(Apodidae)
Class: Aves
Order: Apodiformes
Suborder: Apodi
Family: Apodidae
Thumbnail description
Small to medium-sized, fast flying birds with narrow swept-back wings, square to deeply forked tails, small bills and feet, and sometimes distinctive screaming vocalizations
Size
3.4–9.6 in (9–25 cm); 0.2–7.6 oz (5–205 g)
Number of genera, species
18 genera; 99 species
Habitat
Wide-ranging aerial foragers, but dependent on cliffs, hollow trees, or caves for nesting and roosting sites
Conservation status
Endangered: 1 species; Vulnerable: 5 species; Near Threatened: 5 species; Data deficient: 6 species
Distribution
Nearly worldwide except for coldest and treeless areas of northern hemisphere; most numerous in warmer tropical regions
Evolution and systematics
Swifts are highly specialized, generally fast flying, aerial foragers that rarely come to earth except at nest or roost sites. There have been suggestions of a swift-swallow relationship and an affinity between swifts and hummingbirds. Caprimulgids have also been frequently mentioned as a possibly closely allied group. The view that swifts and hummingbirds are related is influenced by the greatly shortened humerus and elongated distal portion of the wings. More recent DNA evidence has supported this.
The intrafamilial relationships of the swifts have also been debated over the years. Today, consensus supports there being three subfamilies: Cypseloidinae, Chaeturinae, and Apodinae. The Cypseloidinae (two genera; 12 species), with the least derived, or shortened, humerus, are presumed to be the most primitive of the swifts and are today confined to North and South America and the West Indies. Their distinctive habit of nesting and roosting in dark damp areas near or behind waterfalls or in sea caves set them apart from the other swifts.
The Chaeturinae are often called spinetails as many have stiffened shafts with bare tips to the tail feathers and use them for support while perching on vertical surfaces. This subfamily consists of 58 species in 10 genera. It includes the bewildering array of morphologically similar swiftlets or cave swiftlets of Southeast Asia, northern Australia, and many islands in the Indian and Pacific oceans. Determining the species limits among this group has been considered to be among the most daunting in ornithology and has led to most of the variation in the number of species, from 90–99, attributed to the family. Their use of echo-location and the commercial exploitation of their nests further contributes to the uniqueness of the swiftlets.
The Apodinae have a unique lateral grasping foot where toes one and two oppose toes three and four. The Apodinae, the second largest subfamily, consists of 29 species in six genera, including the widespread genus Apus as well as five species of palm-swifts.
Physical characteristics
Swifts are true masters of the aerial environment. With their long narrow swept-back wings, their flight is rapid with frequent changes in direction to capture prey. In some, the rushing noise of their wings or screaming cries announce their presence even before they come into view. Swifts are similar in having narrow-pointed wings with seven to nine secondaries and 10 primaries, the outermost being greatly elongated and more than twice the length of the innermost. They have different tail shapes with many being slightly rounded, square, or shallowly notched, while others are deeply forked. Swifts also differ greatly in size. Body lengths vary from 3.5–9.8 in (9–25 cm) and weights from about 0.17–7.2 oz (5 g–205 g). Among the smallest are the pygmy swiftlet (Collocalia troglodytes) of the Philippines and the pygmy palm-swift (Micropanyptila furcata) in Venezuela. The white-naped swift (Streptoprocne semicollaris) of western Mexico and the purple needletail (Hirundapus celebensis) from the Celebes vie for honors as the largest, while the widespread and familiar white-collared swift (Streptoprocne zonaris) in the Neotropics and the alpine swift (Apus melba) are intermediate in mass between these extremes.
Generally, swifts are dull-colored creatures with only the chestnut-collared swift (Cypseloides rutilus) and Tepui swift (Cypseloides phelpsi) having a rich chestnut red in addition to the browns, grays, black, and varying amounts of white found in other species. Several swifts have a faint to pronounced greenish irridescent sheen to their freshly molted feathers, but this is largely unobservable except for birds in the hand. In all but a few species, males and females are not separable by external appearance and, in nearly all, there is no appreciable difference in size or mass. The bill of all swifts is small but the gape is large, facilitating capture of aerial prey.
Distribution
Swifts are decidedly more common in the tropical parts of the world where up to eight or nine resident species may occur sympatrically. However, one or more species are also found at high latitudes on all continents other than Antarctica. They are found nesting from sea level to several thousand feet/meters elevation in mountainous regions of Europe, North America, and in the Andes of South America. Three swifts, the common swift (Apus apus), alpine swift (Apus melba), and pallid swift (Apus pallidus), occur as breeding species in Europe and migrate to Africa in the winter. In North America, three species similarly migrate south in winter, two of which, the chimney swift (Chaetura pelagica) and black swift (Cypseloides niger), travel all the way to South America. The white-throated swift similarly migrates from the more northerly portions of its range but is a permanent resident in much of the southwestern United States and Mexico. Two Asiatic breeding swifts, the white-throated needletail (Hirundapus caudacutus) and fork-tailed or Pacific swift (Apus pacificus), make extended migrations to winter in Australia and New Zealand, and occasionally reaching sub-Antarctic Macquarie Island.
Habitat
Swifts occur in a wide variety of habitats from densely forested tropical areas to nearly the limit of suitable nest trees in northern Scandanavia. Others that depend more on rock cliff, or manmade equivalents, for nest sites can utilize extensive nearby open savanna or grassland habitats.
Behavior
Swifts typically leave their nesting and roost sites at dawn and in many cases do not return until dusk. Swifts spend their entire day aloft, only landing at nest sites when feeding young or at nighttime roosts. The daily foraging range of these highly mobile birds has not been adequately studied, but some species may regularly forage in excess of 30–38 mi (50–60 km) from nest or roost sites. Although they may occur singly or in small groups, swifts are highly social birds and flocks of hundreds are often found in suitable feeding areas. During migration, huge flocks of swifts can occur.
Nesting is largely dependent on the nest-type and location; some species such as the lesser swallow-tailed swift (Panyptila cayennensis) build solitary nests under rock clefts or under large limbs of forest trees. Others, notably some of the swiftlets, may nest in dense aggregations where more than 100,000 individuals utilize a single cave system. Swifts conduct much of their courtship and social activities on the wing. The common swift in Europe is noted for its ability to spend nights aloft. These same swifts make extended movements away from breeding areas during spells of cold rainy weather and seek out food by following high-pressure zones; their chicks back in the nest may drop their body temperature and go torpid until both warmer weather and the adults return. Social interactions include a lot of high-speed, usually noisy, aerial chases with intermittent glides and raised-wing displays. In spring, interlocked birds sometimes make dramatic tumbling display falls of nearly 200 ft (60 m) or more which have been interpreted, at least in the white-throated swift (Aeronautes saxatalis), as aerial copulation.
Among the most interesting capacities of the swiftlets in the genus Aerodramus is the ability to echolocate, which means they emit sharp pulses of sound, or clicks, and utilize the echo of these sounds to enable them to navigate to and from nest or roost sites in the inner regions of caves. These syringeallyproduced sounds are typically brief audible clicks ranging from 2–10 kHz in frequency, with most of the sound energy between 2 and 5 kHz. Repetition rates vary between five and 20 clicks per second. Click rates are lowest in the twilight zone at the cave entrance and increase abruptly when the swiftlets enter the zone of total darkness. The sensitivity of this echolocation system does not allow them to discriminate targets much below 0.19–0.23 in (5–6 mm), which precludes them using it to locate food items.
Vocalizations in swifts are not particularly elaborate or even musical. They range from staccato chatter or chippering in such as the chimney swift to the more drawn out screams of Apus and Aeronautes swifts. Social screaming parties are a familiar sight and sound at dawn and dusk near breeding sites of the common swift in towns throughout much of Europe. The call of the greater swallow-tailed swift (Panyptila sanctihieronymi) in southern Mexico may be an exception in that it has been described as a high clear and descending musical whistle.
Swifts show an extreme long-term fidelity to breeding sites; some are regularly or continuously used for a half century or more. In some cases, nests are used again and again by the same pair of swifts year after year, with only the addition of some new nest material. Others build less durable nests that have to be built anew each season. Swifts are long-lived and generally monogamous. Life-spans may approach 15–20 years or more; annual survival rates can exceed 80%, although they are lower in some of the highly migratory species.
Feeding ecology and diet
Swifts gather all of their arthropod food on the wing, most of which are insects supplemented by the occasional spider.
The specific items taken vary extensively from place to place and even from day to day, particularly in temperate areas where substantial differences in local weather conditions occur. Swifts tend to forage at higher altitudes and take larger food items on bright sunny, warm days when more potential prey are carried aloft in wind currents and thermals. During cooler, overcast or rainy weather, they forage lower and seek out areas of continued insect productivity, such as lakes and settling ponds. Food for the young is generally carried in the mouth and consolidated into a bolus with the addition of a sticky glue-like substance produced by seasonally enlarged salivary glands. A given bolus may contain as many as 1,500 individual insects representing perhaps 50–60 different species. Swifts are quick to exploit local abundances of prey as found in the mating swarms of mayflies, termites, ants, and other insects. The foraging ecology of cypseloidine swifts is substantially different in that they appear to specialize in feeding on the swarms of winged reproductive ants and termites. They also store these prey in the esophagus thus enabling them to forage more widely in search of patchily distributed swarms of these energy-rich prey and then return with a larger load than a single bolus. Their chicks are provisioned only once or twice a day compared to the one to three times per hour for the other swifts. However, they may receive multiple regurgitations on each return of the adults to the nest.
Reproductive biology
Breeding in tropical swifts usually coincides with the onset of the seasonal rains and a flush of insect prey. In swifts in the temperate zone, it begins with the longer and warmer days of late spring and summer, and chick provisioning and fledging occurs when aerial food supplies are near their peak. This timing is also important to adults that must increase fat reserves prior to their lengthy migrations.
The nests of swifts are highly variable and have proven useful in assessing intrafamilial relationships. The very large needletail swifts (Hirundapus) build no nest at all and lay their eggs in a scrape in the detritus at the bottom of hollow trees, and the white-naped swift utilizes a scrape in the sand on ledges in caves. The Horus swift (Apus horus) usurps burrows of bee-eaters and othe burrowing birds while two other African species, the white-rumped swift (Apus caffer) and Bates's swift (Apus batesi) utilize old swallow nests that they reline with feathers. The most elaborate structures are the tubular nests of the swallow-tailed swifts (Panyptila) of Central and South America, which are made of plant seed floss glued together with salivary cement into a felt-like material. In the chaeturine swifts in the genera Chaetura, Telecanthura, and Neafrapus, and presumably other spinetails, the nest is a neat half-saucer of twigs glued to the inside of a hollow tree. Other swifts build simple or bulky nests of plant material and feathers glued together and located on cliffs, in crevices, or, in the case of the palm swifts, attached to the hanging palm fronds. The cypseloidine swifts are unique in that their nests are a simple pad or cone of mosses or seaweed placed in a damp dark niche near or even behind waterfalls or in sea caves. The swiftlets include varying amounts of plant material and feathers in their nests along with copious amounts of salivary cement. Some such as those of the edible-nest swiftlet (Aerodramus fuciphagus) are made entirely of salivary secretions and are the highly prized ingredient of birds-nest soup and extensively harvested for this delicacy.
Swift eggs are almost uniformly ovate and glossy white in color. Clutch sizes range from only one or two in the cypseloidine swifts and swiftlets to as many as five to seven in some species of Chaetura and Aeronautes. Incubation periods tend to be long, ranging from 19–20 days to about 28 days. Newly hatched chicks are blind, helpless, and devoid of any natal down. A fluffy down-like semi-plume covering starts growing within a few days, which precedes the growth of the first typical contour plumage. Development of the chicks is slow compared to other similar-sized birds and takes from four to six and a half weeks (an extraordinary nine weeks in the spot-fronted swift [Cypseloides cherriei]). In many of the spine-tailed swifts (Chaetura), the young may leave the crowded nest and cling to the walls of the hollow tree nest site for a week or more before actually fledging. Others such as the cypseloidine swifts and cave-inhabiting swiftlets remain in the nest until fledging.
The molt period in swifts is generally prolonged with only a few flight feathers being replaced at the same time. This necessitates extensive overlap of breeding and molt periods in temperate zone species that complete their molt prior to fall migration. Others delay the molt until reaching the wintering ground, or interrupt it shortly after its start to accommodate the energy cost of migration.
Conservation status
Because of their highly specialized nesting requirements, a number of swifts have been included on various lists of threatened or vulnerable species. Destruction or disturbance of localized nesting areas, especially caves, can have an appreciable negative impact on populations of these birds, particularly island populations. A deficiency of detailed information on the breeding biology and status of a number of species of swifts has led to their being tentatively included on lists of those possibly needing protection. One species (Guam swift, Collocalia bartschi) is listed as Endangered by the IUCN. An additional five species are considered Vulnerable and another five species are deemed Near Threatened. Detailed information is badly needed for other poorly studied species.
Significance to humans
Swifts are often found nesting in close association with humans and common swifts used to be encouraged to nest in man-made rooftop compartments where the young could be harvested for food. Even today, swiftlet nests are widely harvested for food and perceived medicinal properties. The feather-free "white" nests of the edible-nest swiftlet (Aerodramus fuciphagus), which are made entirely of the sticky secretions of the salivary glands, are important economic assets throughout their range in the oriental region. So-called "black" nests of the black-nest swiftlet (Aerodramus maximus), which contain substantial amounts of feathers intermixed with the salivary secretions, are also harvested but command a much lower price. Today, the harvest of these nests is closely monitored and regulated. Properly managed, this can be a renewable resource providing a sustained yield year after year. However, illegal harvesting and excessive disturbance threaten the long-term viability of some colonies and the nest-collecting industry. Cross-fostering of young edible-nest swiftlet chicks into nests of other species as a means of establishing new colonies in places where a carefully regulated harvest will be possible is a promising new program.
Species accounts
White-collared swiftBlack swiftChimney swiftEdible-nest swiftletWhite-rumped swiftletWhite-throated needletailCommon swiftAlpine swiftAfrican palm swiftResources
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Chantler, P. Swifts: A Guide to the Swifts and Treeswifts of the World. 2nd edition. New Haven: Yale University Press, 2000. del Hoyo, J., A. Elliot, and J. Sargatul, eds. Handbook of the Birds of the World. Vol 5. Barn Owls to Hummingbirds, Barcelona: Lynx Editions, 1999.
Francis, C. M. The Management of Edible Bird's Nest Caves in Sabah. Sandikan: Wildlife Section, Sabah Forest Department, 1987.
Lack, D. Swifts in a Tower. London: Methuen and Co. Ltd., 1956.
Sibley, C. G., and J. E. Ahlquist. Phylogeny and Classification of Birds: A Study in Molecular Evolution. New Haven: Yale University Press, 1990.
Periodicals:Collins, C. T. "The Comparative Biology of Two Species of Swifts in Trinidad, West Indies." Bulletin of the Florida State Museum 11 (1968): 257–320.
Collins, C. T. "Subfamilies of the Swifts (Apodiformes: Apodidae): Schoutedenapus Revisited." Durban Museum Novitates 25 (2000): 36–39.
Collins, C. T., and R. K. Brooke. "A Review of the Genus Hirundapus (Aves: Apodidae)." Natural History Museum of Los Angeles County, Contributions in Science 282 (1976): 1–22.
Lack, D., and E. Lack. "The Breeding Biology of the Swift." Ibis 93 (1951): 501–546.
Marin A. M., and F. G. Stiles. "On the Biology of Five Species of Swifts (Apodidae, Cypseloidinae) in Costa Rica." Proceedings of the Western Foundation of Vertebrate Zoology 4 (1992): 287–351.
Medway, Lord, and J. D. Pye. "Echolocation and the Systematics of Swiftlets." Evolutionary Ecology, B. Stonehouse and C. Perrins, eds.; 226–238. Baltimore: University Park Press, 1977.
Medway, Lord. "The Swiftlets (Collocalia) of Niah Cave, Sarawak." Ibis 104 (1962): 45–66, 228–245.
Tarburton, M. K. "Breeding of the White-rumped Swiftlet in Fiji." Emu 86 (1986): 214–227.
Tarburton, M. K., and E. Kaiser. "Do Fledglings and Prebreeding Common Swifts Apus apus Take Part in Aerial Roosting? An Answer from a Radiotracking Experiment." Ibis 143 (2001): 255-263.