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Gleaning is a feeding strategy by birds and bats in which they catch invertebrate prey, mainly
arthropod Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...
s, by plucking them from foliage or the ground, from crevices such as rock faces and under the eaves of houses, or even, as in the case of ticks and lice, from living animals. This behavior is contrasted with hawking insects from the air or chasing after moving insects such as ants. Gleaning, in birds, does not refer to foraging for seeds or fruit. Gleaning is a common feeding strategy for some groups of birds, including
nuthatch The nuthatches () constitute a genus, ''Sitta'', of small passerine birds belonging to the family Sittidae. Characterised by large heads, short tails, and powerful bills and feet, nuthatches advertise their territory using loud, simple songs. Mo ...
es, tits (including chickadees),
wrens Wrens are a family (biology), family, Troglodytidae, of small brown passerine birds. The family includes 96 species and is divided into 19 genus, genera. All species are restricted to the New World except for the Eurasian wren that is widely di ...
, woodcreepers, treecreepers,
Old World flycatcher The Old World flycatchers are a large family, the Muscicapidae, of small passerine birds restricted to the Old World (Europe, Africa and Asia), with the exception of several vagrants and two species, bluethroat (''Luscinia svecica'') and north ...
s, Tyrant flycatchers, babblers,
Old World warbler The Old World warblers are a large group of birds formerly grouped together in the bird family Sylviidae. They are not closely related to the New World warblers. The family held over 400 species in over 70 genera, and were the source of much taxo ...
s, New World warblers, Vireos and some
hummingbird Hummingbirds are birds native to the Americas and comprise the Family (biology), biological family Trochilidae. With approximately 366 species and 113 genus, genera, they occur from Alaska to Tierra del Fuego, but most species are found in Cen ...
s and
cuckoo Cuckoos are birds in the Cuculidae ( ) family, the sole taxon in the order Cuculiformes ( ). The cuckoo family includes the common or European cuckoo, roadrunners, koels, malkohas, couas, coucals, and anis. The coucals and anis are somet ...
s. Many birds make use of multiple feeding strategies, depending on the availability of different sources of food and opportunities of the moment.


Techniques and adaptations

Foliage gleaning, the strategy of gleaning over the leaves and branches of trees and shrubs, can involve a variety of styles and maneuvers. Some birds, such as the
common chiffchaff The common chiffchaff (''Phylloscopus collybita''), or simply the chiffchaff, is a common and widespread leaf warbler which breeds in open woodlands throughout northern and temperate Europe and the Palearctic. It is a bird migration, migratory ...
of
Eurasia Eurasia ( , ) is a continental area on Earth, comprising all of Europe and Asia. According to some geographers, Physical geography, physiographically, Eurasia is a single supercontinent. The concept of Europe and Asia as distinct continents d ...
and the Wilson's warbler of
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
, feed actively and appear energetic. Some will even hover in the air near a leaf or twig while gleaning from it; this behavior is called "hover-gleaning". Other birds are more methodical in their approach to gleaning, even seeming lethargic as they perch upon and deliberately pick over foliage. This behavior is characteristic of the bay-breasted warbler and many vireos. Another tactic is to hang upside-down from the tips of branches to glean the undersides of leaves. Tits such as the familiar
black-capped chickadee The black-capped chickadee (''Poecile atricapillus'') is a small, nonmigratory, North American passerine bird that lives in deciduous and mixed forests. It is a member of the Paridae family (biology), family, also known as tits. It has a distin ...
are often observed feeding in this manner. Some birds, like the
ruby-crowned kinglet The ruby-crowned kinglet (''Corthylio calendula'') is a very small passerine bird found throughout North America. It is a member of the kinglet family. The bird has olive-green plumage with two white wing bars and a white eye-ring. Males have a r ...
and red-eyed vireo of North America use a combination of these tactics. Gleaning birds are typically small with compact bodies and have small, sharply pointed bills. These features are even seen in gleaning birds that are not closely related. For example, in flycatchers of the family Tyrannidae, in which some member species are more adapted for hawking insects on the wing and others for gleaning, the gleaners have bills that resemble those of tits and warblers, unlike their larger-billed relatives. Also, some members of the
woodpecker Woodpeckers are part of the bird family (biology), family Picidae, which also includes the piculets, wrynecks and sapsuckers. Members of this family are found worldwide, except for Australia, New Guinea, New Zealand, Madagascar and the extreme ...
family, particularly piculets such as the rufous piculet of Southeast Asia, are similarly adapted for gleaning, with small, compact bodies and sharp bills, rather than the long, supportive tails and wedge-shaped bills more typical of woodpeckers. Birds such as the aforementioned piculet are specialized for gleaning the bark of trees, as are nuthatches, woodcreepers, and treecreepers. Most bark-gleaners work their way up tree trunks or along branches, though nuthatches are well known as the birds that can go the opposite direction, facing down and working their way down the trunk, as well. This requires strong legs and feet on the part of the nuthatch and piculet, while birds that face upwards tend to have stiff tail feathers to prop them up. Birds often specialize in a particular niche, such as a particular stratum of forest or type of vegetation. In South and Southeast Asia, for example, the mountain tailorbird is often found gleaning in thickets and stands of
bamboo Bamboos are a diverse group of mostly evergreen perennial plant, perennial flowering plants making up the subfamily (biology), subfamily Bambusoideae of the grass family Poaceae. Giant bamboos are the largest members of the grass family, in th ...
, Abbott's babbler gleans lower-storey foliage in lowland forest, the
rufous-chested flycatcher The rufous-chested flycatcher (''Ficedula dumetoria'') is a species of bird in the family Muscicapidae. It is found in Brunei, Indonesia, Malaysia, and Thailand. Its natural habitats are subtropical or tropical moist lowland forests and subtropi ...
and brown fulvetta are birds of the mid-storey forest, the yellow-breasted warbler gleans in the mid- to upper-storey, and the greater green leafbird specializes in the upper-storey forest. The Javan white-eye is a bird of coastal scrub and
mangroves A mangrove is a shrub or tree that grows mainly in coastal saline or brackish water. Mangroves grow in an equatorial climate, typically along coastlines and tidal rivers. They have particular adaptations to take in extra oxygen and remove sal ...
, while the related black-capped white-eye is restricted to
montane forest Montane ecosystems are found on the slopes of mountains. The alpine climate in these regions strongly affects the ecosystem because temperatures lapse rate, fall as elevation increases, causing the ecosystem to stratify. This stratification is ...
. Further specialization within a habitat is associated with behaviors and morphological adaptations (physical traits of size and shape). Tiny birds are lightweight enough to hang onto the ends of twigs and pluck small prey; the
goldcrest The goldcrest (''Regulus regulus'') is a very small passerine bird in the kinglet family. Its colourful golden Crest (feathers), crest feathers, as well as being called the "king of the birds" in European folklore, gives rise to its Englis ...
of Europe and its counterpart the golden-crowned kinglet of North America exhibit this feeding style. The related common firecrest is very similar in size and shape, but slightly bulkier, and has less of a tendency to glean along twigs and more of a habit of flying from perch to perch. Having a very small bill seems to be good for taking tiny prey from the surfaces of leaves, and small-billed birds such as the
blue tit The Eurasian blue tit (''Cyanistes caeruleus'') is a small passerine bird in the tit (bird), tit family, Paridae. It is easily recognizable by its blue and yellow plumage and small size. Eurasian blue tits, usually resident bird, resident a ...
forage in broad-leafed woodlands. The long-billed gnatwren and speckled spinetail of Central and South America, and the ashy tailorbird and striped tit-babbler of South Asia, show a preference for gleaning in tangles of vines. The ash-browed spinetail of South America specializes in gleaning among
epiphyte An epiphyte is a plant or plant-like organism that grows on the surface of another plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphyt ...
s on moss-covered tree branches. Many
hummingbird Hummingbirds are birds native to the Americas and comprise the Family (biology), biological family Trochilidae. With approximately 366 species and 113 genus, genera, they occur from Alaska to Tierra del Fuego, but most species are found in Cen ...
s take small insects from flowers while probing for nectar, and some species glean actively among bark and leaves. The Puerto Rican emerald is one such hummingbird. Found only on the island of
Puerto Rico ; abbreviated PR), officially the Commonwealth of Puerto Rico, is a Government of Puerto Rico, self-governing Caribbean Geography of Puerto Rico, archipelago and island organized as an Territories of the United States, unincorporated territo ...
, the female subsists on insects and spiders, while the male has a typical hummingbird diet of nectar. Hummingbirds and other gleaners are also sometimes attracted to the sap wells created by sapsuckers. Sapsuckers, which are in the woodpecker family, drill small holes in living tree branches to get the sap flowing. The sap and the insects it attracts are then consumed, and rufous hummingbirds have been observed to follow the movements of sapsuckers and take advantage of this food source. Clusters of dead leaves also often harbor invertebrate prey, and the Bewick's wren and worm-eating warbler of North America have long bills well-suited for probing them, as do certain Asian babblers, such as the rusty-cheeked scimitar-babbler. In Central and South America, foliage-gleaners such as the red-faced spinetail and buff-throated foliage-gleaner are also examples of birds that glean clusters of dead leaves. Crevice-gleaning is a niche particular to dry and rocky habitats. Adaptations for crevice-gleaning are similar to that of bark-gleaning. Just as the Bewick's wren, as mentioned in the preceding paragraph, has a long bill suited for poking around in the small places of woods and gardens, another North American wren, the canyon wren, has an even longer bill, which allows it to probe crevices in rocky cliffs. It also has skeletal adaptations to aid it in reaching deep into small spaces. These same traits are useful for gleaning the sides of buildings, as well. Another kind of rocky habitat is found along mountain streams, where birds such as the
Louisiana waterthrush The Louisiana waterthrush (''Parkesia motacilla'') is a New World warbler, that breeds in eastern North America and winters in the West Indies and Central America. Plain brown above, it is white below, with black streaks and with buff flanks and ...
of North America and the forktails of Asia pick over stream-side rocks and exposed roots for aquatic insects and other moisture-loving prey.


Other foraging techniques

Foraging for invertebrate prey on the ground often involves gleaning the
leaf litter Plant litter (also leaf litter, tree litter, soil litter, litterfall, or duff) is dead plant material (such as leaves, bark, needles, twigs, and cladodes) that has fallen to the ground. This detritus or dead organic material and its constituen ...
of the forest floor, sometimes flicking, flipping, or scratching through dead leaves. Birds can use their bills to flick or toss dead leaves from the ground to reveal prey residing beneath. The leaftossers of Central and South America and the pittas and laughingthrushes of Asia do this. An example of a bird that employs flipping is the
ovenbird The ovenbird (''Seiurus aurocapilla'') is a small songbird of the New World warbler family (biology), family (Parulidae). This bird migration, migratory bird breeds in eastern North America and winters in Central America, many List of Caribbean ...
, a species of North American wood-warbler. It deliberately turns over leaves on the ground to search for spiders, worms, and such underneath. In other parts of the world, similar leaf-flipping behavior has been observed in unrelated birds, such as the
jungle babbler The jungle babbler (''Argya striata'') is a member of the family Leiothrichidae found in the Indian subcontinent. Jungle babblers are gregarious birds that forage in small groups of six to ten birds, a habit that has given them the popular name ...
of
India India, officially the Republic of India, is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area; the List of countries by population (United Nations), most populous country since ...
. Some birds, such as
hummingbird Hummingbirds are birds native to the Americas and comprise the Family (biology), biological family Trochilidae. With approximately 366 species and 113 genus, genera, they occur from Alaska to Tierra del Fuego, but most species are found in Cen ...
s, will use their wings to create a blast of air to roll leaves over. Other birds rake a foot through the leaf litter, like a chicken, for the same purpose. This has been observed in
buttonquail Buttonquail or hemipodes are members of a small family of birds, Turnicidae, which resemble, but are not closely related to, the quails of Phasianidae. They inhabit warm grasslands in Asia, Africa, Europe, and Australia. There are 18 species in ...
s. Some American sparrows, such as the green-tailed towhee, perform a double-scratch by raking both legs simultaneously through the leaf-litter. They then catch prey items dislodged by the disturbance. Ground-foraging birds can be very hard for humans to observe, as they often occupy densely vegetated habitat, as in the case of the Bornean wren-babbler, which specializes in gleaning leaf litter in gullies in the forest of Southeast Asia. A feeding technique that is somewhere between gleaning and hawking is where a bird flies from a perch and takes prey off foliage; this is called "sally-gleaning". The pygmy tyrants of
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
are tiny flycatchers that feed this way. The todies of the
Caribbean The Caribbean ( , ; ; ; ) is a region in the middle of the Americas centered around the Caribbean Sea in the Atlantic Ocean, North Atlantic Ocean, mostly overlapping with the West Indies. Bordered by North America to the north, Central America ...
employ a distinct version of sally-gleaning. These small birds choose a perch within their lush forest and plantation habitats in the
Greater Antilles The Greater Antilles is a grouping of the larger islands in the Caribbean Sea, including Cuba, Hispaniola, Puerto Rico, and Jamaica, together with Navassa Island and the Cayman Islands. Seven island states share the region of the Greater Antille ...
, from which they scan the undersides of leaves above them. Upon spotting an insect or spider, they fly up in an arcing sally, pluck their prey item without stopping, and complete the arcing movement to land on a new perch. An unusual feeding strategy is that of the
oxpeckers The oxpeckers are two species of bird which make up the genus ''Buphagus'', from Ancient Greek βοῦς (''boûs''), meaning "ox", and φάγος (''phágos''), meaning "eater", and family (biology), family Buphagidae. The oxpeckers were form ...
of
Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
. They perch on living animals and glean parasites from the animals' hides. On furry animals, such as buffalo,
giraffe The giraffe is a large Fauna of Africa, African even-toed ungulate, hoofed mammal belonging to the genus ''Giraffa.'' It is the Largest mammals#Even-toed Ungulates (Artiodactyla), tallest living terrestrial animal and the largest ruminant on ...
, and
donkey The donkey or ass is a domesticated equine. It derives from the African wild ass, ''Equus africanus'', and may be classified either as a subspecies thereof, ''Equus africanus asinus'', or as a separate species, ''Equus asinus''. It was domes ...
, these birds run their bills through the fur of the animal, using a scissors-like motion to extract ticks and lice from near the skin. When they pull the insect out to the end of the fur, they catch it and eat it. (On animals with bare hides, such as
rhinoceros A rhinoceros ( ; ; ; : rhinoceros or rhinoceroses), commonly abbreviated to rhino, is a member of any of the five extant taxon, extant species (or numerous extinct species) of odd-toed ungulates (perissodactyls) in the family (biology), famil ...
and
hippopotamus The hippopotamus (''Hippopotamus amphibius;'' ; : hippopotamuses), often shortened to hippo (: hippos), further qualified as the common hippopotamus, Nile hippopotamus and river hippopotamus, is a large semiaquatic mammal native to sub-Sahar ...
, oxpeckers pick at any open wounds the animals happen to have, consuming blood and pus, and possibly keeping the wounds free of maggots.) Historically, rhinoceros and other large wild mammals have been among the favored hosts, but as the populations of large mammals in the African savanna have declined in modern times, the population and range of both red-billed and yellow-billed oxpecker have also changed, and now the birds will use donkeys and domestic
cattle Cattle (''Bos taurus'') are large, domesticated, bovid ungulates widely kept as livestock. They are prominent modern members of the subfamily Bovinae and the most widespread species of the genus '' Bos''. Mature female cattle are calle ...
as hosts. There are other tactics.
Dipper Dippers are members of the genus ''Cinclus'' in the bird family Cinclidae, so-called because of their bobbing or dipping movements. They are unique among passerines for their ability to dive and swim underwater. Taxonomy The genus ''Cinclus'' ...
s forage underwater in fast-moving streams.
Common grackle The common grackle (''Quiscalus quiscula'') is a species of large icterid bird found in large numbers through much of North America. First described in 1758 by Carl Linnaeus, the common grackle has three subspecies. Adult common grackles have a ...
s have been observed to follow farmers’ plows to glean the grubs exposed in the fresh soil. Similarly, on the island of Borneo, the Bornean ground-cuckoo will follow wild pigs and
sun bear The sun bear (''Helarctos malayanus'') is a bear species in the Family (biology), family Ursidae found in the tropical forests of Southeast Asia. It is the only species in the genus ''Helarctos'' and the smallest bear species, standing nearly ...
s as they turn up soil while foraging in the forest. Brewer's blackbirds are often seen in parking lots, where they pick off dead insects from car grilles. Some hummingbirds are known to take prey items from spiderwebs.


Behavioral implications

Gleaning, like other methods of foraging, is a highly visual activity, and as such has some implications for birds. First, to see requires light, and thus time allotted to gleaning is limited to daytime. Second, while a bird focuses on examining an area for prey items, it must necessarily divert its attention from scanning its surroundings for predators. Birds that glean in tree branches will often join together in a flock, and often with other gleaners in a
mixed-species foraging flock A mixed-species feeding flock, also termed a mixed-species foraging flock, mixed hunting party or informally bird wave, is a flock (birds), flock of usually insectivorous birds of different species that join each other and move together while fora ...
. It has been shown that individual birds feeding in flocks are able to spend more time looking for food and less time looking for predators. On the other hand, it is not a universal trait of gleaning birds to join with other species or even to be gregarious with their own kind. The
leafbird The leafbirds (Chloropseidae) are a family of small passerine bird species found in the Indian Subcontinent and Southeast Asia. They were formerly grouped with the ioras and fairy-bluebirds in the family Irenidae. As presently defined, the leaf ...
s of Asia are foliage-gleaners, but are often found singly or in pairs. Also, where multiple species of gleaning birds forage in the same area, they may show niche segregation; for example, one species may stick to conifers while another species inhabits broadleaf trees, or they may even divide up a habitat, with smaller species feeding among higher, smaller tree branches and larger species staying on lower, larger branches.


References

{{Wiktionary, gleaning, glean Bird behavior Bird feeding Ornithology