shorter overall wing length. For identification, the pallid swift appears chunkier and browner than the (Apus apus), with more prominent scaling on the underparts, a paler rump, and a less deeply forked tail; it also shows a paler head with a conspicuous eye-mask. It differs from the (Tachymarptis melba) by its smaller size, lack of white on the belly or underwing, and overall darker, less robust build without the alpine's umber-brown tones and white throat extension.

Vocalizations

The primary vocalization of the pallid swift (Apus pallidus) is a shrill, piercing, wheezy scream rendered as "srreeeeerrr" or "sriiih," typically overslurred with peak frequencies around 5–6 kHz. This call is often delivered in flight during communal displays or while foraging, serving as a contact signal in gregarious flocks. Pallid swifts also produce shorter chirps or peep calls, particularly during , aerial displays, , and nest , where these higher-pitched, brief notes help coordinate interactions between pairs or repel intruders. Breeding pairs engage in antiphonal duetting, alternating screaming calls in a precise ABAB pattern without overlap, which facilitates territorial signaling and pair bonding; acoustic analysis reveals consistent individual frequency features in the terminal portion of these calls, aiding . Nestlings emit calls—high-pitched, repetitive chirps—to solicit food from parents, with recordings capturing these in contexts alongside adult flight calls. Compared to the (Apus apus), pallid swift calls are lower-pitched (initial frequency around 3.8 kHz versus 4.4 kHz), more nasal, drier, and often disyllabic with a downward in the latter part, features that assist in field identification amid mixed-species flocks. These acoustic distinctions, confirmed through multivariate analysis of frequency and temporal patterns, likely contribute to species recognition and .

Distribution and habitat

Breeding range and preferences

The pallid swift (Apus pallidus) breeds across the , encompassing from and Spain eastward to , , and , as well as from to northwestern . Its range extends into the and western as far as southern Iran, with isolated populations on the and . Distinct occupy specific parts of this distribution, such as A. p. brehmorum on the and , and A. p. pallidus in the core Mediterranean and Middle Eastern areas. In its breeding grounds, the pallid swift favors coastal cliffs, gorges, and caves for nesting sites, though it has adapted well to anthropogenic environments, increasingly utilizing urban buildings, , and old structures in regions like , , and . It forages aerially over diverse open landscapes, including farmland, wetlands, and water bodies adjacent to these nesting areas. The species occupies an altitudinal range from to 2,700 m, with breeding recorded up to 1,250 m in southern 's mountainous plateaus and 2,700 m in southwestern . Pallid swifts build shallow, half-cup nests composed of straw, plant material, and feathers, secured with sticky , which they place in sheltered crevices, rock faces, or building apertures. These nests are often constructed colonially, with multiple pairs sharing sites on cliffs or in settings, reflecting the ' tolerance for proximity and disturbance.

Migration patterns

The pallid swift (Apus pallidus) is classified as a medium-distance migrant across much of its range, with populations breeding in southern Europe, North Africa, and the Middle East undertaking seasonal movements to wintering grounds in sub-Saharan Africa, while southernmost breeders in North Africa and some Middle Eastern populations remain resident year-round. Most individuals from European and North African breeding sites are fully migratory, departing after the breeding season to avoid northern winters and returning in spring to exploit seasonal insect abundance. Migration timing varies by population and location, but northern breeders typically arrive in breeding areas from to May, with earlier arrivals (late February to ) recorded in southern regions like and . Departure from breeding sites occurs from August to , following the completion of breeding and moult, with birds reaching wintering grounds by and remaining there until or before northward resumes. This schedule aligns with the species' dependence on aerial , which peak in warmer months at higher latitudes. Western populations from Iberia and northwest Africa primarily follow routes across the into northwest Africa, then southward over the to the zone (15°–20° N) in west and central , while eastern breeders from the , , and the may cross via the or the . A 2021 GPS tracking of individuals from a colony revealed complex intra-African movements, including multiple stopovers and shifts between the , Sudanian savannah (10°–15° N), and Guinea forest zone (0°–10° N), with wintering sites distributed across rather than a single location; eastern populations occasionally winter farther east in coastal . Sahara crossings occur rapidly over 2–6 days without prolonged stopovers, covering 300–400 km per day on average. During migration, pallid swifts exhibit remarkable , capable of non-stop flights exceeding 1,000 km, such as one recorded instance of 1,858 km in approximately two days, and they remain airborne for up to eight months continuously during the non-breeding period, approaching 10 months annually when including breeding-season flights. The species is highly gregarious on passage, with concentrations of up to 10,000 individuals observed daily in key areas like the , facilitating social foraging and navigation along routes.

Behavior and ecology

Foraging and diet

The pallid swift (Apus pallidus) is an exclusively aerial forager, capturing flying insects mid-flight using its wide-gape bill while maintaining high-speed maneuvers over diverse landscapes such as urban areas, farmlands, and coastal regions. This opportunistic strategy allows it to exploit spatiotemporally variable concentrations of airborne prey, often in gregarious flocks that enhance detection and access to insect swarms. Like other swifts, it drinks on the wing by skimming low over water surfaces to scoop up droplets without perching. The diet consists primarily of flying arthropods, encompassing 139 operational taxonomic units (OTUs) from 74 families across 16 orders, with a strong emphasis on but also including airborne spiders (Araneae). Dominant orders include (ants and relatives, present in 66% of samples, primarily Formicidae at 62%), (true bugs, 58%, with at 20%), Coleoptera (, 57%, including Elateridae), and Diptera (flies, 33%). The pallid swift opportunistically consumes species, such as the dipteran Geomyza tripunctata (found in 6% of samples), contributing to natural pest regulation. Prey selection favors small arthropods measuring 2–4 mm in length, which comprise the majority of captures, followed by 4–6 mm and 6–8 mm sizes. Males tend to take larger prey items than females (: χ² = 4.871, p = 0.027), though there are no significant differences between sexes in overall richness, , or preferences. Dietary exhibits marked temporal variations during the season, reflecting adaptations to fluctuating prey availability. In June–July, dominate (up to 46% in June, mostly small Formicidae), coinciding with higher family richness but smaller average prey sizes. By August–September, there is a shift toward larger and Coleoptera (over 70% combined in September), with reduced richness. In October, Diptera become predominant (>70%), as the swift adjusts to declining overall aerial abundance toward season's end. These shifts likely result from opportunistic responses to seasonal changes in phenology and atmospheric conditions.

Breeding biology

The pallid swift breeds from mid-April to early in the Mediterranean region, arriving earlier and departing later than the (Apus apus), which enables some pairs to raise one or two clutches per season. involves aerial chases and vocal displays, with pairs showing marked fidelity to breeding colonies and nest sites across years. Clutches typically consist of 2–3 eggs, averaging 2.44 for first broods and 2.03 for second broods. lasts 20–23 days and is performed by both parents, with the female taking primary responsibility at night. Pairs often nest in colonies within buildings or cliffs, reusing sites from previous seasons. Chicks are brooded continuously for the first 8–10 days by both parents, after which brooding declines sharply as nestlings become more independent. Parents feed the young at rates that increase with brood size, delivering boluses at rates ranging from 0.7 to 2.3 times per nest per hour during the nestling period. Nestlings after 40–45 days, though the full nestling stage to airworthiness extends to 44–49 days. Breeding success for first broods is relatively high, with hatching rates around 86% and fledging success of 70–80% of eggs, yielding about 1.91 young per nest. Second or autumn broods have lower success, primarily due to adverse weather conditions like excessive or dryness, which reduce availability. Double-brooding occurs in approximately 34% of pairs in , contributing modestly to overall annual productivity given the higher failure rates of late broods.

Conservation

The global population of the pallid swift (Apus pallidus) is estimated at 800,000–1,499,999 mature individuals, based on a assessment. The subpopulation comprises 152,000–424,000 mature individuals, equivalent to approximately 76,100–212,000 breeding pairs. Overall trends have remained stable since the 2018 assessment, with no evidence of significant declines across the breeding range. In , short-term trends (2007–2018) indicate stability in key countries like , alongside slight increases in urban areas attributed to greater availability of nesting sites in buildings. Demographic factors supporting this stability include a lifespan exceeding 10 years for some individuals, as recorded through long-term ringing efforts. adult survival rates average around 70–75%, with variations linked to environmental conditions during wintering. is enhanced by the species' capacity for double-brooding in favorable years, allowing pairs to fledge additional young and contribute to maintenance. Populations are monitored primarily through programs and, more recently, GPS tracking devices deployed on adults. Studies in 2021 using GPS tags on individuals from confirmed stable wintering aggregations in western , with birds exhibiting consistent site fidelity and no signs of disruption in non-breeding habitats.

Threats and status

The Pallid swift (Apus pallidus) is currently assessed as Least Concern on the . This status, which has been in place since 2016 with no changes noted through 2021 assessments, is supported by the species' extensive extent of occurrence spanning 18,700,000 km² across , , and parts of the , coupled with a stable global population of 800,000–1,499,999 mature individuals and no detected significant declines. Threats to the pallid swift remain minor and localized, with no major population-level impacts identified to date. Habitat alterations, such as urban development and building renovations that block natural and artificial nesting cavities on cliffs and structures, can disrupt breeding sites in affected regions. Intensified agriculture and associated use may contribute to declines in aerial abundance, potentially affecting resources for aerial insectivores like the pallid swift. Hybridization with the sympatric (Apus apus) occurs in urban and Mediterranean colonies, leading to and potential genetic dilution through , as evidenced by studies revealing both historical and recent hybrid events. exacerbates these pressures by altering migration timing and reducing annual adult survival rates, particularly through Sahel rainfall anomalies that affect wintering grounds in , with survival dropping to as low as 62% in dry years. Conservation efforts for the pallid swift are integrated into broader avian protections rather than species-specific initiatives. It is safeguarded under the EU Birds Directive (Directive 2009/147/EC), which prohibits the deliberate disturbance of breeding sites and promotes habitat maintenance for wild birds across member states. The species benefits from its affinity for urban nesting opportunities, such as rooftops and eaves, which have bolstered populations amid habitat changes in human-modified landscapes. Given its stable trends and wide distribution, no dedicated recovery programs or interventions are deemed necessary at present. The future outlook for the pallid swift is generally positive but carries risks from escalating agricultural intensification, which could further diminish insect prey availability and amplify vulnerability for this aerial .

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