Swallow

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This article is about the family of birds. For the action, see Swallowing. For other uses, see Swallow (disambiguation).

Swallow
Pied-winged swallow Hirundo leucosoma
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Order: Passeriformes
Suborder: Passeri
Family: Hirundinidae
Rafinesque, 1815
Genera

19, see text

The swallows, martins, and saw-wings, or Hirundinidae are a family of

Hirundinidae are known,[1] divided into 19 genera, with the greatest diversity found in Africa, which is also thought to be where they evolved as hole-nesters.[2] They also occur on a number of oceanic islands. A number of European and North American species are long-distance migrants
; by contrast, the West and South African swallows are nonmigratory.

This family comprises two subfamilies:

Pseudochelidoninae (the river martins of the genus Pseudochelidon) and Hirundininae (all other swallows, martins, and saw-wings). In the Old World, the name "martin" tends to be used for the squarer-tailed species, and the name "swallow" for the more fork-tailed species; however, this distinction does not represent a real evolutionary separation.[3] In the New World, "martin" is reserved for members of the genus Progne. (These two systems are responsible for the same species being called sand martin
in the Old World and bank swallow in the New World.)

Taxonomy and systematics

The family Hirundinidae was introduced (as Hirundia) by the French

Pnoepygidae. The two families diverged in the early Miocene around 22 million years ago.[7]

Within the family, a clear division exists between the two subfamilies, the Pseudochelidoninae, which are composed of the two species of river martins,

sister to each other. The phylogeny of the swallows is closely related to evolution of nest construction; the more basal saw-wings use burrows as nest, the core martins have both burrowing (in the Old World members) and cavity adoption (in New World members) as strategies, and the genus Hirundo and its allies use mud nests.[11]

Description

The Hirundinidae have an evolutionarily conservative body shape, which is similar across the clade, but is unlike that of other passerines.[10] Swallows have adapted to hunting insects on the wing by developing a slender, streamlined body and long, pointed wings, which allow great maneuverability and endurance, as well as frequent periods of gliding. Their body shapes allow for very efficient flight; the metabolic rate of swallows in flight is 49–72% lower than equivalent passerines of the same size.[12]

The bill of the sand martin is typical for the family, being short and wide.

Swallows have two foveae in each eye, giving them sharp lateral and frontal vision to help track prey. They also have relatively long eyes, with their length almost equaling their width. The long eyes allow for an increase in visual acuity without competing with the brain for space inside of the head. The morphology of the eye in swallows is similar to that of a raptor.[13]

Like the unrelated

Fanti sawwing, at a mean body mass of 9.4 g (0.33 oz) while the purple martin and southern martin, which both weigh in excess of 50 g (1.8 oz) on average, rival one another as the heaviest swallows.[14] The wings are long, pointed, and have nine primary feathers. The tail has 12 feathers and may be deeply forked, somewhat indented, or square-ended.[10] A long tail increases maneuverability,[15][16] and may also function as a sexual adornment, since the tail is frequently longer in males.[16] In barn swallows, the tail of the male is 18% longer than those of the female, and females select mates on the basis of tail length.[17]

Their legs are short, and their feet are adapted for perching rather than walking, as the front toes are partially joined at the base. Swallows are capable of walking and even running, but they do so with a shuffling, waddling gait.[18] The leg muscles of the river martins (Pseudochelidon) are stronger and more robust than those of other swallows.[10][18] The river martins have other characteristics that separate them from the other swallows. The structure of the syrinx is substantially different between the two subfamilies;[8] and in most swallows, the bill, legs, and feet are dark brown or black, but in the river martins, the bill is orange-red and the legs and feet are pink.[10]

The most common hirundine plumage is glossy dark blue or green above and plain or streaked underparts, often white or rufous. Species that burrow or live in dry or mountainous areas are often matte brown above (e.g. sand martin and crag martin). The sexes show limited or no sexual dimorphism, with longer outer tail feathers in the adult male probably being the most common distinction.[19]

The chicks hatch naked and with closed eyes.[20] Fledged juveniles usually appear as duller versions of the adult.[3]

Distribution and habitat

The family has a worldwide cosmopolitan distribution, breeding on every continent except Antarctica. One species, the Pacific swallow, occurs as a breeding bird on a number of oceanic islands in the Pacific Ocean,[21] the Mascarene martin breeds on Reunion and Mauritius in the Indian Ocean,[22] and a number of migratory species are common vagrants to other isolated islands and even to some sub-Antarctic islands and Antarctica.[23] Many species have enormous worldwide ranges, particularly the barn swallow, which breeds over most of the Northern Hemisphere and winters over most of the Southern Hemisphere.

Northern Rough-winged Swallow
photographed in central Maine, the northeastern limit of the species' breeding range.

The family uses a wide range of habitats. They are dependent on flying insects, and as these are common over waterways and lakes, they frequently feed over these, but they can be found in any open habitat, including grasslands, open woodland, savanna, marshes, mangroves, and scrubland, from sea level to high alpine areas.[10] Many species inhabit human-altered landscapes, including agricultural land and even urban areas. Land-use changes have also caused some species to expand their range, most impressively the welcome swallow, which began to colonise New Zealand in the 1920s, started breeding in the 1950s, and is now a common landbird there.[24]

Species breeding in temperate regions

house martins.[28]

Behaviour and ecology

A tree swallow attending its nest in a tree cavity

Swallows are excellent flyers and use these skills to feed and attract mates. Some species, such as the mangrove swallow, are territorial, whereas others are not and simply defend their nesting sites. In general, the male selects a nest site, and then attracts a female using song and flight and (dependent on the species) guards his territory. The size of the territory varies depending on the species of swallow; in colonial-nesting species, it tends to be small, but it may be much larger for solitary nesters. Outside the breeding season, some species may form large flocks, and species may also roost communally. This is thought to provide protection from predators, such as sparrowhawks and hobbies.[10] These roosts can be enormous; one winter-roosting site of barn swallows in Nigeria attracted 1.5 million individuals.[29] Nonsocial species do not form flocks, but recently fledged chicks may remain with their parents for a while after the breeding season. If a human being gets too close to their territory, swallows attack them within the perimeter of the nest. Colonial species may mob predators and humans that are too close to the colony.[30]

Diet and feeding

For the most part, swallows are insectivorous, taking flying insects on the wing.[10] Across the whole family, a wide range of insects is taken from most insect groups, but the composition of any one prey type in the diet varies by species and with the time of year. Individual species may be selective; they do not scoop up every insect around them, but instead select larger prey items than would be expected by random sampling.[31] In addition, the ease of capture of different insect types affects their rate of predation by swallows.[32] They also avoid certain prey types; in particular, stinging insects such as bees and wasps are generally avoided. In addition to insect prey, a number of species occasionally consume fruits and other plant matter. Species in Africa have been recorded eating the seeds of Acacia trees, and these are even fed to the young of the greater striped swallow.[10][33]

The swallows generally forage for prey on the wing, but they on occasion snap prey off branches or on the ground. The flight may be fast and involve a rapid succession of turns and banks when actively chasing fast-moving prey; less agile prey may be caught with a slower, more leisurely flight that includes flying in circles and bursts of flapping mixed with gliding. Where several species of swallows feed together, they separate into different

Niche separation may also occur with the size of prey chosen.[34]

Breeding

Swallow chicks nesting at the Skomer Marine Conservation Zone, 2017: Video by Natural Resources Wales
American cliff swallows
constructing mud nests

The more primitive species nest in existing cavities, for example in an old

nest parasitism.[35][36] In barn swallows, old mated males and young unmated males benefit from colonial behaviour, whereas females and mated young males likely benefit more from nesting by themselves.[37]

Pairs of mated swallows are monogamous,[38] and pairs of nonmigratory species often stay near their breeding area all year, though the nest site is defended most vigorously during the breeding season. Migratory species often return to the same breeding area each year, and may select the same nest site if they were previously successful in that location. First-year breeders generally select a nesting site close to where they were raised.[39] The breeding of temperate species is seasonal, whereas that of subtropical or tropical species can either be continuous throughout the year or seasonal. Seasonal species in the subtropics or tropics usually time their breeding to coincide with the peaks in insect activity, which is usually the wet season, but some species, such as the white-bibbed swallow, nest in the dry season to avoid flooding in their riverbank nesting habitat.[10] All swallows defend their nests from egg predators, although solitary species are more aggressive towards predators than colonial species.[40] Overall, the contribution of male swallows towards parental care is the highest of any passerine bird.[10]

Parent approaching with food
Transferring the food
A wire-tailed swallow feeding a recently fledged chick

The

incubation duties are shared in some species, and in others the eggs are incubated solely by the females. Amongst the species where the males help with incubation, their contribution varies amongst species, with some species such as the cliff swallow sharing the duties equally and the female doing most of the work in others. Amongst the barn swallows, the male of the American subspecies helps (to a small extent), whereas the European subspecies does not. Even in species where the male does not incubate the eggs, he may sit on them when the female is away to reduce heat loss (this is different from incubation as that involves warming the eggs, not just stopping heat loss). Incubation stints last for 5–15 minutes and are followed by bursts of feeding activity. From laying, swallow eggs take 10–21 days to hatch, with 14–18 days being more typical.[10]

The chicks of swallows hatch naked, generally with only a few tufts of down. The eyes are closed and do not fully open for up to 10 days. The feathers take a few days to begin to sprout, and the chicks are brooded by the parents until they are able to

thermoregulate. On the whole, they develop slowly compared to other passerine birds. The parents do not usually feed the chicks individual insects, but instead feed a bolus of food comprising 10–100 insects. Regardless of whether the species has males that incubate or brood the chicks, the males of all hirundines help feed the chicks. When the young fledge is difficult to determine, as they are enticed out of the nest after three weeks by parents, but frequently return to the nest afterwards to roost.[10]

Calls

Song of the purple martin.

Swallows are able to produce many different calls or songs, which are used to express excitement, to communicate with others of the same species, during courtship, or as an alarm when a predator is in the area. The songs of males are related to the body condition of the bird and are presumably used by females to judge the physical condition and suitability for mating of males.[41] Begging calls are used by the young when soliciting food from their parents. The typical song of swallows is a simple, sometimes musical twittering.

Status and conservation

The Bahama swallow is listed as an endangered species.

Species of hirundine that are

golden swallow, have declined due to forest loss and also competition with introduced species such as starlings and sparrows, which compete with these swallows for nesting sites. The golden swallow formerly bred on the island of Jamaica, but was last seen there in 1989 and is now restricted to the island of Hispaniola.[43]

Relationship with humans

An artificial purple martin nesting colony
birds on postage stamps around the world.[45][46][47]

Swallows are tolerated by humans because of their beneficial role as insect eaters, and some species have readily adapted to nesting in and around human habitation. The barn swallow and

house martin now rarely use natural sites. The purple martin is also actively encouraged by people to nest around humans and elaborate nest boxes are erected. Enough artificial nesting sites have been created that the purple martin now seldom nests in natural cavities in the eastern part of its range.[48]

Because of the long human experience with these conspicuous species, many myths and legends have arisen as a consequence, particularly relating to the barn swallow.

Korean folktale of Heungbu and Nolbu, which teaches a moral lesson about greed and altruism through the mending of a swallow's broken leg.[50]

During the 19th century, Jean Desbouvrie attempted to tame swallows and train them for use as messenger birds, as an alternative to war pigeons. The swallows would have a light load of course, as a laden swallow could only travel about half as far as an unladen swallow in the same trip. He succeeded in curbing the migratory instinct in young birds and persuaded the government of France to conduct initial testing, but further experimentation stalled.[49][51] Subsequent attempts to train homing behaviour into swallows and other passerines had difficulty establishing a statistically significant success rate, although the birds have been known to trap themselves in a cage repeatedly to get to the bait.[49]

According to a sailing superstition, swallows are a good omen to those at sea. This probably arose from the fact that swallows are land-based birds, so their appearance informs a sailor that he or she is close to shore.[52] An old term of venery for swallows is a "flight" or "sweep".[53]

Species list

The family contains 89 species in 21 genera.[54]

Image Genus Species
Pseudochelidon Hartlaub, 1861
Psalidoprocne
 Cabanis, 1850
Neophedina
 Roberts, 1922
Phedinopsis
 Wolters, 1971
Phedina
 Bonaparte, 1855
Riparia Forster,T, 1817
Tachycineta Cabanis, 1850
Atticora Gould, 1842
Pygochelidon Baird, SF, 1971
Alopochelidon
 Ridgway, 1903
Orochelidon Ridgway, 1903
Stelgidopteryx Baird, SF, 1858
Progne Boie, F, 1826
Pseudhirundo
 Roberts, 1922
Cheramoeca
 Cabanis, 1850
Ptyonoprogne
 Reichenbach, 1850
Hirundo Linnaeus, 1758
Delichon Moore, F, 1854
Cecropis Boie, F, 1826
Atronanus De Silva, 2018
Petrochelidon Cabanis, 1850

References

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  4. ^ Rafinesque, Constantine Samuel (1815). Analyse de la nature ou, Tableau de l'univers et des corps organisés (in French). Vol. 1815. Palermo: Self-published. p. 68.
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  25. ^ In 1878, Dr. Elliott Coues, listed titles of 182 papers dealing with the hibernation of swallows ((USGS: Northern Prairie Wildlife Research Center) "Early ideas about migration" Archived 2008-08-27 at the Wayback Machine).
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  46. ^ The American Topical Association lists more than 1000 depictions of swallows and martins on postage stamps around the world. See [1] Archived 2020-03-31 at the Wayback Machine
  47. ^ Wikibooks: World Catalogue of Stamps[permanent dead link]
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External links

Wikiquote has quotations related to Swallows.
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