▪ bird orderIntroductionany member of a cosmopolitan group of birds (bird) containing two very distinct families, the cuckoos (cuckoo) (Cuculidae) and the hoatzin (Opisthocomidae). Family Cuculidae is the much larger group, containing about 140 species of cuckoos, roadrunners (roadrunner), coucals (coucal), couas (coua), malkohas (malcoha), guiras (guira), and anis (ani); cuculids are found in the tropical and temperate zones of all the continents except Antarctica and on many oceanic islands. In contrast, family Opisthocomidae contains one species, found only in South America.The cuckoos are an ancient group with uncertain phylogenetic affiliations and no living near relatives. The cuckoos are of unusual biological, especially ethological, interest because many species are brood parasites—they lay their eggs in the nests of other species, which then rear the young cuckoos. Other cuckoos make their own nests, in which they incubate their eggs and rear their young as do most birds, and still others (Crotophaga and Guira) build communal nests. Some cuckoos are among the few birds that feed extensively on hairy caterpillars.This article provides a detailed overview of the Cuculidae. For more-detailed descriptions of the hoatzin, see galliform and hoatzin.General featuresThe cuckoos (cuckoo) range in size from the small glossy or emerald cuckoos of the genus Chrysococcyx, which are about 15 cm (6 inches) long, to the large ground cuckoos (Carpococcyx) and the larger species of coucals (Centropus), which reach nearly 90 cm (about 3 feet) in length after the often strikingly long tail is considered. Most cuckoos have fairly loose-webbed feathers, varying in colour from subdued browns, grays, olive, and black, to brilliant, iridescent greens and purples and bright yellow. The beak is of moderate length and often slightly downcurved.The cuckoos include the arboreal “typical” cuckoos of both the Old World and the New World, the terrestrial roadrunners (roadrunner) (Geococcyx) of southwestern United States and Mexico, and the more compact but also largely terrestrial coucals (coucal) (Centropus) of Africa and Australasia.DistributionAlthough the family Cuculidae is distributed virtually worldwide in temperate and tropical regions, most of the subfamilies are restricted to one hemisphere or the other. Three—Cuculinae, Centropodinae, and Couinae—are limited in distribution to the Old World; the subfamily Crotophaginae is wholly New World; the Neomorphinae is mostly New World, with one genus in Southeast Asia; and the sixth subfamily, Phaenicophaeinae, is represented in the tropics of both hemispheres. Many cuckoo genera are peculiar to certain parts of the world; others are cosmopolitan. Three genera—Rhamphomantis, Caliechthrus, and Microdynamis, for example—are known only from New Guinea; Dasylophus and Lepidogrammus occur only in the Philippine Islands; and Saurothera (lizard cuckoos) are found only in the West Indies. Some other genera are wide-ranging: the typical cuckoos of the genus Cuculus are found in Europe, Asia, Africa, and Australia; the small, brightly coloured Chrysococcyx species occur in Africa, Asia, Australia, and some Pacific islands; and the coucals, Centropus, live in Africa, Asia, Australia, and adjacent islands.Most cuckoos are solitary, often furtive birds that are inconspicuous even when relatively common. They do not form large flocks or even—except for the communal nesting anis (Crotophaga)—small parties.Natural historyHabitat utilizationAs a group, cuckoos (cuckoo) are forest birds (bird), often inhabiting dense thickets that may make them difficult to observe. Some species, such as many of the genus Cuculus, inhabit rather open woodland. The guira (Guira guira) of South America and many members of the Old World genera Clamator and Chrysococcyx are found in open savanna (grassland), but only where trees are present. The roadrunners live in open scrub and cactus desert, often in the absence of any large vegetation.Food habitsCuckoos are largely insectivorous, preying mostly on crawling insects. Orthoptera (grasshoppers, locusts, and mantids) are often taken. The greater, or North American, roadrunner (Geococcyx californianus) is reported to take flying grasshoppers by leaping after them from the ground. The larger cuckoos (such as the lizard cuckoos, roadrunners, and coucals (coucal)) take substantial numbers of lizards, snakes, and other small vertebrates, sometimes including birds. Among the Cuculidae, only the Madagascar couas (coua) (Coua) are reported to eat some fruit.VocalizationsCompared with other birds generally, cuckoos must be considered a highly vocal group. A variety of songs (birdsong), contact calls, and alarm notes are known for most species, some melodious and many harsh and discordant. The “song,” associated with territorial assertion and courtship, is usually characterized by the repetition of loud, short notes, often on a descending scale or with a downward break in the middle. The individual syllables have variously been described as whistling, piping, cooing, tooting, laughing, grating, and clicking, depending on the species. The familiar clear, two-note call of the common cuckoo (Cuculus canorus) of Europe, Asia, and Africa is uttered by the male alone, the female giving a low bubbling call; apparently in most other species of cuckoos as well, the song is given by the male alone.In addition to vocal sounds, at least one cuckoo, the North American, or greater, roadrunner, makes nonvocal sounds, clacking the mandibles to produce a rattling noise.Courtship behaviourExcept for features of parasitism, the behaviour of cuckoos is one of the least-studied aspects of their natural history. A few species, in several subfamilies, have been observed closely, and some generalizations are possible. In territorial advertisement and in courtship, the singing male (in both parasitic and nonparasitic cuckoos) may posture, spreading the tail feathers, which are white-tipped in most species, and drooping the wings (wing). Courtship feeding of the female by the male has been observed in several species.NestingTwo basic types of nesting behaviour are found among members of the order: parasitic and nonparasitic. About three-fifths of the Cuculidae mate, nest, and rear their young in the manner seen in most birds.About 50 species of cuckoos, including all of the subfamily Cuculinae (sometimes called “typical” cuckoos) and three species of the Neomorphinae, exhibit brood parasitism (that is, they lay their eggs in the nests of other species, which then rear the young cuckoos). Brood parasitism in the common cuckoo was recognized by Aristotle in the 4th century BCE. In addition, earlier evidence of brood parasitism in the koel (Eudynamys scolopacea) appears in the ancient Vedic literature of India, dating from about 2300 to 1800 BCE. Nonetheless, the scientific study of this form of parasitism remained largely neglected until the late 19th century, when intensive study by many observers revealed a number of remarkable behavioral and physiological adaptations in the cuckoo.In western Europe the common cuckoo is migratory, arriving at its breeding area in spring. The female establishes a territory and defends it against intrusion by other female cuckoos. The male is less restricted to an individual territory and more tolerant of other males; often more than one male will be in attendance on one female. Moving about within her territory, the female learns of potential host nests largely by watching their construction. Males of some species of cuckoos may accompany the females in searching for potential host nests. By the time she is ready to lay her eggs the female cuckoo has under surveillance several possible host nests and knows when the first eggs are laid by their owners. Most of the species parasitized by the common cuckoo are smaller than the cuckoo and lay their eggs at the rate of one each day until the clutch is complete, only then beginning incubation. Visiting when the hosts are away from the nest the cuckoo is able to place her egg in the nest after the host has started the clutch but before incubation has begun. It is well established that the common cuckoo lays eggs at 48-hour intervals, and apparently some glossy cuckoos lay one every 24 hours. The cuckoo places only one egg in each host nest and so must find a new nest for each of her 15 to 20 eggs. When parasitizing a species that builds a domed or globular nest, the female cuckoo clings to the outside of the nest and lays her egg through the opening without entering the nest. Cuckoo eggs have been found repeatedly in nests too small to admit a female cuckoo or too delicate to bear her weight. These findings have led some authorities to maintain that the cuckoo lays the egg on the ground and carries it to the nest in her beak, but extensive observations have failed to substantiate this supposition. Some of the glossy cuckoos, however, parasitize nests so small and frail that mandibular egg placement cannot be ruled out.In western Europe common cuckoos lay their eggs in the nests of many species, mostly songbirds (songbird) (Passeriformes (passeriform)); more than 140 host species have been reported, including some that could hardly rear a young cuckoo successfully. Careful study of this and several other cuckoos indicates, however, that despite the wide range of hosts recorded, a few host species bear most of the load of raising the young of each cuckoo species. In some instances, such as two of the little Australasian glossy cuckoos (Chrysococcyx), the parasite relies mainly on a single host species.Many birds react to a foreign object in the nest by deserting the nest, building another nest on top of the first, or removing the offending object. It is therefore of great advantage to a brood parasite to lay eggs that resemble those of its host, or, conversely, to parasitize species whose eggs are the same colour as its own. The utilization of several host species by a single population of cuckoos presents a difficult adaptive problem, for the host species may have many different egg colours. Early naturalists noted that there was often a marked resemblance between the egg of a cuckoo and those of the host, and a German ornithologist, Eduard Baldamus, in 1892 showed that the frequency and degree of similarity were too great to be coincidental. Subsequent studies by a number of workers, especially by the English naturalist, Edgar P. Chance, have revealed much of the basis for the resemblance, which is now called egg mimicry.Each female common cuckoo lays eggs of a particular shape and coloration throughout her life, but the eggs of different females vary widely. Most authorities now agree that a given female normally restricts her parasitism to the single host species by which she herself was reared, and that the common cuckoo population is composed of numerous clans. Each clan of parasitic cuckoos is called a gente, and each gente parasitizes only one species of bird for which the females of that gente have evolved egg mimicry. Only the gente of the female is important in egg coloration; the foster parentage of the male (that is, his gente) has no effect on his choice of females, and his genetic makeup apparently does not influence the coloration of eggs laid by his female offspring. Gentes are not given taxonomic designations but are sometimes known by their host species (“ redstart cuckoo,” “ dunnock cuckoo,” etc.). Evidence for this theory is largely circumstantial because of the difficulty in establishing the identity of individual cuckoos, but no plausible alternative theory has been proposed.The presence of cuckoo eggs in a wide variety of host nests in some small areas has been taken as evidence refuting the single-host hypothesis. Nevertheless, most of the nests with nonmatching eggs are found in areas in which the ecology has been seriously altered by human activity. As a result, many of female cuckoos residing in disturbed areas have suddenly found themselves without adequate numbers of the proper host species and have been forced to utilize hosts for which their own egg colours are inappropriate. In the forested portions of northern Europe, where the habitat is relatively undisturbed and consists of homogeneous microecological areas (such as reedbeds and birch forests), the common cuckoo parasitizes mainly a small thrush, the redstart (Phoenicurus phoenicurus), since both lay unspotted bluish eggs. In the reedbeds around Hungarian lakes, the usual host is the great reed warbler (Acrocephalus arundinaceus), and the cuckoo eggs are strongly blotched with gray and black, like those of the warbler.Studies of the reaction of the host to the presence of a cuckoo egg have indicated that egg mimicry is of considerable importance to the success of the parasite. Only about 5 percent of well-matched eggs are rejected by their hosts, compared with up to 72 percent of mismatched eggs. Few cuckoos have been studied intensively in terms of egg mimicry, but the phenomenon is known to occur in at least some species. The great spotted cuckoo has an egg pattern mimicking that of the magpie (Pica pica), its usual host in southern Europe. In Africa, where it is apparently a recent colonist, this cuckoo exhibits what has been called an “evolutionary escape from specialization.” Its new hosts, certain starlings (starling), nest in holes, in which the semidarkness renders the dissimilarity of eggs less visible and hence unimportant. Among the 12 species of emerald cuckoos, the degree of host specificity and egg resemblance is variable, some species being highly host-adapted, others less so. The genus as a whole shows no clear continuity of evolution in egg adaptation; individual species evidently have evolved their egg mimicry separately.A notable feature of the egg of the common cuckoo is its small size; its weight is only about 2.5 percent of that of the adult cuckoo. The eggs of other parasitic cuckoos weigh more than 6 percent of the adult weight, and those of birds that rear their own young vary from 8 to more than 20 percent of adult weight. The adaptive value of small eggs is not well understood, but it is assumed to be related to the reliance of the common cuckoo on small hosts. Experiments have shown, however, that incubating birds show no adverse reaction to eggs significantly larger than their own.Once the egg has been accepted and incubated by the hosts, the newly hatched cuckoo faces the problem of securing enough to eat from its small foster parents. The young cuckoo has evolved an astonishing form of behaviour, that of nest-mate eviction, that ensures that it will not have to compete with members of the foster brood for food. Within a few hours of hatching, the blind, naked, young cuckoo develops a strong urge to evict any objects, such as eggs or other nestlings, from the nest. It does this by working itself under the offending object and, aided by the presence of a depression between the shoulder blades, heaving the object over the rim of the nest. Within about 24 hours of hatching, the young cuckoo has the nest and the attentions of its foster parents to itself. The eviction habit is well developed in some parasitic cuckoos and seems to operate effectively in favour of the evictor but is surprisingly absent in many others, such as the crested cuckoos (Clamator). Another habit that, like nest-mate eviction, tends to reduce the competition faced by the young cuckoo is the removal of one or more of the host's eggs by the female cuckoo, sometime before or after or at the time of the laying of her own egg.The young cuckoo, while still in the nest, grows rapidly in response to the efforts of its foster parents, who regard it as their own. By the time it is ready to leave the nest, the young bird dwarfs its smaller hosts. Normally it does not receive attention from its real parents, but adult common cuckoos have occasionally been observed feeding young cuckoos; in the genus Chrysococcyx, such behaviour appears to be more frequent than in other genera.Nonparasitic cuckoosMost of the nonparasitic cuckoos form stable pair bonds and defend territories, within which they build their nests and rear their own young. The guira and the anis (ani) are exceptional in that they live in flocks of five to 20 individuals, each flock defending a territory within which its members feed and nest. Several birds of the flock may cooperate in building a nest, in which two or more females may lay eggs and share incubation. Many members of the flock participate in feeding the young.The nests of nonparasitic cuckoos are loose platforms of twigs placed in low vegetation or, rarely, on the ground. The coucals are unusual in that they build sizable domed nests of grass and twigs, on or near the ground, with side entrances.Like most insectivorous birds in temperate climates, temperate-zone cuckoos migrate toward or across the Equator for the winter. The migrations of some species are remarkable in that the young of the year travel completely independently of their parents and may cross up to 3,200 km (about 2,000 miles) of open ocean unguided. Adults and young of some of the small Australasian glossy cuckoos separately migrate long distances to relatively small oceanic islands. It is obvious that the means of orientation and navigation must be unusually precise, but what they are remains as yet unknown.Form and functionIn general body plan, most cuckoos (cuckoo) resemble the “perching birds” (order Passeriformes). They are slender-bodied, with medium to long wings (wing) and medium to extremely long tails, the latter sometimes constituting more than half the total length of the bird. The tail is usually graduated (the outer feathers (feather) shorter), and the individual feathers often are white-tipped.The foot structure of cuckoos is distinctive and at close range distinguishes these birds from any with which they might be confused. The foot of cuckoos is zygodactylous, or yoke-toed (that is, the outer toe is directed backward).The cuckoo bill, although usually slender, is occasionally quite stout, as in the ground cuckoos and coucals (coucal), in which it makes a formidable weapon for subduing prey. In the anis (ani) the bill is heavy, with a strong ridge along the culmen (top), and in the channel-billed cuckoo (Scythrops novaehollandiae) of Australasia it is extremely large, almost like that of toucans (toucan). In addition, a number of cuckoos are usually brightly coloured around the eye or between the eye and the bill. Prominent crests, a feature rare in cuckoos, are well developed only in the genus Clamator.Evolution and paleontologyThe order Cuculiformes is not well represented in the fossil record, but cuckoos (cuckoo) apparently existed by the early Paleogene Period, some 60 million years ago. The dearth of fossil evidence is not surprising in view of the group's preference for forest habitats; woodland animals, particularly small ones and particularly birds, are less frequently fossilized than those in open areas or aquatic habitats.Although not a large group, the modern species of cuckoos have highly diverse and specialized breeding behaviours. These behaviours range from extremes of social cooperation among pairs, such as the combination of eggs into a single nest in which incubation is shared, to the obligate brood parasitism of other species. The behaviour of brood parasitism has evolved multiple times in family Cuculidae. Some species—such as roadrunners (roadrunner), ground cuckoos and couas (coua)—are highly terrestrial.ClassificationOrder CuculiformesFamily Cuculidae (cuckoos (cuckoo), roadrunners (roadrunner), coucals (coucal), couas (coua), guiras (guira), malkohas (malcoha), and anis (ani))35 genera, about 140 species.Family Opisthocomidae ( hoatzin)Long neck and small head, with prominent crest of long feathers; beak short, stout, laterally flattened. Wings large and rounded; tail long and broad. Brown above, marked with white and russet; buff below. Nestling with 2 free, clawed digits on each wing. Adult size large, length about 61 cm (24 inches). 1 extant genus with 1 species; river systems of northern South America.Critical appraisalFormerly, the turacos (turaco) were a family (Musophagidae) within order Cuculiformes; however, most ornithologists treat them as a separate order (Musophagiformes) at present. The parrots (Psittaciformes (psittaciform)) are often considered on osteological and myological evidence to be the closest relatives of the cuculiforms, but even this relationship, if it exists at all, is not a close one. DNA studies have defined six distinct groups: the Old World parasitic cuckoos, the malkohas (malcoha) and couas, the coucals (coucal), the American cuckoos, the anis (ani), and the New World ground cuckoos. Some authorities recognize these groups, especially the coucals of the Old World tropics, as separate families.Herbert FriedmannAdditional ReadingEdgar P. Chance, The Cuckoo's Secret (1922), is an important and highly readable book about the common cuckoo. See also Ian Wyllie, The Cuckoo (1981). D.E. Davis, “Social Nesting Habits of the Smooth-Billed Ani,” Auk, 57:179–218 (1940), presents the results of extensive fieldwork on anis. Herbert Friedmann, The Parasitic Cuckoos of Africa (1948), Evolutionary Trends in the Avian Genus Clamator (1964), and The Evolutionary History of the Avian Genus Chrysococcyx (1968), are technical papers on Old World tropical cuckoos. Reginald E. Moreau, “A Contribution to the Biology of the Musophagiformes, the So-Called Plantain-Eaters,” Ibis, 2: 639–671 (1938), is one of the few detailed studies of musophagids.Herbert Friedmann Ed.
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