passeriform

/pas"euhr euh fawrm', peuh ser"euh-/, adj.

of or pertaining to the order Passeriformes; passerine.

[ < NL Passeriformes, equiv. to L passer sparrow + -iformes -IFORMES]

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▪ bird

Introduction

also called  passerine  or  perching bird 

   any member of the largest order of birds (bird) and the dominant avian group on Earth today. The passeriform birds are true perching birds, with four toes, three directed forward and one backward. Considered the most highly evolved of all birds, passerines have undergone an explosive evolutionary radiation in relatively recent geological time and now occur in abundance on all continents except Antarctica and on most oceanic islands. Their rapid evolution and adaptation to virtually all terrestrial environments resulted in a large number of species, some 5,700, compared with only about 4,069 species of all other birds.

      The order Passeriformes is divided by most taxonomists into two suborders: Tyranni and Passeri. The first suborder, containing about 1,250 species, is considered more primitive and is often grouped informally as the “suboscines (suboscine).” Birds of suborder Passeri are often grouped as the “oscines (oscine),” or songbirds (songbird), for convenient comparison with the suboscines. Passeri is a very large group made up of about 4,500 species.

General features

Size range and structural diversity

      Passerines are small to medium-sized land birds (bird), ranging from about 7.5 to about 117 cm (3 to 46 inches) in overall length. Among the tiniest species are some of the New World flycatchers (flycatcher) (Tyrannidae), New Zealand wrens (wren) (Xenicidae), titmice (titmouse) (Paridae), flowerpeckers (flowerpecker) (Dicaeidae), tanagers (tanager) (Thraupidae), and waxbills (waxbill) (Estrildidae). The heaviest are the lyrebirds (lyrebird) (Menuridae) of Australia and the ravens (raven) (Corvus). The longest species, the ribbon-tailed bird of paradise (bird-of-paradise) (Astrapia mayeri), is actually not so large in body bulk but has extremely long tail feathers (feather). Most passerine species fall within the range of about 12.5 to 20 cm (5 to 8 inches) in length and from 15 to 30 grams (0.5 to 1 ounce) in weight. A house sparrow (Passer domesticus (house sparrow)), for example, is 12 to 15 cm (5 to 6 inches) long and weighs about 26 grams (0.9 ounce); a cardinal (Cardinalis cardinalis) is 20 to 23 cm (8 to 9 inches) long and weighs approximately 44 grams (1.6 ounces).

      Passerines have evolved a great diversity of feeding adaptations. The majority are insectivorous, at least at certain times of their lives. Members of the order have evolved many ways for finding insect food: swallows (swallow) (Hirundinidae) are aerial feeders; New World flycatchers “hawk” insects by flying out from a perch; vireos (vireo) (Vireonidae) glean insects from small twigs and foliage; woodcreepers (woodcreeper) (Dendrocolaptidae), nuthatches (nuthatch) (Sittidae), and creepers (creeper) (Certhiidae) search for insects in crevices in tree bark; and many other species pick and scratch on the ground and in leaf litter. More-specialized passerines eat aquatic insects (dippers (dipper): Cinclidae), fish (some New World flycatchers: Tyrannidae), fruit (cotingas: Cotingidae; and many others), leaves (plantcutters (plantcutter): Phytotoma), nectar (sunbirds (sunbird): Nectariniidae), small land vertebrates (shrikes (shrike): Laniidae), and seeds (finches (finch) and many others). For these different food habits, various structural specializations have developed, especially in the bill and feet (see below Form and function (passeriform)).

Importance to humans

Aesthetic and economic importance

      Since prehistoric times, people have enjoyed watching and listening to songbirds (songbird). The almost infinite variety of colours, patterns, behavioral traits, songs, and calls found in these birds appeals to people's aesthetic tastes. As objects of beauty and interest, passerines have been incorporated into human culture, folklore, poetry, music, sculpture, and painting. Songbirds have also been used as symbols; for example, the European goldfinch (Carduelis carduelis) represented the Passion of Christ in Renaissance art, and the raven (Corvus corax) sometimes signified a messenger of the Devil, an evil omen.

 Passerines are widely kept as cage birds (pet). The origins of this practice are lost in antiquity, but it is known that by the 5th century BC the Greeks kept a variety of songbirds, including finches, nightingales (nightingale) and other thrushes (thrush), magpies (magpie) (Pica), and starlings (starling) (Sturnidae). Canaries (canary) (Serinus canaria) were brought to Europe from their native Canary Islands in the 16th century and have since been developed into many varieties by domestication and breeding. Other passerines now widely kept as pets are the cardueline and estrildine finches and the starlings (particularly Asian mynahs (mynah), Gracula). The magnitude of the cage-bird “fancy” is indicated by importation statistics on wild and semidomestic birds: in one year alone during the late 20th century, over 420,000 passerines (excluding canaries) were legally imported into the United States as cage birds, a number far exceeding that of parrots (parrot), the only other bird group whose members are commonly kept as pets. Many countries, including the United States and Great Britain, prohibit the capture and sale of nearly all native songbirds.

      Songbirds are economically important in other ways also. Although seldom considered food in economically advanced areas, they are nonetheless important dietary items in many rural or heavily populated countries. China, Japan, and other Asian countries, for instance, have highly developed techniques for catching small birds; in cities such as Hong Kong and Tokyo, passerines are commonly sold in food markets. In Italy, France, and Belgium the capture of migratory songbirds for the pot or for cage birds is still extensive. Laws against such activities are difficult to enact or enforce in areas in which the habit has become part of the culture.

      Killing songbirds for their feathers (feather) is no longer as prevalent as it once was. Until the early 20th century, however, there were few protective laws, and the wearing of embalmed birds and bird parts (especially on women's hats) was common. In 1886 a young ornithologist reported that he had counted feathers from no fewer than 40 bird species, including 22 kinds of passerines, on hats seen on two afternoon walks in a fashionable part of New York City.

      Other cultures have used songbird feathers for personal adornment, but usually for men rather than women. This practice often came about not only for the beauty of the feathers themselves but also because the feathers were used as symbols of such bird qualities as speed and aggressiveness. Most noteworthy are the feathers of male birds of paradise (Paradiseidae), used as headdresses by tribesmen of New Guinea. An estimated 80,000 adult birds are still being killed annually for this purpose. Other ancient uses of passerine feathers have now largely been terminated, either because the birds are extinct (in the case of Hawaiian feather cloaks) or because more suitable modern substitutes have been found (Melanesian feather money).

      Some passerines, on the other hand, are serious economic pests (pest). In areas in which one-crop agriculture is extensive, certain bird species have undergone population explosions because of almost unlimited food availability; in turn, their crop depredations can be serious. One example of this is in Africa, where immense flocks of a small weaver, the red-billed quelea, or Sudan dioch (Quelea quelea), numbering as many as 20 million birds in one flock, do millions of dollars worth of damage to various small grain crops each year. Other serious pests are the Java sparrow (Padda oryzivora) in Asian rice fields and mixed flocks of New World blackbirds (blackbird) (Icteridae) and European starlings (Sturnus vulgaris) in grainfields in the United States. The same starling and the house sparrow, both introduced to the United States from Europe, have become urban pests by fouling buildings with excrement and blocking rain gutters and ventilators with their nests. Starlings occasionally have been implicated in accidents; in 1960 a flock at the airport in Boston was sucked into a jet's engines and the resultant crash killed 61 people.

Ecological importance

      The greatest importance of passerines is ecological. As the dominant form of birdlife in virtually all terrestrial environments, the perching birds are a major component of the world's ecosystems (ecosystem). They consume great quantities and varieties of food—grains, fruits, insects and other invertebrates, small amphibians and reptiles, and even small mammals—and in turn serve as food for other animals; they act as hosts for parasites and are occasionally parasitic themselves; they both propagate and distribute plants by pollinating (pollination) flowers and carrying viable seeds to new locations; and they have the mobility (through migration) to utilize habitats that are available only at certain times of the year. A few aspects of the ecological impact of passerines are known, but, until the science of ecology has advanced, the true magnitude of their importance cannot be evaluated with precision.

Natural history

Reproduction

Territoriality and courtship

      The breeding behaviour of passerines is diverse. Most species are solitary nesters, a single monogamous pair of birds (bird) maintaining a territory that is large enough to support all their activities during the breeding season: courtship, mating, nesting, and food gathering. Others have similar territories but forage outside the defended area for most of their food (e.g., the North American redwinged blackbird, Agelaius phoeniceus). Still others are colonial nesters, defending only the nest site and a small area immediately adjacent to it. Some species build individual nests close together in a colony (oropendolas (oropendola), Icteridae; some swallows (swallow); the house sparrow), and others construct massive communal nests in which the breeding pair defends only its own nest cavity ( palm-chat, Dulus; several weavers (weaver), Ploceidae). In a few species, polygynous (polygamous) males establish special display territories (leks (lek)) for courtship and mating in which no nesting takes place. In these courtship arenas the males, usually brilliantly coloured, attract females through song and posturing and sometimes by dancing, manipulation of objects, and other elaborate displays. The best-known arena-displaying males are the cocks-of-the-rock (cock-of-the-rock) (Rupicola), manakins (manakin) (Pipridae), birds of paradise (bird-of-paradise), and bowerbirds (bowerbird) (Ptilonorhynchidae). After mating in or near the lek, a female leaves to build a nest and raise the young without assistance from the male. Still other species build no nest at all but are brood parasites (some cowbirds (cowbird), Icteridae; whydahs (whydah), Estrildidae): the female lays her eggs in the nests of other (usually smaller) species, and the young are raised entirely by the foster parents.

Nesting

      Nest sites are varied: they include holes in the ground, trees, banks, and rock crevices; they may be on ledges, on the surface of the ground, within the larger nests of other species (including nonpasserines) or near wasp nests (presumably for the protection the wasps afford), and in a wide variety of vegetation—grasses, shrubs, and trees.

      Passerine nests are usually elaborately constructed and may contain many different kinds of materials: mud, grasses, hair and feathers (feather), strips of bark, plant fibres and downs, rootlets, twigs and sticks, leaves, string, spiderwebs, cast snake skins, lichens, and many other substances. Most species build open nests, usually cup-shaped. Others form domed or ball-shaped closed nests, with an entrance at the side (occasionally at the top or bottom). One of the most famous closed nests is that of the South American ovenbirds (ovenbird) of the genus Furnarius (Furnariidae), whose name derives from its thick-walled mud “oven” nest, often built on top of a fence post or some other exposed site. The North American ovenbird, Seiurus aurocapillus (a wood warbler, Parulidae), also builds a domed oven-shaped nest, but of plant materials on the forest floor. Some species, especially members of the Icteridae, make soft hanging nests that range to 0.6 metre (2 feet) or more in length. The thorn birds (Phacellodomus), as well as many other Furnariidae, build huge nests of twigs suspended from the ends of tree branches; these nests, which may be more than 2 metres (nearly 7 feet) long and contain many compartments, are used by only a single nesting pair, sometimes with nonbreeding helpers (probably the young of the previous season). These nests are often appropriated by troupials (Icterus icterus), which evict the owners, even destroying the eggs and young in the process. a few other species also take over nests for their own use, notably the piratic flycatcher (Legatus leucophaius, a tyrannid) and the bay-winged cowbird (Molothrus badius).

      Nests of many passerines are constructed with amazing skill. The tailorbirds (tailorbird) of Asia (Orthotomus) are noted for nests built in a pocket that the birds make by sewing together the edges of one or more leaves, using plant fibres or other materials. Some species, especially the weavers, are able to tie knots with strips of grass or palm leaves and thus weave an exceptionally tight and compact nest. Others build equally firm nests by felting the materials together. In contrast, a few passerines build flimsy nests (some Cotingidae), apparently as an adaptation toward lessened visibility to predators, for such nests are attended minimally by the parents, seemingly to draw as little attention to the site as possible. Other birds excavate their nests in soft earthen banks, use old woodpecker holes, or find natural crevices in trees or rocks. The type of nest built by the members of a single family may be varied (extremely so in the Furnariidae) or consistent: all woodcreepers (woodcreeper) nest in holes; all vireos (vireo) weave a cup between the arms of a forked branch.

incubation and parental care

 Passerines lay clutches of 1 to 14 eggs, clutch size being unrelated to the size of the bird. The largest species, the two lyrebirds (lyrebird) (Menura), lay a single egg; some of the smaller titmice (titmouse) (Parus) have been recorded with the biggest clutches. In most passerines the female incubates the eggs alone, but in some groups—such as the antbirds (antbird) (Formicariidae), certain grosbeaks (grosbeak) (Pheucticus), and others—the male shares equally in incubation. Males of most species help to feed the young. Some passerines have only one nest per breeding season, but others may have two or more, especially if one nest is destroyed before the young fledge. The incubation period generally varies from 11 to 21 days depending on the species but is well over a month in lyrebirds. The hatchlings are typically blind, sparsely covered with down, and helpless; some species hatch completely naked, and a very few are densely covered with down at hatching (some cotingas, antbirds of the genus Formicarius, and some Campephagidae). The young remain in the nest for 8 to 30 or 35 days (about 42 in the lyrebirds) but most commonly from 10 to 15 days. After they fledge, they require some days or weeks to become fully independent of their parents.

Sound production

      An outstanding aspect of passerine behaviour is the ability to sing (birdsong). Song is best developed in the oscines, which have a highly complex vocal organ or syrinx, but even the more primitive suboscines (suboscine) are capable of a variety of vocal sounds. The woodcreepers (Dendrocolaptidae), ovenbirds (Furnariidae), and antbirds (Formicariidae) sing relatively simple songs, consisting of a few notes or whistles, often repeated rapidly in a trill, roll, or rattle. Manakins (Pipridae) also utter simple trills or whistles: in addition, some species are capable of a loud nonvocal snapping sound, which is produced by specialized wing feathers. The cotingas sing a wider variety of songs, from quiet musical notes to the incredibly loud and far-carrying “gongs” of the bellbirds (bellbird) (Procnias). The New World flycatchers are well known for their range of distinctive call notes, and many species sing well and melodiously. In some groups (notably the Empidonax complex), the plumages (plumage) of closely related species are so similar that the birds can be distinguished in the field only by their calls and songs. Both the lyrebirds and scrub-birds (scrub-bird) (Atrichornithidae) are known for their loud and complicated songs. They are also accomplished mimics; lyrebirds mimic the songs of almost all birds within their hearing, as well as many mechanical sounds. Many species of oscines (oscine) have complicated and beautiful songs, notable examples being the nightingales (nightingale) (Luscinia) and some other thrushes (thrush), larks (lark) (Alaudidae), mimic thrushes (Mimidae), and wrens (wren) (Troglodytidae). The possession of the complex oscine syrinx does not guarantee a complex song, however, and many “songbirds (songbird),” such as waxwings (waxwing) (Bombycilla) and swallows, utter simpler sounds than do many suboscines.

      Only the male of most passerine species sings a true song, although the female can produce a variety of call notes and other sounds. In some species in which the female sings, she seldom does so during the breeding season unless it is a duet with her mate. Such duetting or antiphonal singing of paired birds is so well developed in certain species that it is difficult to determine that the song is coming from two individuals. In the African black-and-red shrike (Laniarius barbarus erythrogaster), the reaction time between the male's start of song and the female's response has been timed at 0.135 second.

Interactions with ants

Anting

      A characteristic but poorly understood behaviour pattern of passerines is the practice of anting. This peculiar ritual has two forms: active anting, in which a bird picks up worker ants (ant) in its bill and wipes them on its feathers (feather) in a stereotyped manner, and passive anting, in which the bird squats or lies down in a group of ants and assumes an exposing stance so that the ants will crawl up into its feathers. Birds may also apply ants to their plumage while passively anting, but species that use the active stance (the majority of recorded passerines) apparently never use the passive stance. Birds show definite discrimination in the type of ants used, avoiding stinging species and selecting those that exude or spray formic acid or other defense fluids (ants of the subfamilies Formicinae and Dolichoderinae of the family Formicidae).

      A great deal of controversy has existed over the function of anting. Some authorities have theorized that it is a form of self-stimulation, but most ornithologists conclude that anting is a type of feather maintenance. Formic acid and other ant fluids are known to be insecticidal; dressing the feathers with ants would thus kill or deter avian parasites, such as lice and mites. Additional components of ant fluids include essential oils, which could be used by birds to supplement the oils from their own uropygial (preen) gland. After a bout of anting, birds often continue feather-maintenance activities by bathing, oiling (from the uropygial gland), and preening. Recent studies have shown anting to be most prevalent during molt, when the bird's skin is irritated by the growth of new feathers. Anting clearly is innate behaviour, and its remarkable uniformity in at least 30 passerine families, both oscine and suboscine, implies that it has real importance to the bird. Some individuals have been seen to ant with such things as cigarette butts, orange peels, mothballs, and smoke, apparently reacting to the pungent fumes of these objects as to the strong odours of ants. A few nonpasserines have also been observed going through motions that are similar to anting, but, as yet, true anting is known only in the Passeriformes. Another specialized form of behaviour associated with ants is the practice known as ant-following.

Ant-following

 In the New World tropics, nomadic army ants move in huge troops, swarming over the forest floor in columns as wide as 10 metres (about 30 feet) or more. Because the ants devour all the small animal life in their path, a moving column of them is edged by fleeing insects, spiders, millipedes, isopods, small frogs, and lizards. The ant columns are accompanied by troops of birds that seize the fugitives. Ant-following birds apparently do not eat the ants but only the insects and other small animals trying to escape. A number of passerine species, notably several antbirds (antbird), are believed to be entirely dependent on army ants for finding food. Many other birds also follow ants when they come upon them; these include woodcreepers (woodcreeper), manakins (manakin), New World flycatchers (tyrant flycatcher), tanagers (tanager), wrens (wren), and occasional ovenbirds (ovenbird). Even some nonpasserines may join a troop of ant followers—motmots (motmot) (Momotidae), tinamous (tinamou) (family Tinamidae), and hawks (hawk)—although the hawks may be more attracted by the ant-following birds than by the insects. The same ant-dependent species have also been known to follow large animals, including man, that stir up insects with their feet.

      A few passerines, although not ant followers, will escort large quadrupeds, such as cattle, buffalo, and deer, to catch the insects that fly up around them and to feed on the ticks and flies parasitizing the animals themselves; especially noted for this behaviour are the cattle tyrant (Machetornis rixosa, Tyrannidae) tickbirds or oxpeckers (oxpecker) (Buphagus, Sturnidae), and several cowbirds. In Australia, yellow robins (Eopsaltria) follow the much larger lyrebirds as they scratch and feed along the ground.

Form and function

External features

Feet and legs

 The single feature that distinguishes passerines from all similar birds (bird) is their “perching” foot. In this foot type, all four toes are well developed and free from one another; in some families (wrens (wren) and most suboscines (suboscine)), the front toes may be partially fused at the base, but the distal portions (extremities) are functionally free. The hind toe (hallux) is joined on the same level with the front toes and opposes them, so that the foot can grip a perch. The only exception to this passerine foot type is found in the well-named Paradoxornis paradoxus, or three-toed parrotbill (Panuridae), in which the outer toe is reduced to a short clawless stump, fused to the middle toe; other species of Paradoxornis have normal feet.

      Although all passerines can perch, not all do so habitually. A number of species (some tapaculos (tapaculo), Rhinocryptidae; larks (lark); pipits (pipit), Motacillidae) are largely terrestrial and have feet modified for walking and running; the terrestrial foot is differently proportioned from the typical perching one, often with longer toes and longer, straighter claws (particularly on the hallux), probably as an aid in maintaining balance when running. The dippers (dipper), or water ouzels (ouzel) (Cinclus), are semiaquatic, but, although they successfully swim on the water surface and walk underwater searching for food on stream bottoms, they have retained the typical passerine foot. The single slight difference in the Cinclus foot is that the claw of the middle toe sometimes has a thin horny flap (of unknown function) on its inner border. Some other passerines, notably swallows (swallow), live a largely aerial life and have small and weak feet. The typical arboreal songbird has a well-developed foot, with the middle front toe longer than the others. Birds such as woodcreepers (woodcreeper) and nuthatches (nuthatch) that often cling to vertical surfaces have strong, curved, sharp claws. Those that spend much of their time walking and scratching on the ground (although not limited to terrestrial activity) tend to have heavy, straighter, and rather blunt claws. Most passerines, however, have moderately curved sharp claws that are suited to grip a variety of rounded or rough surfaces.

      The lower leg of passerines, the tarsometatarsus (usually called simply the tarsus), is normally covered by a horny sheath (podotheca). Exceptions include some swallows, which have feathered (feather) tarsi. Although the various different patterns of scale size and distribution of the normal unfeathered podotheca have been used by some taxonomists to differentiate families or groups of families, study has revealed so much variability in the tarsal patterns of certain families that it is no longer considered a reliable family character; it may still be useful as a generic or specific character. In most oscines (oscine) the posterior (plantar) surface of the tarsus is bilaminate—that is, covered by two long plates, or laminae.

Bill (beak)

 The bills of passerines are extraordinarily diverse in size, shape, and proportions. This diversity was long thought to be indicative of the birds' relationships and so was used as a prime taxonomic character. It is now believed, however, that bills are evolutionarily plastic, reacting with relative ease to selective pressures, particularly to changes in feeding habits. Thus, on a broad scale, a passerine's bill shape reveals less about its family affinities than it does about its food preferences, and, although bill shape may be an aid to determining a bird's relationships, it must be considered in the light of other features and of the degree of variation found in the family. Two frequently cited examples of the adaptiveness of bills are the Darwin's finches (finch) of the Galapagos Islands and the Hawaiian honeycreepers (honeycreeper), Drepanididae (see evolution: Adaptive radiation (evolution)). Each is a closely interrelated group of birds with different kinds of bills in the several species and genera. Bills of the drepanidids range from heavy, seed-cracking, grosbeaklike (grosbeak) bills through thin, pointed, insectivorous types to the long, decurved (curved downward) bills of nectar feeders. These Hawaiian birds are now thought to be members of a single family of nine genera. On the basis largely of bill shape, they were once classified into four different families and 18 genera.

      Most birds, including passerines, show little sexual dimorphism (difference between sexes) in bills except for minor differences in size (reflecting general body size differences) and sometimes in colour. The most outstanding exception is the extinct huia (Heteralocha acutirostris, Callaeidae), originally classified as two different species. The male of this New Zealand bird had a strong chiselling bill, whereas the female had a long, decurved, pliable bill. Reportedly, the two sexes fed cooperatively, the male digging in decaying wood and the female probing in crevices to extract grubs. The species unfortunately was prized by the Maoris, who used the white-tipped tail feathers in ceremonial headdresses, as well as by Europeans, and, after most of its habitat had been destroyed, the huia was hunted to extinction about the end of the 19th century.

      Passerine bills may be broadly classified into eight morphological and functional types:

● Insectivorous: a generalized type found in many passerines, ranging from relatively straight and pointed (as in the wood warblers (warbler), Parulidae), through bills with a slight or pronounced hook (some New World flycatchers (flycatcher)), to those that are short, with a wide gape and usually surrounded by rictal bristles (stiff hairlike feathers)—as in aerial feeders, such as swallows. Most insectivorous bills are relatively light in build, but this depends on the type of insect usually taken by the species and also on how generalized a feeder it is.

● Omnivorous: unspecialized in shape and function but usually strongly built, as in crows and jays (jay) (Corvidae).

● Toothed: strongly hooked at the tip and with a “tooth” (notch) on either tomium (cutting edge) of the upper mandible; adapted to tearing up large, relatively soft prey. This is the typical bill of shrikes (shrike) (Laniidae) but is also found in some unrelated birds, such as the Australian bell-magpies (bell-magpie) (Cracticidae) and some tanagers (tanager).

● Tearing: a relatively light bill with a strong hook at the tip, for tearing open objects, such as flowers, to obtain the insects and nectar within. Found in flower piercers (Diglossa, Thraupidae).

● Probing: relatively narrow and often downcurved; slender in species that probe flowers for tiny insects and nectar (sunbirds (sunbird); some Hawaiian honeycreepers) but more heavily constructed in those that probe in wood or under tree bark (creepers (creeper), Certhia; some woodcreepers).

● Frugivorous: variable but usually rather wide; ranges from lightly built with a wide gape for swallowing whole fruits (found in some cotingas, and in the swallow-tanager, Tersina) to more heavily built for tearing apart tougher fruits (some tanagers).

● Serrated: conical, with a finely serrated edge, adapted for feeding on leaves, buds, shoots, and fruit. Found only in the plantcutters (plantcutter) (Phytotoma, Cotingidae).

● Conical: adapted for seed eating. Ranges from exceedingly stout and blunt (such as the hawfinch, Coccothraustes, which can crack remarkably hard objects, such as cherry pits) to relatively small and pointed (siskins (siskin), Carduelis). Some forms specialized for particular kinds of seed extraction (such as crossbills (crossbill), Loxia, which feed on pine seeds).

      This classification indicates morphological and functional types of bills, but it does not imply that a species with a particular type of bill will feed only on the food for which it is best adapted. Although some birds are extremely specialized in their feeding habits, most are opportunistic feeders, seizing upon whatever food is readily available and can be “handled” with the bill. Hence, many basically granivorous or frugivorous birds catch insects, especially when feeding nestlings, and many insectivorous species exploit seasonally available plant food. Yellow-rumped warblers (Dendroica coronata) and tree swallows (Iridoprocne bicolor), for example, feed on bayberries in fall and winter, and eastern kingbirds (kingbird) (Tyrannus tyrannus) and other New World flycatchers eat a variety of fruits and berries in season.

      The mandibles of passerines, like those of all other birds, are composed of bone covered with a horny sheath, the ramphotheca. The ramphotheca is worn down by normal use and, in most birds, is capable of growing to replace the lost material. In individuals with damaged bills or those (such as cage birds) that do not have the opportunity to wear down the constantly growing ramphotheca, the bills overgrow at the tip.

plumage and pterylosis

  The colours, patterns, and textures of passerine feathers are considered important taxonomic characters, especially in determining genera, species, and subspecies. plumage is also occasionally used in a very broad way to indicate evolutionary levels. Spots, streaks, and dull colours are generally considered more primitive than bold or complicated patterns and bright colours, but there are many exceptions to this rule.

      Passerines often are sexually dimorphic in their plumage, with adult males wearing brighter colours and more striking patterns than do females. In some families, notably tanagers (Thraupidae), wood warblers (Parulidae), and New World orioles (oriole) (Icteridae), the temperate zone species show more sexual dimorphism than do tropical members of the same families. In addition, many species (especially those in temperate climates) are seasonally dimorphic, with a bright plumage during the breeding season and a dull one in winter. Juvenile plumages of both sexes tend to be cryptically coloured (that is, adapted for concealment), as is that of the adult female.

      Virtually any colour may be found in one passerine or another, and the order offers a wide array of specialized feather types, such as the waxlike tips on the flight feathers of waxwings (waxwing) (Bombycillidae); the tufts of stiff feathers in some honeyeaters (honeyeater) (Meliphagidae); iridescent “spangles” in some manakins (manakin), sunbirds, and tanagers; and the almost unbelievable array of “wires,” iridescent gorgets, velvety ruffs, racquet tails, and filamentous plumes of the birds of paradise.

      Another taxonomically important character is the number and distribution of feathers (pterylosis) on the bodies of passerines. From external appearance all birds seem to be more or less evenly covered by feathers; in actual fact, however, most birds have their feathers growing from relatively narrow tracts (pterylae) in the skin. From the pterylae the feathers fan out and cover the remainder of the bird's body. In passerines, the feathers are arranged in eight distinguishable tracts, with apteria (relatively bare skin) between them. Variations in tract width and length and especially differences in feather number and distribution are often useful in determining relationships. Of particular interest are the occurrence of apteria within tracts and the configuration of the ventral tract. Also used in classification are the numbers of flight feathers. The remiges (flight feathers on the wings (wing)) of most passerines consist of 10 primaries on the “hand” (manus) and 9 secondaries on the forearm (ulna). In all perching birds the 10th (outermost) primary is reduced to some degree, and in many families only 9 may be found. The number of secondaries is more variable, with some species having as many as 14 (the satin bowerbird, Ptilonorhynchus violaceus). Tail feathers (rectrices) also vary; most passerines have 12, but the number ranges from 6 to 16.

      Of importance in some species is the relative length of the primaries. This “wing formula” is often useful to differentiate between species of such difficult groups as the New World flycatchers and the Old World warblers (Sylviidae).

Internal features

syrinx

      In a group of birds as vocal as the passerines, it is natural that the structure of the vocal apparatus should have evolutionary significance. Differing from the mammalian larynx in both location and structure, the syrinx consists of a resonating chamber at the lower end of the windpipe (trachea), with associated membranes, cartilages, and muscles. These modifications involve elements of the bronchi (the two tubes connecting the trachea with the lungs) as well as those of the trachea. Since the mid-19th century the basic subdivisions of the order Passeriformes have been based primarily on the structure of the syrinx. Syrinx morphology has also provided characters useful for modern taxonomic revisions of such groups as the tyrant flycatchers (tyrant flycatcher) (Tyrannidae).

      Syringeal muscles are classified into two groups: extrinsic muscles, which connect the syrinx with other parts of the anatomy, and intrinsic muscles, which extend from one part of the syrinx to another. The number, shape, and attachments of the intrinsic muscles are likely to remain important in passerine classification. Those birds in which the muscles are inserted on the middle of the bronchial semi-rings (C-shaped cartilages that strengthen the bronchi) are sometimes called mesomyodian (most members of the suborder Tyranni), and those with the insertion on the ends of the semi-rings are acromyodian (Menuridae, Passeri). The broadbills (Eurylaimidae) and a few others have no intrinsic muscles. Further distinction is made in the number of pairs of intrinsic muscles, most importantly in the Passeri, which have four.

      The passerine syrinx exists in four basic types:

● Unspecialized: relatively little modification of the tracheobronchial region; few, if any, cartilaginous specializations, and no intrinsic muscles; found in broadbills (broadbill) (Eurylaimidae), pittas (pitta) (Pittidae), New Zealand wrens, asities (asity) (Philepittidae), plantcutters, most cotingas, and a few manakins and tyrant flycatchers.

● Tracheophone: most of the specializations limited to the tracheal region; intrinsic muscles number zero to two pairs; pessulus (a bony bar lying at the junction of the bronchi) absent; found in all members of the Furnarioidea (South American ovenbirds (ovenbird), woodcreepers, antbirds (antbird), and tapaculos).

● Intermediate tracheobronchial: various modifications of cartilages and membranes; one or two pairs of intrinsic muscles; pessulus present or absent; found in the sharpbill (Oxyruncus) and most manakins and tyrant flycatchers.

● Oscine (acromyodean): complex musculature involving four pairs of intrinsic muscles (but three pairs in lyrebirds and scrub-birds (scrub-bird)); some cartilaginous specializations; pessulus present (except in larks).

Skeleton

      Of the many variations in passerine skeletal structure, only a few that are important in classification are mentioned here.

      In the skull the bony palate, composed of a number of small bones, is termed aegithognathous; also found in swifts (swift) (Apodidae), this palatal type is characterized by the shape and type of fusion of the small bones of the palate. Within this basic type the many minor variations in shape, size, and position of the component bones are useful in delimiting closely related groups of birds, especially suboscines.

      Elsewhere on the head, variations in the hyoid apparatus, a complex of small bones that supports the tongue, have been used in passerine classification.

      In the sternum (breastbone) the shape of the anteriormost spine (spina sternalis) and the number of notches in the posterior border are of great interest. The spina sternalis, which is short and forked in most passerines, is long and simple in the Eurylaimidae (one exception), the Philepittidae, and a few of the Cotingidae. All oscines and most suboscines have a single pair of posterior sternal notches; only the tapaculos and certain of the terrestrial antbirds (Conopophaga, Pittasoma, Hylopezus, Myrmothera) have two pairs. The sternum of lyrebirds differs from those of all others in the order in being very thick, long, and narrow; it may have no posterior notches at all, or it may have a single shallow pair.

Musculature

      A number of different muscle systems have been important in passerine classification. Important examples, in addition to those of the syrinx, are the muscle complexes controlling the tongue, the jaws, the wings and pectoral girdle, and the legs and pelvic girdle. One character that has been used since the 19th century is the condition of the deep plantar tendons. These narrow straps extend from the bellies of the two deep flexor muscles on the leg and down the back of the tarsometatarsus and attach to the toes. They act to close the toes (hence to grasp a perch). In the Eurylaimidae these tendons are connected by a short band (vinculum), but in all other passerines they are entirely separate. This difference has been used by some to divide the passerines into two major groups: the Desmodactyli (vinculum present) and the Eleutherodactyli (vinculum absent).

Evolution and paleontology

      Passeriforms are now the dominant group of modern birds (bird). The 5,700 extant species make up 60 percent of all birds of the world. Ornithologists dispute many details of their evolutionary history, but almost all agree that they are monophyletic; that is, they are derived from a single ancestral lineage. Passeriforms are distinguished unambiguously by a series of unique derived characters.

      Preceded by coraciiform and piciform birds as the dominant land birds of the early Paleogene Period, passerines first appeared in the fossil record of the late Oligocene Epoch (some 34–23 million years ago) of France. Passerines of any kind are absent from the abundant fossils of landbirds of the preceding Eocene Epoch, and some early fossils of passerines have been reclassified to other taxa. Prior to the Oligocene, any forms must have been rare indeed. By the early Miocene Epoch (some 11.6 to 5.3 million years ago), however, passerines became very abundant and diverse as they outnumbered all other birds combined in the lower Miocene deposits of the Wintershof-West in the mountains of southern Germany. Basic family lineages with modern genera that included crows (crow) (Corvidae), thrushes (thrush) (Turdidae), wagtails (wagtail) (Motacillidae), Old World warblers (warbler) (Sylviidae), shrikes (shrike) (Laniidae), and wood warblers (wood warbler) (Parulidae) were established by the this time.

      During the Pliocene Epoch (5.3–2.6 million years ago) the warm, dry conditions of the Miocene continued, and all the living passerine families diversified through speciation. Most ornithologists believe that most modern species of birds arose during the early Pleistocene Epoch (about 1.8 million to 11,800 years ago), a period of cooling temperatures, shifts in habitats, and advancing glaciers. Most of the passerines in the fossil record are from the Pleistocene or Holocene and represent either living species or close relatives. Evolution since the retreat of the last ice sheet (about 11,800 years ago) has been mainly at the subspecies level.

      The evolutionary success of passerine birds begs for explanation. Most ornithologists have rejected the possibility that one key feature is responsible. Instead, as summarized by American ornithologist John Fitzpatrick, the large brain size, behavioral plasticity, and rapid population turnover of small-sized species may have facilitated more-rapid morphological evolution and speciation than in nonpasserines. The combination of a flexible body plan and superior neural capacities enabled passerines to explore and adapt to novel environments. Added to those traits, American ornithologist Nicola Collias suggested that the complex nest-building behaviours of passerine birds released them from the obligatory cavity-nesting behaviours of their predecessors and the move into new habitats and ecological zones.

Classification

Distinguishing taxonomic features

      Passerine birds are distinguished by a suite of unique derived characters that include (1) aegithognathous (that is, characterized by the fusion of the vomer bone in the forward part of the skull) structure of the palate, (2) syringeal anatomy, (3) multiple characters of the anisodactyl foot, such as a large, incumbent, rear-directed hind toe (hallux) capable of independent action, (4) insertion of the forearm muscle tensor propatagialis brevis, and (5) spermatazoa with a coiled head.

      In the mid-20th century, taxonomists began reexamining the generally accepted family groupings of passerine birds that had been in place since the 19th century. In these investigations, some of the features on which earlier classifications were based (such as bill shape and tarsal scutellation) were deemed to be either the result of convergent evolution or too variable to be useful in certain groups. Consequently, passerine taxonomists have been left with a rather sparse body of morphological information upon which to base a classification. Beginning in the mid-20th century, ornithologists made a concerted effort both to augment some of the century-old work on passerine anatomy and to explore new avenues of morphology, behaviour, reproductive patterns, biochemistry, and zoogeography. These explorations helped to define and relate the many families of perching birds. Nevertheless, some of this work is still in progress and has not yet been incorporated into classification systems.

      Among the traditionally studied taxonomic features are external characters such as rictal bristles and other specialized feathers (feather), colours and patterns of the fleshy parts of the mouth, morphology of the bill and nostrils, colour patterns of adults and young; internal anatomical characters such as the number of cervical (neck) vertebrae, the condition of the deep plantar tendons, anterior and posterior spines and processes of the sternum, syringeal muscles, palatal and other bones of the skull, feather tracts, jaw and tongue musculature, hyoid (tongue) apparatus, aortic arch system, pneumatic fossa (cavity) of the humerus, and types of spermatozoa; biochemical analysis of substances such as egg white, eye lens, plasma proteins, and hemoglobins; and an array of behavioral traits such as reproductive behaviour, nest building, and methods of scratching. Nonetheless, analyses of DNA sequences, especially the cytochrome b gene of mitochondrial DNA, have produced the most significant advances in the understanding of the relationships between passerine birds. Also notable are the pioneering studies of Charles Sibley and Jon Edward Ahlquist, American ornithologists who exposed the unique radiation of songbirds in Australia and contingas in South America. Their studies also revealed a major division between the passerine birds of family Corvidae and other families, a division which has been corroborated by other ornithologists. As the taxonomic understanding of passeriforms increases, improved phylogenies of many groups of species are published regularly in the ornithological literature.

Annotated classification

      The classification and sequence of families given here conservatively integrate modern biochemical evidence with classic morphological evidence.

Order Passeriformes (perching birds, or passerines)

 Land birds with a characteristic “perching” foot; 4 toes (never webbed) joined at the same level, with the 1st toe (hallux) directed backward and never reversible. Oil gland unfeathered. Wing eutaxic (no gap between the 4th and 5th secondaries), usually with 9 or 10 primaries. The young are altricial—that is, hatched almost or completely naked of feathers (a few exceptions), helpless, and requiring a considerable period of parental care. About 5,700 species.

      Suborder Tyranni

 Syrinx usually more complex; muscles variable; pessulus present or absent. Sternum with short spina sternalis, forked (exceptions noted below); posterior border with 1 or 2 pairs of notches. Hallux strong. Clavicles well developed.

      Family Xenicidae (New Zealand wrens (Xenicidae))

 Small birds, 7.5 to 10 cm (3 to 4 inches), look and act much like true wrens (Troglodytidae). Legs long and slender; 3rd and 4th toes fused basally, and all toes (especially the hallux) with long claws. Considered evolutionary relicts, perhaps the oldest lineage of passeriformes according to DNA-DNA hybridization data. 2 species strongly arboreal, the other 2 terrestrial. 1 of the latter, the now-extinct Stephen Island rock wren (Xenicus lyalli), was markedly terrestrial and may have been flightless; if so, it would have been the only known passerine to have completely lost the ability to fly. The 3 extant species are very weak flyers, with short wings and very short tails. Forest and scrub; New Zealand.

      Family Pittidae (pittas (pitta))

 A relatively homogeneous family; stout-bodied, long-legged, largely terrestrial, 15 to 28 cm (6 to 11 inches); with a wide range of colours in the loose-webbed plumage: reds, greens, blues, as well as browns, black, and white. Wings short and rounded but strong; tail very short. Syrinx simple, lacking intrinsic muscles and cartilaginous modifications; pessulus absent in most species. Egg white protein pattern differs from that of all other suboscines. About 30 species, Old World tropics in forests and scrublands. Africa through Malaysia, Australia, Solomon Islands.

      Family Eurylaimidae (broadbills (broadbill))

 Generally brightly coloured, chunky birds, with large heads, short necks; 12.5 to 28 cm (5 to 11 inches) in length. Bill broad and flattened, covered with a crest in some; gape wide. Front toes partially joined; 10 or 11 primaries. Syrinx simple, tracheobronchial, lacking intrinsic muscles and cartilaginous specialization; pessulus present. Usually 15 cervical vertebrae (all other passerines have 14). Deep plantar tendons of different type from all other passerines. 8 genera, about 14 species, primarily in forests or cloud forests. Indo-Malaysia, Africa, Philippines.

      Family Philepittidae (asities (asity) and false sunbirds (false sunbird))

 The 2 species of asities (Philepitta) are black or yellowish green, rather pittalike, with stout bodies and long legs. The 2 false sunbirds (Neodrepanis) are very different externally, looking much like true sunbirds (Nectariniidae), with which they were long classified; small, blue, yellow, and greenish with short tails and long, slender, curved bills. All philepittids have bare skin or wattles or both around the eyes. The simple structure of their syrinx and the unforked spina sternalis suggest they are allied to the broadbills (Eurylaimidae). 2 genera, 4 species. Forests of Madagascar.

      Family Tyrannidae (tyrant flycatchers (tyrant flycatcher))

 Large family of generally (but not exclusively) arboreal birds with plumage in grays, browns, olive greens, some with black, white, and yellow, occasionally brighter colours. Many with erectile crowns or crests, often more colourful than the rest of the plumage; length 7.5 to 40.5 cm (3 to 16 inches). Bill extremely variable but commonly broad, somewhat flattened, and hooked at tip; nostril rounded without an operculum or narrow with membranous operculum; most species with well-developed rictal bristles (stiff hairlike feathers around the mouth). Feet weak except in the few terrestrial species; front toes variably but never strongly fused. Syrinx characterized by 1 pair of intrinsic muscles (none or 2 in some), generally variable in other features. Approximately 100 genera, about 400 species, with a wide range of habitats. Northern Canada and Alaska through North and South America to Tierra del Fuego, also Falkland and Galapagos islands. (The family's common name, New World flycatchers, serves to distinguish it from the large oscine family of Old World flycatchers, Muscicapidae.)

      Family Dendrocolaptidae (woodcreepers (woodcreeper))

 Characteristically slender arboreal birds, olive brown to rufous, usually streaked or barred; 14.5 to 37 cm (5.5 to 15 inches). Strong probing bills, laterally compressed, short and straight to long and downcurved; nares holorhinal (see below Furnariidae). Short legs, powerful feet; front toes partially joined at base. Tail stiff with spiny tips. Many hitch up trees like woodpeckers. Posterior border of sternum with one pair of notches. Syrinx with cartilaginous specializations and two pairs of intrinsic muscles. Pterylosis with distinctive ventral tract type. 13 genera, approximately 54 species, forest or brushland. Mexico through South America.

      Family Furnariidae (ovenbirds (ovenbird))

 A large and extraordinarily diverse group, an excellent example of adaptive radiation. In South America, especially in the southern Brazil-Argentina-Chile region, this family has evolved to fill a broad range of ecological niches; various furnariids look and behave like wrens, thrushes, dippers, creepers, larks, wood warblers, titmice, nuthatches, nonpasserine woodpeckers (Picidae), and even sandpipers (Scolopacidae). If such diversity can be summarized, ovenbirds are generally small, dull brown birds, darker above and paler below; 12 to 28 cm (about 5 to 11 inches). A distinctive characteristic is the shape of the external nares (nostrils), which are schizorhinal (slitlike) at their posterior border rather than holorhinal (rounded), as in other passerines. Front toes partially joined at base. Sternum with 1 pair of notches (some exceptions). Syrinx with 2 pairs of intrinsic muscles. Pterylosis distinctive within suboscines but of same type as many oscines. Approximately 60 genera, about 246 species, wide variety of habitats; Mexico through South America.

      Family Formicariidae (antbirds (antbird))

 A large and diverse family, with loose-webbed plumage, generally in browns, grays, black, and white. Authorities inclined to split this group into 2, the Thamnophilidae (typical antbirds) and Formicariidae (ground antbirds); relationships between gnateaters (Conopophaga) and ground antbirds uncertain. Overall size range 9.5 to 37 cm (less than 4 to nearly 15 inches), including several large terrestrial species with long legs but very short tails (hence a deceptively short “body length”). Bill strong, hooked to variable degrees. Front toes slightly joined at base. Sternum with 1 or 2 pairs of notches, the 2 pairs found in most but not all the long-legged terrestrial species. Syrinx of 2 types: the 1st with 1 pair of ventrally originating intrinsic muscles (largely the arboreal species), the 2nd with no intrinsic muscles. Pterylosis variable; dorsal tract with reduced or absent posterior element (primarily the arboreal species); several different dorsal types among the terrestrial species. Wings generally short and rounded, flight weak. 52 genera, 244 species, largely in dense forests and brushland; neotropical, Mexico to northern Argentina.

      Family Conopophagidae (gnateaters (gnateater))

 1 genus (Conopophaga), 8 species.

      Family Rhinocryptidae (tapaculos (tapaculo))

 A small, little-known, secretive group of usually terrestrial species. Plumage dull, generally in grays, browns, and black; 9 to 25.5 cm (less than 4 to more than 10 inches). Bill distinguished by a movable flap (operculum) over the nostrils. Legs, feet, and claws strong. Wings short and rounded, flight feeble. Sternum with 2 pairs of notches. Syrinx with 1 pair dorsally originating intrinsic muscles (some exceptions). Pterylosis distinctive in ventral tract (exceptions). 12 genera, 55 species, grasslands, scrublands, and dense forest undergrowth. Central and South America, especially Chile and Patagonia.

      Family Cotingidae (cotingas, plantcutters (plantcutter), and sharpbills (sharpbill))

 An extremely diverse, probably composite family; ranges from plain-coloured flycatcher-like to bright, extravagantly adorned birds, such as the cock-of-the-rocks: 9 to 45.5 cm (about 4 to 18 inches). Some with extraordinary plumes and lappets (the crow-sized umbrella bird, Cephalopterus); others renowned for their loud voices, fleshy wattles, and white plumage (white being unusual in land birds). The family as traditionally constituted can no longer be characterized anatomically: the syrinx is of 2 types; the tarsal envelope is of several different kinds; the joining of the front toes is variable; the dominant artery of the thigh is either the sciatic or femoral; the spina sternalis is forked except in some bell birds (Procnias). Several species with powder downs, a specialized type of feather otherwise known in passerines only in the oscine woodswallows (Artamidae). 35 genera, approximately 100 species, forests. U.S.–Mexico border to Bolivia, Peru, and Argentina.

      Family Pipridae (manakins (manakin))

 Most piprids are fairly uniform in external appearance; generally small (8.5 to 16 cm [3.5 to 6.5 inches]), rather stubby, with short wings and tail (a few long-tailed species). Most males black with patches of brilliant colour (red, yellow, blue, etc.); females are generally drab olive green. Several species, such as the broad-billed manakin (Sapayoa aenigma), are externally and anatomically unlike the majority of manakins and may not be properly included in this family. Bill short, rather broad at base, notched, with a slight hook at tip. Some species with specialized feathers, particularly in the wings (also true of some cotingas and tyrannids). 3rd toe partially fused at base to 2nd or 4th. Syrinx highly variable, as family is presently constituted; about 16 genera, approximately 57 species, tropical and subtropical forest. Southern Mexico to Paraguay and northern Argentina.

      Suborder Passeri (songbirds (songbird), or oscines (oscine))

 Syrinx with 4 pairs of intrinsic muscles. Sternum with spina sternalis short and forked, and posterior border with 1 pair of notches. Hallux variable in strength. Clavicles well developed. All with same complex of syringeal muscles, with only minor variations.

      Family Climacteridae (Australian treecreepers (treecreeper))

 Small, creeperlike climbing birds, 12.5 to 17.5 cm (5 to 7 inches); of uncertain ancestry and affinities. Legs short; toes long, claws long, curved, strong, especially that of hallux; tail rounded, soft; bill long, somewhat downcurved. Grayish brown to black above, streaked below, often a lighter eye stripe and wing bar; sexes similar but with slight recognizable differences. Rather solitary, sedentary birds; feed by spiralling up tree trunks, some forage for insects on ground. Voice high-pitched whistles. 2 genera, 7 species. Forests of Australia and New Guinea.

      Family Menuridae (lyrebirds (lyrebird))

 Among the largest members of the order in length, 75 to more than 100 cm (30 to 40 inches), with some weighing over 1,100 grams (about 2.5 pounds), lyrebirds are aberrant passerines in virtually all features. In size, brownish plumage, and terrestrial habits, they look much like pheasants. The tail of 16 feathers is extraordinary: in males of Menura superba the outermost pair of feathers is curved like a lyre, the next 6 pairs are light and filamentous, and the innermost pair is modified into narrow “wires”; the tail of M. alberti is shorter and simpler; females of both species have rather long but not extremely modified tails. The alula (free digit on the leading edge of the wing) on the short rounded wings has 6 feathers rather than the usual 3 or 4. Rictal bristles present. Legs and feet strong and heavy. Sternum long, narrow, and thick, unique within passerines; posterior border may be entire or with 1 pair of shallow notches. Syrinx of the oscine type, but with only 3 pairs of intrinsic muscles; it is capable of a remarkable variety of loud ringing calls, and the birds are famous for their vocal mimicry. 2 species, dense forests; Tasmania (introduced) and eastern Australia.

      Family Atrichornithidae (scrub-birds (scrub-bird))

 Wholly unlike lyrebirds in external appearance, rather like large wrens, long-tailed, brownish, and terrestrial; 16.5 to 23 cm (6.5 to 9 inches). Virtually flightless, with very small wings and clavicles small and separated (not fused into a furcula, “wishbone”), a condition unique within passerines. Sternum unlike Menura, with 1 pair of deep posterior notches. Legs and feet very strong. Rictal bristles lacking. Similar to lyrebirds in the quality of their strong voices, nesting habits, and basic syringeal structure. Scrub-birds, however, have only 2 pairs of intrinsic muscles in the syrinx. 2 species; Atrichornis clamosus long thought extinct but recently rediscovered. Scrublands; in western and eastern Australia.

      Family Ptilonorhynchidae (bowerbirds (bowerbird))

 10-primaried oscines of medium size, 23 to 37 cm (about 9 to 15 inches); sexes alike in a few, but males usually much brighter coloured, often with nuchal (neck) crest but never with plumes or facial wattles. Stout bill, straight to slightly curved; wings rounded; legs and feet stout, hind toe shorter than middle toe. Solitary, largely terrestrial birds; eat berries, seeds, fruits, insects, small animals. Males of most species build elaborate stages or bowers for display and courtship. Loud ringing calls, good mimics. 8 genera, 20 species, in forests of New Guinea and northern Australia.

      Family Maluridae (Australian fairy wrens (fairy wren) or wren-warblers (wren-warbler))

 Small-bodied birds, 7.5 to 25 cm (3 to 10 inches), that carry the long tail cocked up over the back. Bill small, weak; wings short, rounded; legs and feet medium. Emu-wrens (Stipiturus) have rectrices reduced to 6 loose-barbed shafts. Most species are brightly patterned in contrasting browns, reds, shiny blues, black, and white; sexes alike or unalike. Food largely insects and other invertebrates. Many are good singers, mimics. About 5 genera, 29 species in Australo-Papuan region, including New Zealand; in forests, scrublands, heaths.

      Family Meliphagidae (honeyeaters (honeyeater))

 Small to medium-sized, 10 to 40 cm (4 to 16 inches); long, protractile, brush-tipped tongue curled at the sides to form a tube. Bill slender, pointed, downcurved, upper cutting edge serrated; nostrils unfeathered, with leathery operculum. Wings long, pointed, 10th primary about half the length of 9th; tail medium to long. Legs short to medium, tarsus scaled anteriorly. Drab browns, yellows, grays, or bold patterns of black, red, white; sexes alike or unalike. Food chiefly nectar, insects, some fruit. Voices loud, varied; musical song in many. About 43 genera, approximately 180 species, from Australia and New Zealand through the Papuan region, north to Marianas, east to Hawaii, 2 species in southern Africa. Habitat forests, brushlands, cultivated lands.

      Family Pardalotidae (pardalotes and bristlebirds)

 Small to medium-sized songbirds, 9–27 cm (3.5–11 inches). Pardalotes once allied to the similar flowerpeckers (Dicaeidae), but DNA studies revealed that they were relicts of old radiation of Australian songbirds. Plumage ranges from colourful to sombre brown. Bills stubby (pardalotes) or medium-length and pointed (bristlebirds). Pronounced bristles at bill base responsible for name of these terrestrial birds with pleasant squeaky songs. 2 genera, 7 species. Australia.

      Family Acanthizidae (Australian warblers)

 Tiny to small songbirds 8–12 cm (3.1–4.7 inches), some with beautiful songs. The weebill is Australia's smallest bird. Mostly drab brown and gray in colour and difficult to identify. Includes thornbills (Acanthiza) and fairy warblers (Gerygone). About 15 genera, 62 species. Australia, New Guinea, New Zealand, Indonesia, the Philippines, and Southeast Asia.

      Family Petroicidae (Australasian robins)

 Small thrushlike and chatlike songbirds, 11–22 cm (4.3–8.7 inches). Some flycatcher-like in habits, but also engage in wing and tail flicking. Drab brown to colourful (yellow and red) plumages. DNA revealed they are not related to similar species of Eurasia but instead part of the radiation of Australian songbirds related to other corvids. About 14 genera, approximately 46 species. Australia, New Zealand, and New Guinea through Southeast Asia, China, and the Philippines.

      Family Irenidae (leafbirds (leafbird) and ioras)

 Small to medium-sized birds, 13 to 25 cm (5 to 10 inches); bill fairly long, slightly downcurved or hooked; legs short; wings rounded; tail square to rounded. Like bulbuls, some have hairlike feathers on nape and long fluffy rump feathers but are more brightly coloured, brown or black with contrasting yellow, green, or blue; sexes usually dissimilar. Eat fruit, berries, buds, some insects. Good singers, musical whistles and fluty notes. 3 genera, 14 species in forests and cultivated lands from India to Philippines, south to Borneo, Java, Sumatra.

      Family Orthonychidae (logrunners and chowchillas)

 Medium-sized terrestrial birds, 17–28 cm (about 7–11 inches). Once linked to babblers of Asia and thought to be part of whipbird family (Eupetidae) but now known to be part of the endemic Australasian avifauna. Females build domed stick nests with side entrance. Bold brown, rufous black and white plumage colours. 3 species. Rainforests of Australia and New Guinea.

      Family Pomatostomatidae (Australo-Papuan babblers)

 Medium-sized terrestrial songbirds, 18–25 cm (7–10 inches), with long bills and tails, like mockingbirds (Mimidae) of the New World. Conspicuously social in family groups. Bold white throats, caps, eyelines, wing bars and tail tips highlight dark brown and rufous plumage. 2 genera, 5 species. Australia. New Guinea.

      Family Laniidae (shrikes (shrike))

 Rather small to medium-sized, 15 to 36 cm (6 to 14 inches); 10-primaried, with proportionately large heads; stout, strong, sometimes toothed, sharply hooked bills; strong legs with sharp claws; tarsus scutellate anteriorly, lamellate (plated) laterally; wings medium; tail usually long, narrow. Plumage soft, black, gray, or brown above, usually paler below in true shrikes; yellows, reds, greens in some African bush shrikes; sexes alike or unalike. Food large insects, small vertebrates; some impale prey on thorns for storage and to tear apart. Voices varied, some with well-developed song, several species sing antiphonally. 12 genera, 81 species. Essentially Old World, across temperate Eurasia, Africa, east to Philippines, and south to New Guinea and Timor; only 2 species in North America to southern Mexico; usually solitary birds, in open forests, clearings, brushlands, cultivated areas. Bush-shrikes of Africa increasingly treated as separate family (Malaconotidae).

      Family Vireonidae (vireos (vireo))

 Rather plainly coloured, small arboreal birds, 10 to 17.5 cm (4 to 7 inches), mostly brownish gray to olive green above, yellow, grayish, or white below; plumage never streaked or spotted; some with light eye-rings, eye stripes, wing bars; sexes alike. Bill fairly heavy, slightly hooked and notched; nostrils ovate, operculate, partly exposed; rictal bristles inconspicuous; legs short, strong. Wings long, pointed to short, rounded, 10th primary very short or vestigial. Family now includes peppershrikes (Cyclarhis) and shrike-vireos (Vireolanius) once separated in their own respective families. Usually solitary inhabitants of forest edges; seek insects on leaves and branches, eat some berries, fruit. Compulsive singers; songs of repeated, often melodious, phrases; harsh scolding notes. 4 genera, 58 species in all types of woodlands from central Canada to Uruguay and Argentina.

      Family Cinclosomatidae (quail-thrushes and whipbirds (whipbird))

 Medium-sized terrestrial birds, 17–30 cm (7–12 inches). Shy, secretive, terrestrial quail-thrushes boldly patterned with rufous, black and white colours. They flush fast with whirring wings like quail from hiding places. About 6 genera, approximately 16 species. Southeast Asia, Indonesia, Australia, and New Guinea.

      Family Pachycephalidae (whistlers (thickhead) and allies)

 A diverse group of small to medium-sized, stout-bodied birds, 13 to 28 cm (5 to 11 inches). Roundish heads; rather heavy bill sometimes hooked at the tip. Wings rather long, pointed, with very short 10th primary; tail medium long, usually rounded. A few species crested or with wattles or bare patches at base of bill. Most are greenish gray to brown above, lighter below, many with yellow or dull red markings; sexes alike, a few unalike; many juvenile plumages spotted or like that of female. Group includes not only the wide-ranging, colourful whistlers, but also shrike-tits (Falcunculus) and the poison-feathered pitohuis of New Guinea. Diet predominantly insects with some fruit. Melodious fluty calls, pairs often duet. 13 genera, about 60 species in Australo-Papuan region, Malaysia, Philippines, and Oceania; in forests, brushlands, mangroves, savannas.

      Family Corvidae (crows (crow), jays (jay), ravens (raven), rooks (rook), choughs (chough), nutcrackers (nutcracker), and magpies (magpie))

 Medium to large passerines, including the heaviest oscines, the ravens (Corvus); 17.5 to 70 cm (7 to 28 inches). 10 primaries, 10th (outer) always much shorter than 9th but longer than primary coverts. Bill strong, powerful, longer than rest of head or nearly so; nostrils usually covered with bristles. Crows (Corvus) and allies, large, black or black and gray or white; wings long, tail shorter than wing. Jays generally smaller, often coloured in blues, greens, yellows; wings rounded; tail sometimes two-thirds of total length. Voices harsh, loud. About 24 genera, approximately 124 species, almost cosmopolitan (absent from southernmost South America, Antarctica, some oceanic islands, introduced to New Zealand); varied habitats, prefer woodlands, open brushlands.

      Family Paradisaeidae (birds of paradise (bird-of-paradise))

 Small to medium-sized, 14 to 117 cm (about 5.5 to 46 inches), greatest length due to streaming tail feathers; 10-primaried; greatly varied colours, most males with spectacular plumes on head, flanks, wings, or tail, some with wattles or bare skin on the head; females plain browns or grays. Bill slender to rather heavy, hooked or sickle-shaped in some. Wings rounded; legs short; feet rather stout. Solitary forest birds, rather weak fliers; males of most species have elaborate courtship displays. Eat fruit, seeds, insects, small animals. Prolonged whistles, loud shrill calls. About 18 genera, approximately 46 species; New Guinea, northern and eastern Australia, Moluccas, and adjacent islands.

      Family Corcoracidae (white-winged chough and apostlebird)

 2 genera, 2 species.

      Family Grallinidae (mudnest builders (Grallinidae))

 10-primaried oscines of medium size, 19 to 50 cm (7.5 to 20 inches). Differ from most corvids in lacking nasal bristles. Legs long, strong; wings long and pointed to short and rounded. Blackish to gray in colour. Usually feed on the ground. Distinguished and united by their unique bowl-like mud nests stiffened with hair, feathers, and grass. Have complex communal social units. Weak fliers with peculiar jumping gait. Eat insects, snails, seeds, soft fruit. Melodious whistles, harsh and plaintive notes. 2 genera, 2 species; limited to Australia and western New Guinea, in woodlands, marshes, cultivated lands, usually near water.

      Family Cracticidae (bell-magpies (bell-magpie))

 Medium to large oscines, 25 to 58 cm (10 to 23 inches). Most black, white, and gray; some have brown phase; sexes alike or unalike. Large, heavy bill, slightly to strongly hooked; nostrils bare. Wings long, pointed; legs strong, medium to long. Eat insects, small animals, some fruit and seeds. Fine singers with ringing, gonglike calls. About 5 genera, approximately 13 species. Australia, New Guinea, and nearby islands; gregarious inhabitants of brushy plains, open forests, mangrove shores, cleared lands.

      Family Artamidae (woodswallows (woodswallow) or swallow-shrikes)

 Chunky-bodied, medium-sized, 15 to 21 cm (6 to 8.5 inches); unique among oscines in having powder downs. Bill stout; broad at base, moderately long, decurved, pointed; legs short; feet strong. Wings long, pointed; tail short, nearly square. Plumage compact, soft, in plain solid browns, grays, or black above, usually lighter below; sexes similar. Sometimes roost communally. Live on insects caught in flight. Voice a harsh nasal twittering. 2 genera, about 16 species; in open lands, forest clearings, Australia east to Fiji Islands, north to Philippines, Indochina, India.

      Family Oriolidae (Old World orioles (oriole) and figbirds)

 Medium-sized birds, 18 to 30.5 cm (7 to 12 inches); brightly coloured, predominantly in yellows, greens, and black; sexes unalike, female duller, young streaked below. Bill strong, pointed, slightly hooked; long, pointed wings with 10 primaries; medium to long tail of 12 feathers. Strictly arboreal birds that feed on insects, fruit. Flight strong, undulating. Voices loud, with harsh calls, but melodious songs. 2 genera, 29 species in forest and open woodland across Eurasia, in Africa, East Indies, Australia, Philippines. For New World orioles, see below Family Icteridae.

      Family Campephagidae (cuckoo-shrikes (cuckoo-shrike) and minivets (minivet))

 Small to medium-sized birds, 12.5 to 35.5 cm (5 to 14 inches). Rather stout, slightly downcurved bill, notched, hooked at tip; nostrils partly concealed by short bristles; wings medium, pointed; long tail rounded or graduated; legs short, feet weak to strong. Feathers of back and rump usually with heavy shafts, thickly matted and loosely attached. Cuckoo-shrikes plain gray, black, or whitish birds, often with barred underparts; minivets brightly coloured in reds, yellows. Sexes often completely dissimilar. Eat insects, berries, small fruits. Noisy birds, with whistles and harsh calls. About 7 genera, approximately 80 species. In forests; Africa, India to Japan, East Indies, Australia.

      Family Rhipiduridae (fantails (fantail))

 Small, active songbirds, 15–21 cm (5.9–8.3 inches), all similar in proportions and behaviour; hence just 1 genus. Long tails of rounded and fanned shape which are waved in a distinctive pattern—that is, side-to-side through an upward arc, while holding their wings downward. Most widespread Australian fantail species is the Willie Wagtail (Rhipidura leucophrys). 47 species. India, Southeast Asia, Australasia, New Zealand, and Pacific islands.

      Family Dicruridae (drongos (drongo))

 Small to medium-sized birds, 18 to 63.5 cm (7 to 25 inches), the longest being those with exceptionally long tails; usually black with purple or greenish sheen, some crested or with spangled neck and head feathers, iris of eye usually red; sexes alike. Bill stout, arched, slightly hooked and notched; long, strong rictal bristles; legs short, feet stout. Wings long; tail variable, of 10 or 12 feathers, usually forked; some species with racquet tails. Arboreal birds that fly well but seldom long or far. Food mostly insects caught on the wing. Voice varied, melodious; capable mimics. 2 genera, 24 species. Africa, South Asia, Malaysia, northern Australia, and east to Solomon Islands, in woodlands, savannas, and cultivated regions.

      Family Monarchidae (monarch flycatchers (monarch))

 Small to medium-sized oscines, 13–53 cm (5–21 inches). Many brightly and or boldly coloured. Some species, especially paradise flycatchers, have spectacularly long tails. Insectivorous, woodland birds with few exceptions. Some are sit-and-wait predators, others are active searchers. Some are sexually dimorphic. Bill morphologies vary from small and slender to the huge bill of yellow-breasted boatbill (Machaerirhynchus flaviventer). Once classified with Old World flycatchers (Muscicapidae), the other mudnest builders (Corcoracidae). Magpie-lark is now included in this family, based on DNA studies. About 19 genera, approximately 100 species. Widespread through Africa, southern Asia, New Guinea, Australia, and Pacific islands.

      Family Prionopidae (helmet shrikes (helmet-shrike) and allies)

 Small to medium-sized, 10 to 25.5 cm (4 to 10 inches). Diverse group. Family expanded on the basis of DNA studies to include flycatcherlike wattle-eyes and batises. Most with conspicuous wattles surrounding the eyes. Bill stout, hooked; legs short. Plumage black above, white or buffy below in bold patterns; sexes similar. Helmet shrikes are gregarious arboreal birds that hunt in small flocks for insects. Other species more solitary. Harsh chattering notes, nasal humming sounds; some species snap bills audibly. 6 genera, about 44 species, in brushlands, scrub, and forests of Africa, south of the Sahara.

      Family Vangidae (vanga shrikes (vanga-shrike))

 Small to medium-sized, 12.5 to 32.5 (5 to 13 inches), 10-primaried, arboreal oscines of varied aspect; some resemble flycatchers, thrushes, nuthatches, or typical shrikes. Bill stout, heavy, hooked, notched; with enlarged ridge in 1 part, thin, downcurved in another; legs and feet strong; wings fairly long, rounded; tail square or rounded, moderately long. Plumage rather soft, loose, typically black or blue above, white below, some with brown or gray patches; sexes alike or unalike. Eat insects, small reptiles, amphibians. Chattering calls, harsh notes, shrill whistles. 13 genera, 15 species, restricted to forests, scrublands, mangrove swamps, of Madagascar.

      Family Callaeatidae (wattlebirds (wattlebird))

 Medium-sized, 25.5 to 53.5 cm (10 to 21 inches); black, brown, or blue-gray, with fleshy blue or orange wattles at the gape; sexes may differ in size, wattles, and bill shape. Weak fliers but hop strongly. Sternum weak; 10-primaried wings short, rounded; tail long. Bill stout and short to long and curved; legs long, feet stout. Eat fruit, nectar, insects. A variety of musical notes and whistles. 2 species limited to primeval forests of New Zealand; huia (Heteralocha) extinct; wattlebird (Callaeas) and saddleback (Creadion) rare.

      Family Bombycillidae (waxwings (waxwing))

 Small, arboreal, soft-plumaged, crested, 15.5 to 19 cm (6 to 7.5 inches); distinguished by waxy red “droplets” at tips of secondary wing feathers of most individuals. Bill short, swollen, slightly hooked; nostrils almost concealed by feathers, rictal bristles absent. Long, pointed wing with rudimentary 10th primary; square to slightly rounded, short tail. Velevety plumage blended with browns, grays, and yellow; tail tipped with yellow or red; sexes alike. Eat berries, fruit, buds, flowers, insects. Weak chattering song, soft lisping calls. 3 species in temperate Northern Hemisphere, in evergreen or birch forests.

      Family Ptilogonatidae (silky flycatchers (silky flycatcher))

 Small, arboreal, crested, 18 to 24.5 cm (7 to 9.5 inches). Bill short, broad, deeply cleft; nostrils exposed, bordered by membrane, rictal bristles present. Wings and legs rather short; 10th primary well developed; tail long. Soft silky plumage of solid grays, black, brown, white spots in wings and tail, some with yellow markings. Rather shy, active birds that feed on berries and on insects caught in flight. Weak warbling song. 3 genera, 4 species, found from southwestern U.S. to Panama; in brush country, desert scrub, open forest.

      Family Dulidae ( palm-chat)

 Medium-sized, about 19 cm (7.5 inches); deep laterally compressed bill, strongly curved culmen, circular nostrils wholly exposed. Wings rounded, 10th primary less than half as long as 9th; tail longish; legs and toes stout. Plumage stiff and harsh, olive brown above, yellowish white streaked with brown below; sexes alike. Lives on fruits and flowers. Noisy, gregarious birds with harsh chattering notes. The single species limited to cultivated areas and open woodlands of Hispaniola and Gonave Island, West Indies.

      Family Cinclidae (dippers (dipper))

 The only largely aquatic oscines; small birds, 14 to 19 cm (5.5 to 7.5 inches), with plump bodies, short concave wings, short, square or rounded tail. Tarsi and toes long and stout. Bill slender, straight, pointed; nostrils with broad operculum; rictal bristles absent. Dense compact plumage plain gray, brown, or black, some with white patches on throat, breast, head; sexes alike. Food: insects, aquatic larvae, and other organisms gleaned from stream bottoms. Have shrill penetrating whistled calls and chattering songs. 5 species, found along swift, rocky streams in Eurasia, North Africa, and North and South America, south to Argentina.

      Family Turdidae (thrushes (thrush))

 A large, almost cosmopolitan group of small to medium-sized birds, 11.5 to 33 cm (4.5 to 13 inches). Bill rather slender; legs and feet fairly stout, tarsus usually booted (smooth sheath, not divided into scales); 10-primaried wing rounded to pointed; tail medium, truncate or graduated, forked in a few. Colours brown, blue, gray, often blended, or in bold black, white, yellow, or red patterns; sexes alike or unalike; young usually spotted below. Eat mostly fruit, insects, also seeds, leaves, worms, and mollusks. Most are fine singers. About 21 genera, approximately 184 species. Virtually worldwide; absent originally only from New Zealand (now introduced there), some oceanic islands, Antarctica, and parts of the Arctic. Usually arboreal, many terrestrial, in varied habitats—forests, deserts, brushlands, grasslands, cultivated fields.

      Family Muscicapidae (Old World flycatchers (flycatcher), chats (chat), and wheatears (wheatear))

 A large family of small insectivores, 7.5 to 22.5 cm (3 to 9 inches). The flycatcher forms with typically flat, broad bills, well-developed rictal bristles, and short, weak legs and feet. The thrushlike forms include chats, redstarts, scrub-robins, wheatears, and forktails. Wings short and rounded to long and pointed; tail short. Varied colours, many plain browns, grays, some with bright blues, reds, and black and white; some crested or with facial wattles; sexes alike and unalike, young usually spotted. Voices usually weak, monotonous, but well developed, song in a few. About 48 genera, approximately 250 species; Eurasia from tree line south through Africa, Australia, and in Pacific islands to Hawaii; forests, scrublands, cultivated and riverine areas.

      Family Platysteiridae (wattle-eyes (wattle-eye))

 About 6 genera, approximately 31 species.

      Family Sturnidae (starlings (starling), mynahs (mynah), and oxpeckers (oxpecker))

 Stocky, medium-sized, 16.5 to 42 cm (6.5 to 16.5 inches); very short but visible 10th primary. Bill pointed, straight, or slightly arched, swollen near the tip in oxpeckers (Buphagus); tongue flat, not tubular (as in some relatives). Legs and feet stout, strong, tarsus with unbroken plates behind. Typically dark coloured with metallic sheen, some brown or gray with white, yellow, or red markings, some with facial wattles, a few crested; sexes alike or unalike. Almost omnivorous; wide range of vegetable and animal foods. Garrulous, varied notes, calls, whistles, some excellent mimics. About 26 genera, approximately 148 species in all types of wooded and agricultural lands; temperate Eurasia through Africa, northern Australia, East Indies, east to Tuamotu Archipelago; a few introduced widely elsewhere (such as North America).

      Family Mimidae (mockingbirds (mockingbird), catbirds (catbird), and thrashers (thrasher))

 Slender, medium-sized, 20 to about 30 cm (8 to 12 inches); bill medium to long, nearly straight to strongly downcurved; nostrils always exposed, with overhanging membrane; rictal bristles present, but few and somewhat weak. Legs rather long, feet strong, middle toe joined at base to outer toe but not to inner toe. Wings short, rounded; tail long. Coloured brown, gray, black, bluish, underparts usually pale, often white, spotted, or streaked, some solidly coloured, some with white in wings or tail; sexes alike. Eat fruit, seeds, insects. Famous as fine singers and mimics. Exclusively New World, the 11 genera and approximately 35 species range from southern Canada, the West Indies, Galapagos, to southern Argentina and Chile.

      Family Sittidae (nuthatches (nuthatch))

 Small, stocky, climbing birds, 9.5 to 19 cm (3.5 to 7.5 inches), with thin, pointed, usually straight bills; rounded, nonoperculate nostrils partly concealed by feathers; short rictal bristles present; tarsus short; long, laterally compressed claws on large toes, hallux equal to outer toe, inner toe reaching only to second joint of middle toe; wings rather long, pointed; tail short, square, soft. Typically gray to blue above, white or brownish below; sexes alike or nearly so. Forage on tree trunks and large branches for insects, eat some seeds and small fruits. Simple call notes and songs. 2 genera, 25 species; typical nuthatches (Sitta) distributed across North America and Eurasia to Malaysia and the Philippines, aberrant forms in Philippines, Australo-Papuan region. Typically in forests, a few in rocky areas.

      Family Certhiidae (creepers (creeper))

 Small, slender, climbing birds, 9.5 to 19 cm (3.5 to 7.5 inches), with curved bills as long as or longer than rest of head; operculate nostrils free of bristles or feathers; rictal bristles absent. Legs short, thin, outer toe always shorter than middle toe but much longer than inner toe; claws long, hind claw as long as or longer than hind toe; wings rounded or pointed; tail feathers long, stiff, with pointed tips. Brownish above, streaked with black or light brown, white below; sexes alike. Solitary birds, spiral up tree trunks from base probing bark for insects. Soft calls, sweet but weak songs. 2 genera, 7 species (with the inclusion of the African spotted creeper, Salpornis, a controversial point) in temperate woodlands of Eurasia, Africa, North America, south to Nicaragua. 2 species inhabit rocky cliffs.

      Family Troglodytidae (wrens (wren))

 Small, chunky birds, 9.5 to 22 cm (3.5 to 8.5 inches), mostly brown, usually barred, spotted, or streaked, with white, black, or browns; sexes alike. Bill slender, medium to long, often downcurved; nostrils with operculum; rictal bristles usually indistinct but sometimes obvious, with 1 or 2 well developed. Stout legs and feet, front toes partly joined at base. Wings short, rounded, well-developed 10th (outermost) primary, at least half as long as 9th; tail short, square to rounded, often carried cocked up. Most forage on or near ground in undergrowth for insects, worms, other invertebrates. Highly developed song, musical, variety of bubbling, flutelike to growling notes, harsh chattering calls; some species duet. About 16 genera, approximately 75 species range through most of North and South America; 1 species (Troglodytes troglodytes) also found in Eurasia and North Africa; inhabit brushlands, forests, forest edges, rocky slopes, deserts, grassy marshes.

      Family Polioptilidae (gnatcatchers (gnatcatcher) and gnatwrens)

 Dainty, slender, tiny, 10 to 14 cm (4 to 5.5 inches), with long, thin, pointed bills, operculate nostrils partly exposed, and rictal bristles. Rounded wing with 10th primary much less than half as long as 9th; long, rounded tail constantly moving. Blue-gray or brown above, lighter below, most with white in outer tail feathers, some with black markings on head; sexes alike or nearly so. Food essentially small insects. Rather weak songs of trills and warbles, simple short call notes. 3 genera, 15 species. Active arboreal birds of open forests and semiarid deserts from southern Canada to Argentina.

      Family Paridae (titmice (titmouse) and chickadees (chickadee))

 10-primaried oscines with short to medium rounded wings; tail short to long; bill rather stout, pointed, shorter than remainder of head; nonoperculate nostrils concealed by thick feathers. Small birds, 7.5 to 20 cm (3 to 8 inches), with thick plumage usually strongly patterned in grays, yellows, browns, black, or white, never streaked, barred, or spotted; sexes usually alike. Active, gregarious little birds; feed on insects, seeds. Many species use advanced spatial memory to relocate stored food caches. Chattering notes, whistled calls. About 5 genera, 55 species in forests and brushlands of the Philippines, Malaysia, Eurasia, Africa, and North America to Guatemala.

      Family Remizidae (penduline tits)

 Tiny songbirds, 7.5–11 cm (3–4.3 inches), closely related to the true tits (Paridae). Bill short, conical and sharply pointed. Wing short and rounded with 10 primaries. Mostly gray in colour, some species with yellow. Sexes similar. Acrobatic and often hang upside down to find food. Use feet to hold food. Build elaborate suspended nests of soft materials. Includes verdin of deserts of southwestern United States. 5 genera, approximately 12 species. Eurasia, southern Asia, Africa, and North America.

      Family Aegithalidae (long-tailed tits)

 Very small songbirds, 10–12 cm (4–5 inches), with short wings and long tails. Stubby, short bill. Long, slender legs. Fluffy plumage mostly gray or brown in colour, some species with bold black and white face patterns. Black or pale eyes. Active, acrobatic insect feeders. Highly social, defending group territories and nesting cooperatively. Sexes similar. Not related to Paridae as previously thought. Includes bushtit of North America. 3 genera, 8 species. Eurasia, western North America.

      Family Hirundinidae (swallows (swallow))

 A distinctive family placed low on the oscine family tree because of primitive syringeal characters, particularly the double bronchial rings, unique in Passeri. Small birds, 9.5 to 23 cm (about 4 to 9 inches), with compact plumage, often with metallic sheen, usually lighter below; sexes usually alike or nearly so. Wings long, pointed, primaries reduced to 9. Tail medium to long, truncate to forked. Legs short, feet small, weak. Bill short, flat, but wide gape enhanced by rictal bristles. Walk with difficulty but fly strongly, feed on insects caught in flight. Voice usually a twittering or squealing, sometimes melodious. The river martins of Africa and Thailand differ by large, brightly coloured bills, stronger feet, and syringeal characters. About 15 genera, approximately 90 species. Worldwide except polar regions and certain oceanic islands; many are migratory.

      Family Regulidae (kinglets (kinglet))

 Tiny, active songbirds with short slender bills and drab olive plumage except for colourful crest feathers that are inconspicuous unless displayed. Often members of mixed flocks in coniferous forests. DNA studies reveal they are not closely related to leaf warblers (Phylloscopus), as once thought, and may be an ancient oscine lineage. 1 genus, 6 species. Eurasia and North America.

      Family Pycnonotidae (bulbuls (bulbul))

 Medium-sized birds, 14 to 28 cm (5.5 to 11 inches), with soft, fluffy plumage, especially soft on lower back and rump; many species with hairlike, vaneless feathers on nape of neck. Bill usually slender, slightly downcurved; rictal bristles well developed; feet and legs rather small; wings short; tail medium to long. Usually drably coloured in gray, brown, greenish, some with yellow, white, or red patches on head and under tail; sexes alike. Eat berries, fruit, some insects. Noisy, some good singers, mimics. About 21 genera, approximately 142 species. Africa, across southern Asia to Japan, Philippines, Moluccas, and Borneo; in woodlands, brushlands, cultivated regions.

      Family Hypocoliidae ( hypocoly)

 Medium-sized, 17.5 cm (7 inches); short, broad bill, moderately curved culmen, operculate nostrils; nasal bristles lacking and rictal bristles poorly developed. Tarsus rather short, heavily scutellated (scaled). Soft plumage blue-gray above, lighter below, with black facial, wing, and tail markings, white wing tips, slightly crested; sexes alike. Fruit eater with low calls and weak song. The single species inhabits semiarid scrublands, palm groves, gardens in Tigris-Euphrates valley of Iraq, wanders to Afghanistan, western India, Arabia, Red Sea coast.

      Family Zosteropidae (white-eyes (white-eye))

 Relatively uniform group of little birds, 10 to 15 cm (4 to 6 inches) long, yellowish or reddish brown to olive green above, lighter below, typically with a conspicuous white eye-ring; sexes alike. Bill short, slender, pointed, slightly decurved; tongue brush-tipped; rictal and nasal bristles absent. Wings short, rounded, 10th primary usually lacking; tail medium, square; legs and claws short, feet strong. Travel in restless flocks, feed on nectar, insects, fruit. Weak voices, pleasant twittering, warbling songs. 13 genera, approximately 110 species in forests, brush country, mangroves; Africa south of Sahara, across southern Asia to Korea, Japan, south to Australia, New Zealand, east to Carolines, Samoa.

      Family Sylviidae (Old World warblers (warbler) and wrentits (wrentit))

 Typically small, 9 to 26 cm (3.5 to 10 inches), with slender bill; longitudinal nostrils with an operculum; medium rounded wings of 10 primaries; short to medium legs. Plumage usually browns, grays, olive greens with little pattern; some streaked or barred; some brightly, boldly marked; sexes alike or similar, young never spotted below. DNA studies suggest a broadened definition of this family to include babblers (Timaliidae), wrentit (Chamaea fasciata), and parrotbills (Panuridae). Of uncertain affinities to these species are the odd rock fowl (Picathartes) and rock jumpers (Chaetops) of Africa. African warblers split by some as the family Cisticolidae. 119 genera, nearly 300 species, worldwide except in polar regions, South America, some oceanic islands, mostly in woodlands, some in brushlands, marshes. Food essentially insects; voices pleasant, varied, well-developed song in some.

      Family Timaliidae (babblers)

 50 genera, 265 species.

      Family Alaudidae (larks (lark))

 A distinctive and well-defined group, the only oscines with back of tarsus rounded and scaled instead of sharp and unsegmented. Syrinx also unique among Passeri in lacking the usual bony pessulus. Small ground birds, 12 to 23 cm (about 5 to 9 inches), usually cryptically coloured in browns and grayish buffs, plain or streaked, lighter below (several species black below); sexes similar. Bill usually pointed, slightly downcurved; wings long, pointed; legs rather long, hind toe usually with long, straight claw. Food: seeds, insects, and other invertebrates. Sing beautifully, often a soaring flight song. About 18 genera, approximately 93 species. Open fields, plains, beaches of Old World, with only one species, the horned lark (Eremophila alpestris), also in North America south to Colombia. Many species migratory.

      Family Dicaeidae (flowerpeckers (flowerpecker))

 Small, chunky birds, 7.5 to 18 cm (3 to 7 inches), with short necks, legs, tails. Bill usually short, stout, relatively straight; slender and curved in a few, edges of distal (outer) 3rd serrated; distal half of short tongue deeply cleft, the edges curled into 2 slender semitubular tips. Wings rather long, 10th primary usually vestigial, absent in a few. Bright-coloured males dark, glossy (rarely metallic) above, lighter below, often with red or yellow patches on crown, rump, breast; females usually much duller; some species dull-coloured in both sexes. Active arboreal birds, feed around flowers on nectar, small insects, berries (particularly mistletoe). Sharp metallic twittering calls, a few have warbling song. 2 genera, 44 species; in forests, scrublands; India to southern China, Philippines, east through Solomons, and south to Australia.

      Family Nectariniidae (sunbirds (sunbird) and spiderhunters)

 Small, 9 to 22 cm (3.5 to 8.5 inches). Long, slender, downcurved, pointed bill, finely serrated near tip; tongue not brush-tipped but partly tubular, projectile, divided at tip; nostrils rounded, open, operculate; no rictal or nasal bristles. Wings rounded, short, 10th primary variable in length but always present; tail square, medium to long, pointed, sometimes with elongated central feathers. Legs short, stout, tarsus with anterior transverse scales; hallux and claws short. Many sunbirds with vivid colours, metallic sheens, and bright patches in male; females much duller; other sunbirds and spider hunters dully coloured in both sexes. Active arboreal birds, sunbirds are Old World counterparts of New World hummingbirds but not as skilled fliers. Feed on nectar, insects, some small fruit. Weak call notes, faint songs. The 2 species of sugarbirds (Promerops) of South Africa may be related to either honeyeaters (Meliphagidae) or starlings (Sturnidae). About 13 genera, approximately 131 species. Forests, scrub country, mangroves; Asia and Africa south of Sahara, east to central China, Philippines, Solomons, south through Malaya to northern Australia.

      Family Melanocharitidae (berry-peckers)

 Small to medium-sized songbirds, 9–21 cm (3.5–8 inches), of uncertain affinities. Colours vary from drab olive to dramatic blue and gray with yellow, or yellow and black. Sexually dimorphic. Resemble small honeyeaters in general behaviour. Hover-glean small fruits and invertebrates. Some with small, stubby bills. Cup-shaped nests built in fork of a branch. About 5 genera, 12 species. New Guinea.

      Family Motacillidae (pipits (pipit) and wagtails (wagtail))

 Small, slender-bodied ground birds, 12.5 to 23 cm (5 to 9 inches). Pipits similar to larks in appearance but differ in having a bilaminate tarsus and pointed wing with 9 primaries. Wagtails have longer tails, brighter colours. Bill thin, pointed; legs long, slim, with elongated hind toe and hind claw (with some exceptions); tail usually edged with white or yellow. Plumage brown, streaked, mottled, sexes similar (pipits); or marked in bold black, white, or yellow patterns, sexes unalike (wagtails). Feed on insects, spiders, mollusks, some vegetable matter. Simple repetitive song often given in flight; short, sharp call notes. About 5 genera, approximately 64 species; worldwide except polar regions and some oceanic islands; in open grasslands, deserts, shores, cultivated areas.

      Family Prunellidae (accentors (accentor) and hedge sparrows)

 Small, drab, 12.5 to 17.5 cm (5 to 7 inches); slender, pointed bills, wide at base, culmen (ridge of upper bill) slightly rounded. Tarsus rather short, feet strong. Wings rounded to pointed with very short 10th primary; tail shorter than wing, square or emarginate. Colour browns or grays, usually streaked or spotted; sexes similar. Feed on or near ground on insects, berries, seeds. Thin chattering, twittering songs, metallic calls, often in flight. 1 genus (Prunella), 13 species; across Eurasia and northern Africa; various habitats, parks and gardens, brushlands, barren mountain slopes below snow line.

      Family Ploceidae (weaverfinches, queleas (quelea), bishops (bishop), and allies)

 Small, stoutly built seedeaters, mostly 10 to 25 cm (4 to 10 inches), a few long-tailed species to 50 cm (20 inches; males only). Short, stout, conical, pointed bills; gonys less than half length of upper bill; rictal bristles lacking; tarsus relatively short. Wings short to long, rounded to pointed, 10th primary present, much reduced; tail short to very long. Colours, patterns widely varied; sexes alike or unalike. In some classifications, Old World sparrows (Passer) and rock-sparrows (Petronia) of Passeridae are included here in Ploceidae. Usually gregarious; food mainly seeds, some other vegetable matter, insects. Harsh, monotonous voices, song poorly developed. About 19 genera, approximately 148 species in all terrestrial habitats, mainly in Africa, also in Eurasia, Madagascar, Malaysia; a few (such as the house sparrow, Passer domesticus) widely introduced elsewhere.

      Family Estrildidae (waxbills (waxbill), whydahs (whydah), and allies)

 Small, 7.5 to 15 cm (3 to 6 inches), bill stout, short, pointed; gonys less than half the length of upper bill. Wings rounded, short to pointed, medium length; 10th primary present but almost vestigial; tarsus relatively short. Great variety of bright and sombre colours and patterns; sexes alike or unalike. Highly gregarious, active, ground feeders. Food: seeds, some berries, insects. Song poorly developed, weak chirps, chatterings, buzzes. 29 genera, approximately 162 species; forest edges, clearings, grasslands, and marshes of Africa, southern Asia, East Indies, Australia, some South Pacific islands.

      Family Fringillidae (cardueline finches and chaffinches (chaffinch))

 Small to medium-sized, 10 to 25 cm (4 to 10 inches), with short, stout to slender pointed bills (mandibles crossed in one genus); gonys less than half length of upper bill. Wings somewhat rounded, medium length, 10th primary present but less than half length of 9th; tarsus relatively short. Colours variable but browns, reds, and yellows predominate; some streaked or mottled, some plain patterns; sexes alike or unalike. Gregarious birds with strong, undulating flight. Eat seeds, buds, berries, some insects. Highly developed songs. About 22 genera, approximately 144 species. Worldwide in woodlands and brushlands except Pacific Islands; introduced in Australia.

      Family Drepanididae (Hawaiian honeycreepers (Hawaiian honeycreeper))

 Small forest birds, 11.5 to 22 cm (4.5 to 8.5 inches), limited to the Hawaiian Islands; evolved from fringillid ancestors. Bill extremely varied among species: short to long, thin to thick, straight to extremely downcurved, pointed to hooked, but never serrate or notched. Wings pointed, 10th primary vestigial or absent; tail medium, truncate or slightly forked; legs short to medium; feet strong. Plumage plain brown, olive green, yellow, red, gray, or black, never metallic or glossy; sexes unalike or similar with female smaller. Found solitary or in small flocks. Feed on nectar, insects, seeds, fruit. Warbling songs, clear calls. About 18 genera and approximately 23 species living, most of them rare and local; remnants of more-diverse assemblage now known only from subfossil remains.

      Family Emberizidae (buntings (bunting), New World sparrows (sparrow), and allies)

 Small, 9 primaries; 10 to 27 cm (4 to 11 inches), with short, stout, conical, pointed bills. Commissure distinctly angled or deflexed at the base, mandibular tomium elevated, often toothed, gonys (midline ridge of lower bill) more than half length of upper bill; rictal bristles usually obvious. Wings short, rounded to long, pointed; tail short to long, tarsus relatively long. Colours and patterns varied; sexes alike or unalike. Gregarious or solitary, terrestrial or arboreal birds; food mainly seeds, some other vegetable matter, insects. Call a simple chirp, often well-developed song; flight song rare. Family includes buntings, juncos, longspurs, Darwin's finches, yellowhammer (Emberiza citrinella), and most sparrows. About 73 genera, approximately 300 species almost cosmopolitan (absent from southern Indonesia, Madagascar, Antarctica; introduced in Australia, New Zealand) in all terrestrial habitats.

      Family Parulidae (wood warblers (wood warbler))

 Dainty, small, 10 to 18.5 cm (4 to 7.5 inches); pointed wings of 9 primaries, medium 12-feathered tail. Bill usually slender, pointed, culmen slightly downcurved, flattened with pronounced rictal bristles in a few but never notched or hooked; tongue moderately slender, tip variably bifid (divided) or fimbriate (fringed). Legs medium, hind claw never elongated. Colours varied but never metallic or glossy; grays and olive browns dominant ground colours, often brightly patterned with yellows, reds, blue, black, or white; sexes alike or unalike. Active, arboreal (a few terrestrial); feed mostly on insects, occasional fruits, berries, seeds. Voices typically weak, thin, high-pitched; a few with loud and well-developed songs. Includes the wrenthrush (Zeledonia), a shy resident of the wet rainforests of Costa Rica and Panama, previously classified as a separate family. About 25 genera, approximately 120 species in forests and brushlands from tree line of North America to southern South America.

      Family Thraupidae (tanagers (tanager) and allies)

 Small to medium, compactly built, 9 primaries; 8 to 30 cm (about 3 to 12 inches), most under 20 cm (about 8 inches). Bill with commissure (meeting of edges) not abruptly angled or bent at the base; mandibular tomium not distinctly angled (never toothed) near the base. Bill usually conical, short to medium, generally notched, toothed, or hooked at tip; rictal bristles present. Nectar-feeding species with specialized bill and brush-tipped tongue. Wings pointed, short to long; tail short to medium, truncate, emarginate, or rounded. Generally brightly coloured, in contrasting patterns of black, white, yellow, red, blue, green, brown; sexes alike or unalike. Includes euphonias (Euphonia), honeycreepers (Cyanerpes), conebills (Conirostrom), flower piercers (Diglossa), plushcap (Catamblyrhynchus), and the swallow tanager (Tersina). Some classifications move tanagers of genus Piranga to Cardinalidae, based on DNA phylogeny. Arboreal birds; flight strong but not sustained in nonmigratory species. Eat berries, fruit, insects, nectar. Voices varied, song well developed in a few but usually short calls and warblings. About l04 genera, approximately 416 species in forests, brushlands; temperate North America to Brazil, Argentina.

      Family Cardinalidae (cardinals (cardinal) and allies)

 Medium-sized, colourful songbirds, 11–28 cm (4–11 inches). Cardinals and grosbeaks have large bills for cracking seeds; buntings have smaller but still robust bills. Versatile diets include seeds, fruits, insects and flowers. Mostly monogamous. Build open cup nests. Loud, wonderful songs. Birds of both scrub and open woodlands. Many taxonomic questions remain. About 13 genera, approximately 46 species.

      Family Icteridae (New World orioles (oriole) and allies)

 A heterogeneous group, medium to large, 16 to 54 cm (about 6 to 21 inches). Straight, pointed, unnotched, conical bill, with culmen extending more or less onto forehead; operculate nostrils, never concealed; rictal bristles lacking. Wings mostly long, pointed; tail short to rather long; legs and feet strong. Plumage often solid black with metallic gloss, or multicoloured in bold patterns of black and red, orange, yellow, brown; sexes usually differ, female smaller. Many species gregarious; some colonial or semicolonial nesters. Includes traditional blackbirds as well as oropendolas and caciques. Eat a wide range of animal and vegetable foods. Usually loud-voiced, with harsh, whistling, or bubbling calls; well-developed song in some. About 29 genera, approximately 112 species. Distributed throughout Western Hemisphere (except polar regions), in a wide variety of habitats.

Critical appraisal

      Since the late 19th century, when the many advances in taxonomic thought of the previous century began to crystallize and bear fruit, three main passerine sequences have dominated the world bird lists. The first, proposed originally by English ornithologist R.B. Sharpe of the British Museum in 1877 but based in part on the published and unpublished work of others, placed the crows (crow) at the summit of avian evolution (hence at the end of a modern lineal sequence), ostensibly on the basis of their alleged high intelligence. This sequence was adopted by the German ornithologist Ernst Hartert in his monumental Die Vögel der paläarktischen Fauna (“The Birds of the Palearctic Fauna”), published in 1903, and subsequently by most other European ornithologists. The second sequence, which placed the thrushes (thrush) at the end, was in general usage, particularly in North America, until the late 1920s. The third sequence, originally proposed in 1926 by two American ornithologists, Alexander Wetmore (Wetmore, Alexander) and Waldron De-Witt Miller, but also based partly on the earlier work of others, placed the crows near the base of the oscine family tree and placed the so-called nine-primaried oscines, dominated by the seed-eating fringillids, at the top. This sequence immediately became the standard for North American and certain international works. It has remained so ever since, with only minor departures and rearrangements.

      In one case, a group of small South American ground birds known as gnateaters (gnateater) and antpipits (antpipit) was separated from the antbirds (antbird) in 1882 as the family Conopophagidae; this arrangement was generally recognized for nearly 90 years. Careful study in 1968, however, revealed that the family was an artificial one; one genus (Conopophaga) has been placed in family Conopophagidae, the other (Corythopis) placed in the Tyrannidae.

      DNA studies since the 1990s have clarified many of the relationships between suboscine birds. From DNA evidence, many authorities suggest that antbirds themselves make up two distinct assemblages of species and that they warrant recognition as separate families. Some ornithologists divide this group into the “typical antbirds” contained within family Thamnophilidae (which include antbirds, antwrens, and antshrikes) and the ground antbirds contained within family Formicariidae (which include ant-thrushes and antpittas). In addition, the enigmatic asities (asity) (Philepittidae) of Madagascar are allied to the broadbills (broadbill) (Eurylaimidae) based on internal anatomy and biochemical data. DNA analysis has also supports the placement of sharpbills (sharpbill) (Oxyruncus) and leaf-eating plantcutters (plantcutter) (Phytotoma) in Cotingidae.

      Within the oscines, one of the greatest problems is a satisfactory delineation and arrangement of the many superficially similar groups. Most taxonomists agree that the oscines contain three large groups: the crows, Old World orioles (oriole), birds of paradise (bird-of-paradise), and diverse insect-eating birds of Australia and New Guinea, (2) the thrushes, babblers, Old World flycatchers (flycatcher), Old World warblers (warbler), kinglets (kinglet), and allies, and (3) the finches (finch), icterids, tanagers (tanager), and allies. Some taxonomies regard the distinction between babblers (Timaliidae) and Old World warblers (Sylviidae) to be false, resulting in a redefinition and enlargement of the family Sylviidae. Other groups thought to be related were not, resulting in the further separation of the African warblers into the family Cisticolidae.

      In addition, some Old World flycatchers (flycatcher) (Terpsiphone, Monarcha) are related to other members of the corvoid assemblage (a group containing the crows, shrikes, vireos, kinglets, and others) rather than the muscicapine flycatchers. Instead, the muscicapine flycatchers and many of the thrushlike chats (chat), wheatears (wheatear), and other terrestrial species form a natural assemblage, the new Muscicapidae, a family that has undergone greater changes in its definition and composition than any other family of birds.

      The finch-billed oscines of the world have been subdivided into at least four major, probably unrelated groups—the Fringillidae, Emberizidae, Cardinalidae, and Estrildidae. A fifth group, the finchlike oscines of South America, appear to be seed-eating terrestrial tanagers that continue to expand the perimeter of that already impressive adaptive radiation. DNA studies suggest that the most familiar of the North American tanagers, such as the scarlet and western tanagers (tanager) (Piranga), are not tanagers after all. Instead, they appear to be more closely related to the grosbeaks (grosbeak) (Pheucticus, Cardinalidae) than to the tropical tanagers.

Mary Heimerdinger Clench Oliver L. Austin, Jr. Frank Gill

Additional Reading

Specialized works and technical review articles include Peter L. Ames, The Morphology of the Syrinx in Passerine Birds (1971); Charles G. Sibley, A Comparative Study of the Egg-White Proteins of Passerine Birds (1970); K.E.L. Simmons, “Anting and the Problem of Self-Stimulation,” Journal of Zoology, 149:145–162 (1966); and E.F. Potter, “Anting in Wild Birds: Its Frequency and Probable Purpose,” Auk, 87:692–713 (1970). Taxonomic works include Jean Delacour and Charles Vaurie, A Classification of the Oscines (Aves) (1957); and Ernst Mayr and J.C. Greenway, Jr., “Sequence of Passerine Families,” Breviora, vol. 58 (1956).Mary Heimerdinger Clench Oliver L. Austin, Jr. Ed.

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