Antbird

The antbird family Thamnophilidae used to be considered a subfamily, Thamnophilinae, within a larger family Formicariidae that included antthrushes and antpittas. Formerly, that larger family was known as the "antbird family" and the Thamnophilinae were "typical antbirds". In this article, "antbird" and "antbird family" refer to the family Thamnophilidae. Thamnophilidae was removed from Formicariidae, leaving behind the antthrushes and antpittas, due to recognition of differences in the structure of the breastbone (sternum) and syrinx, and Sibley and Ahlquist's examination of DNA–DNA hybridization. The Thamnophilidae antbirds are members of the infraorder Tyrannides (or tracheophone suboscines), one of two infraorders in the suborder Tyranni. The Thamnophilidae are now thought to occupy a fairly basal position within the infraorder, i. e. with regard to their relatives the antthrushes and antpittas, tapaculos, gnateaters, and also the ovenbirds. The sister group of the Thamnophilidae is thought to be the gnateaters. The ovenbirds, tapaculos, antthrushes and antpittas are thought to represent a different radiation of that early split. The antbird family contains over 230 species, variously called antwrens, antvireos, antbirds and antshrikes. The names refer to the relative sizes of the birds (increasing in the order given, though with exceptions) rather than any particular resemblance to the true wrens, vireos or shrikes. In addition, members of the genus Phlegopsis are known as bare-eyes, Pyriglena as fire-eyes and Neoctantes and Clytoctantes as bushbirds. Although the systematics of the Thamnophilidae is based on studies from the mid-19th century, when fewer than half the present species were known, comparison of the myoglobin intron 2, GAPDH intron 11 and the mtDNA cytochrome b DNA sequences has largely confirmed it. There are two major clades – most antshrikes and other larger, strong-billed species as well as Herpsilochmus, versus the classical antwrens and other more slender, longer-billed species – and the monophyly of most genera was confirmed. The Thamnophilidae contains several large or very large genera and numerous small or monotypic ones. Several, which are difficult to assign, seem to form a third, hitherto unrecognised clade independently derived from ancestral antbirds. The results also confirmed suspicions of previous researchers that some species, most notably in Myrmotherula and Myrmeciza, need to be assigned to other genera. Still, due to the difficulties of sampling from such a large number of often poorly known species, the assignment of some genera is still awaiting confirmation. == Morphology ==

antbirds are stable and adapted to gripping vertical stems and branches. The leg muscles of the bicoloured antbird make up 13 % of the total body weight. The antbirds are a group of small to medium-sized passerines that range in size from the large giant antshrike, which measures 45 cm (18 in) and weighs 150  g (5.29  oz), to the tiny 8-cm (3 in) pygmy antwren, which weighs 7  g (0.25 oz). In general terms, "antshrikes" are relatively large-bodied birds, "antvireos" are medium-sized and chunky, while "antwrens" include most smaller species; "antbird" genera can vary greatly in size. Members of this family have short rounded wings that provide good manoeuvrability when flying in dense undergrowth. The legs are large and strong, particularly in species that are obligate ant-followers. These species are well adapted to gripping vertical stems and saplings, which are more common than horizontal branches in the undergrowth, and thus the ability to grip them is an advantage for birds following swarms of army ants. The claws of these antbirds are longer than those of species that do not follow ants, and the soles of some species have projections that are tough and gripping when the foot is clenched. Tarsus length in antbirds is related to foraging strategy. Longer tarsi typically occur in genera such as the Thamnophilus antshrikes that forage by perch-gleaning (sitting and leaning forward to snatch insects from the branch), whereas shorter tarsi typically occur in those that catch prey on the wing, such as the Thamnomanes antshrikes. Many species of antbirds have a contrasting 'patch' of white (sometimes other colours) feathers on the back (known as interscapular patches), shoulder or underwing. This is usually concealed by the darker feathers on the back but when the bird is excited or alarmed these feathers can be raised to flash the white patch. dot-winged antwrens puff out white back patches, whereas in bluish-slate antshrikes and white-flanked antwrens the white patch is on the shoulder. Voice calling The songs and calls of antbirds are generally composed of repeated simple uncomplicated notes. The family is one of the suboscines (suborder Tyranni) which have simpler syrinxes ("voiceboxes") than other songbirds. Nevertheless, their songs are distinctive and species-specific, allowing field identification by ear. Antbirds rely on their calls for communication, as is typical of birds in dark forests. Most species have at least two types of call, the loudsong and the softsong. The functions of many calls have been deduced from their context; for example some loudsongs have a territorial purpose and are given when birds meet at the edges of their territories, or during the morning rounds of the territory. Pairs in neighbouring territories judge the proximity of rivals by the degradation of the song caused by interference by the environment. In bouts of territorial defence the male will face off with the other male and the female with her counterpart. Loudsong duets are also potentially related to the maintenance of pair bonds. The functions of softsongs are more complex, and possibly related to pair-bond maintenance. In addition to these two main calls a range of other sounds are made; these include scolding in mobbing of predators. The calls of antbirds are also used interspecifically. Some species of antbirds and even other birds will actively seek out ant-swarms using the calls of some species of ant-followers as clues. == Distribution and habitat ==

is distributed from Mexico to Argentina. It has the hooked bill typical of the antshrikes. The distribution of the antbirds is entirely Neotropical, with the vast majority of the species being found in the tropics. A few species reach southern Mexico and northern Argentina. Some species, such as the barred antshrike, have a continental distribution that spans most of the South and Middle American distribution of the family; others, such as the ash-throated antwren, have a tiny distribution. Antbirds are mostly birds of humid lowland rainforests. == Behaviour ==

Antbirds are diurnal: they feed, breed and defend territories during the day. Many of the family are, however, reluctant to enter areas of direct sunlight where it breaks through the forest canopy. Antbirds will engage in anting, a behaviour in which ants (or other arthropods) are rubbed on the feathers before being discarded or eaten. While many species of antbirds (and other families) may opportunistically feed at army ant swarms, 18 species of antbird are obligate ant-followers, obtaining most of their diet from swarms. With only three exceptions, these species never regularly forage away from ant swarms. A further four species regularly attend swarms but are as often seen away from them. Obligate ant-followers visit the nesting bivouacs of army ants in the morning to check for raiding activities; other species do not. These species tend to arrive at swarms first, and their calls are used by other species to locate swarming ants. Because army ants are unpredictable in their movements, it is impractical for obligate ant-followers to maintain a territory that always contains swarms to feed around. Antbirds have evolved a more complicated system than the strict territoriality of most other birds. They generally (details vary among species) maintain breeding territories but travel outside those territories in order to feed at swarms. Several pairs of the same species may attend a swarm, with the dominant pair at the swarm being the pair which holds the territory that the swarm is in. In addition to competition within species, competition among species exists, and larger species are dominant. In its range, the ocellated antbird is the largest of the obligate ant-following antbirds and is dominant over other members of the family, although it is subordinate to various species from other families (including certain woodcreepers, motmots and the rufous-vented ground cuckoo). At a swarm, the dominant species occupies positions above the central front of the swarm, which yields the largest amount of prey. Smaller, less dominant species locate themselves further away from the centre, or higher above the location of the dominant species, where prey is less plentiful. Breeding Antbirds are monogamous, in almost all cases forming pair bonds that last the life of the pair. Studies of the dusky antbird and the white-bellied antbird did not find "infidelity". In the white-plumed antbird, divorces between pairs are common, but, as far as known, this species is exceptional. In most species the pair defends a classic territory, although the nesting territories of ant followers are slightly different (see feeding above). Territories vary in size from as small as 0.5 ha for the Manu antbird, to 1500  m (5000 ft) in diameter for the ocellated antbird. Ocellated antbirds have an unusual social system where the breeding pair forms the nucleus of a group or clan that includes their male offspring and their mates. These clans, which can number up to eight birds, work together to defend territories against rivals. Pair bonds are formed with courtship feeding, where the male presents food items to the female. In spotted antbirds males may actually feed females sufficiently for the female to cease feeding herself, although she will resume feeding once copulation has occurred. Mutual grooming also plays a role in courtship in some species. has an unusual social system of shared breeding territories. One dominant pair may share a territory with up to six other birds. The nesting and breeding biology of antbirds have not been well studied. Even in relatively well-known species the breeding behaviour can be poorly known; for example the nest of the ocellated antbird was first described in 2004. Nests are constructed by both parents, although the male undertakes more of the work in some species. Antbird nests are cups of vegetation such as twigs, dead leaves and plant fibre, and they follow two basic patterns: either suspended or supported. Suspended cups, which may hang from forks in branches, or between two branches, are the more common style of nest. Supported nests rest upon branches, amongst vines, in hollows, and sometimes on mounds of vegetation on the ground. Each species nests at the level where it forages, so a midstory species would build its nest in the midstory. Closely related species nest in the same ways. For example, antvireos in the genus Dysithamnus are all suspension nesters. constructing a nest Almost all antbirds lay two eggs. A few species of antshrike lay three eggs, and a smaller number of antbirds lay one egg, but this is unusual. Small clutch sizes are typical of tropical birds compared to more temperate species of the same size, possibly due to nest predation, although this is disputed. Both parents participate in incubation, although only the female incubates at night. The length of time taken for chicks to hatch is 14–16 days in most species, although some, such as the dusky antbird, can take as long as 20 days. The altricial chicks are born naked and blind. Both parents brood the young until they are able to thermoregulate, although, as with incubation, only the female broods at night. In common with many songbirds, the parents take faecal sacs for disposal away from the nest. Both parents feed the chicks, often bringing large prey items. When the chicks reach fledging age, after 8–15 days, attending parents call their chicks. As each chick leaves the nest it is cared for exclusively from then on by the parent that was present then. After the first chick fledges and leaves with a parent the remaining parent may increase the supply of food to speed up the process of fledging. After fledging, chicks spend the first few days well hidden as the parents bring them food. Chicks of some species may not become independent of the parents for as long as four months in some antwrens, but two months is more typical for the rest of the family. == Ecology ==

Antbirds are common components of the avifauna of some parts of the Neotropics and are thought to be important in some ecological processes. They are preyed upon by birds of prey, and their tendency to join flocks is thought to provide protection against such predation. The greater round-eared bat preys on some antbird species, such as the white-bibbed antbird and the scaled antbird; the latter is the bat's preferred prey. Nests, including incubating adults, chicks and eggs, are vulnerable to predators, particularly snakes but also nocturnal mammals. Nesting success is low for many species, particularly in areas of fragmented habitat. It was once suggested that the relationship between the obligate and regular ant-followers and the army ants, particularly Eciton burchellii, was mutualistic, with the ants benefiting by having the birds chase prey back down towards them. However, experiments where ant followers were excluded have shown that the foraging success of the army ants was 30% lower when the birds were present, suggesting that the birds' relationship was in fact parasitic. This has resulted in a number of behaviours by the ants in order to reduce kleptoparasitism, including hiding of secured prey in the leaf litter and caching of food on trails. It has been suggested that the depressive effect of this parasitism slows the development of E. burchellii swarms and in turn benefits other ant species which are preyed upon by army ants. The ant-following antbirds are themselves followed by three species of butterfly in the family Ithomiinae which feed on their droppings. Bird droppings are usually an unpredictable resource in a rainforest, but the regular behaviour of ant followers makes the exploitation of this resource possible. == Status and conservation ==

of Brazil is threatened by the loss of deciduous forest and is listed as vulnerable by the IUCN. As of April 2008, 38 species are considered by the IUCN to be near threatened or worse and therefore at risk of extinction. Antbirds are neither targeted by the pet trade nor large enough to be hunted; the principal cause of the decline in antbird species is habitat loss. Additionally, new species are discovered at regular intervals; the Caatinga antwren was described in 2000, the acre antshrike in 2004, the sincorá antwren in 2007, and the description of a relative of the Paraná antwren discovered in 2005 in the outskirts of São Paulo is being prepared. ==See also==