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''Aerodramus'' is a
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of small, dark, cave-nesting
bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
s in the Collocaliini
tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide use of the term in English is in the discipline of anthropology. The definition is contested, in part due to conflict ...
of the
swift Swift or SWIFT most commonly refers to: * SWIFT, an international organization facilitating transactions between banks ** SWIFT code * Swift (programming language) * Swift (bird), a family of birds It may also refer to: Organizations * SWIF ...
family. Its members are confined to
tropical The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...
and
subtropical The subtropical zones or subtropics are geographical zone, geographical and Köppen climate classification, climate zones immediately to the Northern Hemisphere, north and Southern Hemisphere, south of the tropics. Geographically part of the Ge ...
regions in southern
Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
,
Oceania Oceania ( , ) is a region, geographical region including Australasia, Melanesia, Micronesia, and Polynesia. Outside of the English-speaking world, Oceania is generally considered a continent, while Mainland Australia is regarded as its co ...
and northeastern
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
. Many of its members were formerly classified in '' Collocalia'', but were first placed in a separate genus by American
ornithologist Ornithology, from Ancient Greek ὄρνις (''órnis''), meaning "bird", and -logy from λόγος (''lógos''), meaning "study", is a branch of zoology dedicated to the study of birds. Several aspects of ornithology differ from related discip ...
Harry Church Oberholser Harry Church Oberholser (June 25, 1870 – December 25, 1963) was an American ornithologist. Biography Harry Oberholser was born to Jacob and Lavera S. Oberholser on June 25, 1870, in Brooklyn, New York. He attended Columbia University, but di ...
in 1906.ITIS standard report page for ''Aerodramus''
/ref> This is a taxonomically difficult group of very similar species. Echolocation,
DNA sequencing DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, thymine, cytosine, and guanine. The ...
and parasitic lice have all been used to establish relationships, but some problems, such as the placement of the Papuan swiftlet are not fully resolved. These swiftlets can pose major identification problems where several species occur. What distinguishes ''Aerodramus''
swiftlet Swiftlets are birds from the four genera ''Aerodramus'', ''Collocalia'', ''Hydrochous'' and ''Schoutedenapus'', which form the tribe (biology), tribe Collocaliini within the swift (bird), swift family (biology), family Apodidae. The group contain ...
s from other swifts, and indeed almost all other
bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
s, is their ability to use a simple but effective form of echolocation. This enables them to navigate within the breeding and roosting caves. The nests of ''Aerodramus'' swiftlets are constructed with
saliva Saliva (commonly referred as spit or drool) is an extracellular fluid produced and secreted by salivary glands in the mouth. In humans, saliva is around 99% water, plus electrolytes, mucus, white blood cells, epithelial cells (from which ...
as a major component. In two species, saliva is the only material used, and the nests are collected for the famous Chinese delicacy ' bird's nest soup', the over-collection of which puts pressure on the swiftlet populations.


Distribution

The range of these swiftlets is confined to tropical southern Asia, Oceania, northeastern
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
and the
Indian Ocean The Indian Ocean is the third-largest of the world's five oceanic divisions, covering or approximately 20% of the water area of Earth#Surface, Earth's surface. It is bounded by Asia to the north, Africa to the west and Australia (continent), ...
, with the greatest diversity in
Southeast Asia Southeast Asia is the geographical United Nations geoscheme for Asia#South-eastern Asia, southeastern region of Asia, consisting of the regions that are situated south of China, east of the Indian subcontinent, and northwest of the Mainland Au ...
,
Indonesia Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania, between the Indian Ocean, Indian and Pacific Ocean, Pacific oceans. Comprising over List of islands of Indonesia, 17,000 islands, including Sumatra, ...
and
Papua New Guinea Papua New Guinea, officially the Independent State of Papua New Guinea, is an island country in Oceania that comprises the eastern half of the island of New Guinea and offshore islands in Melanesia, a region of the southwestern Pacific Ocean n ...
. Several of the species are restricted to small islands, and their limited range can make them vulnerable, like the
Seychelles Seychelles (, ; ), officially the Republic of Seychelles (; Seychellois Creole: ), is an island country and archipelagic state consisting of 155 islands (as per the Constitution) in the Indian Ocean. Its capital and largest city, Victoria, ...
, Whitehead's and Guam swiftlets.Birdlife International species factsheet: ''Collocalia elaphra''.
Retrieved on 16 July 2007.

Retrieved on 24 July 2007.

Retrieved on 24 July 2007
The Mangaia swiftlet is a recently extinct species known only from fossils.


Description

''Aerodramus'' swiftlets are in many respects typical swifts, having narrow wings for fast flight, and a wide gape and small reduced beak surrounded by bristles for catching insects in flight. They have dull plumage which is mainly in shades of black, brown, and grey. Members of this genus typically have dark brown upper wings and upper body, sometimes with a paler rump, light brown underparts, a paler throat, and brownish-white under-wings with dark brown "armpits". Males and female plumages are similar in appearance, as is that of the juvenile, for those species for which it has been described; in some species the juvenile shows pale fringes to the flight feathers. The legs, as with many swifts, are very short, preventing the birds from perching, but allowing them to cling to vertical surfaces. The flight is mainly gliding due to very long primary feathers and small breast muscles. ''Aerodramus'' swiftlets, depending on species, weigh 8–35 grammes (0.28–1.23 oz) and are 9–16 centimetres (0.28–1.23 in) long. These swiftlets are very similar, and where several species occur, such as
Borneo Borneo () is the List of islands by area, third-largest island in the world, with an area of , and population of 23,053,723 (2020 national censuses). Situated at the geographic centre of Maritime Southeast Asia, it is one of the Greater Sunda ...
,
New Guinea New Guinea (; Hiri Motu: ''Niu Gini''; , fossilized , also known as Papua or historically ) is the List of islands by area, world's second-largest island, with an area of . Located in Melanesia in the southwestern Pacific Ocean, the island is ...
and the
Philippines The Philippines, officially the Republic of the Philippines, is an Archipelagic state, archipelagic country in Southeast Asia. Located in the western Pacific Ocean, it consists of List of islands of the Philippines, 7,641 islands, with a tot ...
, may not be separable in the field.


Behaviour

''Aerodramus'' swiftlets are aerial insectivores, which take prey like
flies Flies are insects of the Order (biology), order Diptera, the name being derived from the Ancient Greek, Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwin ...
on the wing. They roost and breed in caves; during the day they leave the caves to forage for food, and return to roost at night. They are monogamous and both partners take part in caring for the nestlings. Males perform aerial displays to attract females and mating occurs at the nest. The breeding season overlaps the wet season, which corresponds to an increased insect population. Clutch size depends on the location and the food source, but generally ''Aerodramus'' swiftlets lay one or two eggs. The eggs are a dull white, and are laid every other day. Many, if not all, species are colonial nesters; some build their nests in high, dark corners on cave walls. Most ''Aerodramus'' swiftlets live in the tropical
Indo-Pacific The Indo-Pacific is a vast biogeographic region of Earth. In a narrow sense, sometimes known as the Indo-West Pacific or Indo-Pacific Asia, it comprises the tropical waters of the Indian Ocean, the western and central Pacific Ocean, and the ...
region and do not migrate. These birds usually remain in one cave or other roosting/nesting site. Examples of cave sites include the Niah Caves and Gunung Mulu National Park, which are both located in
Sarawak Sarawak ( , ) is a States and federal territories of Malaysia, state of Malaysia. It is the largest among the 13 states, with an area almost equal to that of Peninsular Malaysia. Sarawak is located in East Malaysia in northwest Borneo, and is ...
,
Malaysia Malaysia is a country in Southeast Asia. Featuring the Tanjung Piai, southernmost point of continental Eurasia, it is a federation, federal constitutional monarchy consisting of States and federal territories of Malaysia, 13 states and thre ...
n
Borneo Borneo () is the List of islands by area, third-largest island in the world, with an area of , and population of 23,053,723 (2020 national censuses). Situated at the geographic centre of Maritime Southeast Asia, it is one of the Greater Sunda ...
. Characteristics of behaviour, such as what materials apart from saliva the nests contain, can be used to differentiate between certain species of ''Aerodramus''.


Echolocation

The genus ''Aerodramus'' is of special interest due to its use of echolocation. The swiftlets use this technique to navigate in darkness through the chasms and shafts of the
cave Caves or caverns are natural voids under the Earth's Planetary surface, surface. Caves often form by the weathering of rock and often extend deep underground. Exogene caves are smaller openings that extend a relatively short distance undergrou ...
s where they breed and roost at night. Apart from swiftlets, the only other avian species to use echolocation is the unrelated oilbird. The ''Aerodramus'' swiftlets' echolocating double clicks are within the normal human hearing range and up to 3 milliseconds apart, with the interval becoming shorter in darker locations. Unlike the rest of the genus (for those species which have been studied), the Atiu swiftlet, ''Aerodramus sawtelli'', and the black-nest swiftlet, ''A. maximus'', emit only single clicks. The former species also uses echolocation outside its caves. The use of echolocation was once used to separate ''Aerodramus'' from the other non-echolocating cave swiftlet genera ''Collocalia'' and '' Hydrochous'' (virtually nothing is known about '' Schoutedenapus''). However, recently, the
pygmy swiftlet The pygmy swiftlet (''Collocalia troglodytes'') is a species of Swift (bird), swift in the family Apodidae. It is Endemism, endemic to the Philippines. Its natural habitat is tropical moist lowland forests. At under , it is the world's smallest ...
, ''Collocalia troglodytes'', was discovered making similar clicking noises both inside and outside its roosting cave. It has recently been determined that the echolocation vocalizations do not agree with evolutionary relationship between swiftlet species as suggested by
DNA Deoxyribonucleic acid (; DNA) is a polymer composed of two polynucleotide chains that coil around each other to form a double helix. The polymer carries genetic instructions for the development, functioning, growth and reproduction of al ...
sequence comparison. This suggests that as in bats, echolocation sounds, once present, adapt rapidly and independently to the particular species' acoustic environment. A study suggested that the echolocation subunits were mainly located in the
central nervous system The central nervous system (CNS) is the part of the nervous system consisting primarily of the brain, spinal cord and retina. The CNS is so named because the brain integrates the received information and coordinates and influences the activity o ...
, while the subunits in the vocal apparatus were already present and capable of use before echolocation even evolved. This study supports the hypothesis of independent evolution of echolocation in ''Aerodramus'' and ''Collocalia'', with the subsequent evolution of complex behaviour needed to complement the physical echolocation system, or just possibly that the vocal apparatus-parts of the echolocation system might even be inherited from some
prehistoric Prehistory, also called pre-literary history, is the period of human history between the first known use of stone tools by hominins  million years ago and the beginning of recorded history with the invention of writing systems. The use o ...
nocturnal Nocturnality is a ethology, behavior in some non-human animals characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnality, diurnal meaning the opposite. Nocturnal creatur ...
ancestor. It has been suggested that the giant or waterfall swiftlet, ''Hydrochous gigas'', which cannot echolocate, may be descended from an echolocating ancestor.


Saliva nests

The intricately constructed
saliva Saliva (commonly referred as spit or drool) is an extracellular fluid produced and secreted by salivary glands in the mouth. In humans, saliva is around 99% water, plus electrolytes, mucus, white blood cells, epithelial cells (from which ...
nests of this swiftlet genus, which in some species contain no other material, are collected to make the delicacy bird's nest soup. They therefore command extremely high prices. Authentic bird's nest soup is made from the nests of the edible-nest swiftlet (or white-nest swiftlet), ''Aerodramus fuciphagus'', and the black-nest swiftlet, ''Aerodramus maximus''. Instead of incorporating twigs, feathers and straw like others in the genus, these two swiftlets make their nest only from strands of their gummy saliva, which harden when exposed to air. Once the nests are harvested, they are cleaned and sold to restaurants. Over the past twenty years, the high demand for the nests of these ''Aerodramus'' species has had an adverse effect on their populations. The Niah caves population of black-nest swiftlets plunged from around 1.5 million pairs in 1959 to 150,000–298,000 pairs in the early 1990s through over-harvesting. Early authors had doubts about the material used to make the nest, with whale and fish sperm and sea foam being proposed as the basis for construction. Even in the 1830s, when the use of saliva had been fairly well established, it was believed that it was only a cement to bind a sea plant which provided the bulk of the gelatinous material of the nest.


Lice

As with other taxonomically difficult groups, ectoparasites can give information on relationships.Page, R. D. M., Lee, P. L. M., Becher, S.A., Griffiths, R., Clayton D. H. (1997). "A Different Tempo of Evolution in Birds and their Parasitic Lice
Text retrieved 12 Nov 2007
A study of swiftlet parasites in northern Borneo involved transferring lice between closely related swiftlet species. The survival of lice in most of these transfers was significantly reduced in proportion to the mean difference in feather barb size between the donor and recipient species of hosts. Thus, adaptation to a particular resource on the body of the host appears to govern the specificity of swiftlet lice. In transfers where lice survived, the lice moved to different areas on the body of the host where the mean barb diameter of the feathers on which the lice occurred had the required value.


Papuan swiftlet

The Papuan swiftlet, ''Aerodramus papuensis'', has three toes instead of the usual four in this group. It has the ability to echolocate, but whereas other previously studied species use echolocation primarily while flying in their caves, the Papuan swiftlet appears to be nocturnal or
crepuscular In zoology, a crepuscular animal is one that is active primarily during the twilight period, being matutinal (active during dawn), vespertine (biology), vespertine/vespertinal (active during dusk), or both. This is distinguished from diurnalit ...
and uses echolocation while active outside at night. It uses single, not double, clicks. DNA sequence data provides strong support for a basal relationship between ''A. papuensis'' and other ''Aerodramus'' taxa and suggest that this species and the waterfall swift ''Hydrochous gigas'', are sister
taxa In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
, a relationship that would indicate
paraphyly Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
of the genus ''Aerodramus''.


Taxonomy

The genus ''Aerodramus'' was introduced (as a subgenus of '' Collocalia'') in 1906 by the American ornithologist Harry C. Oberholser. He designated the
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
as ''Collocalia innominata'' Hume. This taxon is now considered as a
subspecies In Taxonomy (biology), biological classification, subspecies (: subspecies) is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (Morphology (biology), morpholog ...
of the Himalayan swiftlet (''Aerodramus brevirostris''). The genus name combines the
Ancient Greek Ancient Greek (, ; ) includes the forms of the Greek language used in ancient Greece and the classical antiquity, ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Greek ...
''aēr'' meaning "air" with ''-dromos'' meaning "-racer" (from ''trekhō'' "to run"). The genus now contains 25 species: * Seychelles swiftlet, ''Aerodramus elaphrus'' * Mascarene swiftlet, ''Aerodramus francicus'' * Indian swiftlet, ''Aerodramus unicolor'' * Philippine swiftlet, ''Aerodramus mearnsi'' * Moluccan swiftlet, ''Aerodramus infuscatus'' – includes Sulawesi swiftlet (''A. i. sororum'') and Seram swiftlet (''A. i. ceramensis'') * Mountain swiftlet, ''Aerodramus hirundinaceus'' * White-rumped swiftlet, ''Aerodramus spodiopygius'' * Australian swiftlet, ''Aerodramus terraereginae'' * Himalayan swiftlet, ''Aerodramus brevirostris'' – includes Indochinese swiftlet (''A. b. rogersi'') * Volcano swiftlet, ''Aerodramus vulcanorum'' * Whitehead's swiftlet, ''Aerodramus whiteheadi'' * Bare-legged swiftlet, ''Aerodramus nuditarsus'' * Mayr's swiftlet, ''Aerodramus orientalis'' *
Mossy-nest swiftlet The mossy-nest swiftlet (''Aerodramus salangana'') is a species of swift in the family Apodidae. Some taxonomists consider it to be a subspecies of the uniform swiftlet. It is found in northern Borneo, the Natuna and the Derawan Islands and ...
, ''Aerodramus salangana'' * Uniform swiftlet, ''Aerodramus vanikorensis'' * Ameline swiftlet, ''Aerodramus amelis'' – split from uniform swiftlet, includes Palawan swiftlet (''A. a. palawanensis'') * Palau swiftlet, ''Aerodramus pelewensis'' * Mariana swiftlet, ''Aerodramus bartschi'' * Caroline swiftlet, ''Aerodramus inquietus'' * Tahiti swiftlet, ''Aerodramus leucophaeus'' * Atiu swiftlet, ''Aerodramus sawtelli'' * Marquesan swiftlet, ''Aerodramus ocistus'' * Black-nest swiftlet, ''Aerodramus maximus'' * Edible-nest swiftlet, ''Aerodramus fuciphagus'' – includes brown-rumped swiftlet (''A. f. vestitus'') and Germain's swiftlet (''A. f. germani'') * Three-toed swiftlet, ''Aerodramus papuensis'' There is also an extinct species known from fossils: *Mangaia swiftlet, ''Aerodramus manuoi'' (prehistoric extinction)


Fossil species

A right
ulna The ulna or ulnar bone (: ulnae or ulnas) is a long bone in the forearm stretching from the elbow to the wrist. It is on the same side of the forearm as the little finger, running parallel to the Radius (bone), radius, the forearm's other long ...
( MNZ S42799) found at the Bannockburn Formation of the Manuherikia Group near the Manuherikia River in
Otago Otago (, ; ) is a regions of New Zealand, region of New Zealand located in the southern half of the South Island and administered by the Otago Regional Council. It has an area of approximately , making it the country's second largest local go ...
,
New Zealand New Zealand () is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and List of islands of New Zealand, over 600 smaller islands. It is the List of isla ...
. Dating from the Early to Middle
Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...
( Altonian, 19-16
million years ago Million years ago, abbreviated as Mya, Myr (megayear) or Ma (megaannum), is a unit of time equal to (i.e. years), or approximately 31.6 teraseconds. Usage Myr is in common use in fields such as Earth science and cosmology. Myr is also used w ...
), probably belongs to ''Aerodramus''.


References


External links

* * {{Authority control Bird genera Cave birds Animals that use echolocation Taxa named by Harry C. Oberholser