galliform

▪ order of birds

Introduction

   any of the gallinaceous (that is, fowl-like or chickenlike (chicken)) birds (bird). The order includes about 290 species, of which the best-known are the turkeys (turkey), chickens, quail, partridge, pheasant and peacock (Phasianidae); guinea fowl (Numididae); and grouse (Tetraonidae). Lesser-known members of the order are the megapodes (megapode) and the chachalacas (chachalaca), guans, and curassows (curassow). Although the hoatzin is here treated with the Galliformes, most taxonomists have assigned the hoatzin to the family Opisthocomidae (order Cuculiformes (cuculiform)).

General features

Size range and diversity of structure

  Most galliforms are medium-sized birds (bird), from the size of a pigeon to that of a domestic chicken, 40 to 60 cm (16 to 24 inches) long and 500 to 2,500 grams (1 to 5.5 pounds) in weight. The smallest members of the order are the sparrow-sized painted quail (quail) (Excalfactoria), about 13 cm (5 inches) long and about 45 grams (about 1.5 ounces) in weight. The heaviest galliform is the common, or wild, turkey (Meleagris gallopavo), wild specimens of which may weigh up to 11 kg (about 24 pounds); the longest is the argus pheasant (pheasant) (Argusianus argus), the male of which reaches 2 metres (79 inches) in breeding plumage, including wing and tail feathers, whose length exceeds one metre.

      The majority of gallinaceous birds are heavy-bodied, with short, rounded wings and strong, four-toed feet, adapted for life on the ground; a few, such as cracids and the hoatzin, live mainly in trees. The bill is short and slightly downcurved. The flight is fast but rarely sustained for long distances, most galliforms being sedentary, nonmigratory birds.

Distribution

 Some members of the order are found in virtually every habitat in subarctic, temperate, and tropical regions of the larger landmasses, and a few species (such as the ptarmigan, Lagopus) live within the Arctic Circle. The phasianids, the 178 species of which constitute by far the largest family, have nearly the distribution of the order. The cracids, with about 34 species, are restricted to tropical woodlands of Central and South America. The 18 species of grouse are found in northern temperate and Arctic regions of both hemispheres. The remaining groups are more restricted in distribution. The 10 species of megapodes (megapode) inhabit forests from the East Indies east to the Fiji Islands and south to central Australia. guinea fowl are restricted to Africa south of the Sahara but have been widely introduced on other continents. The two turkeys are native to North and Central America; the common, or wild, turkey in temperate woodlands of eastern United States and Mexico; and the ocellated turkey (Agriocharis ocellata) in Guatemala, British Honduras, and the Yucatán Peninsula of Mexico. The hoatzin (Opisthocomus hoazin) inhabits wooded river edges in northern and eastern South America.

Importance to humans

      Gallinaceous birds (bird) are unquestionably the most important avian group from the human standpoint. The chicken (Gallus domesticus) was domesticated in southern Asia at least 4,000 years ago from a parental stock of one or more species of jungle fowl (principally G. gallus). Selective breeding has produced well over 100 varieties, most of which are specialized for the production of either eggs or meat. In North America and to a lesser extent in northern Europe, the turkey is raised in numbers second only to those of the chicken. Domestic breeding of the turkey has been primarily for rapid growth and high market weight, the birds being raised solely for meat. Mature toms (males) may reach 23 kg (about 51 pounds). In many parts of the world, guinea fowl are an important barnyard species and are valued for the readiness with which they give alarm at the approach of a predator. (For a complete account of domestic gallinaceous birds, see poultry farming.)

 Galliform birds constitute the large majority of land game birds, in number both of species and of individuals. Populations in North America and in western Europe are often carefully managed through habitat manipulation, supplemental feeding, and artificial rearing to ensure maximum hunting yields. The common pheasant (Phasianus colchicus, called the ring-necked pheasant in the United States) was introduced in North America late in the 19th century and has become widely established. Several species of quail (especially those of the genera Coturnix and Colinus) and partridge (Perdix and Alectoris) provide much sport hunting. A widespread tendency in game bird management has been the introduction of species to areas where they have not been found previously.

      The North American wild turkey, once nearly exterminated by overhunting, has responded to careful management and is now taken in fair numbers in the hardwood forests of the eastern United States. Grouse (including ptarmigan) are hunted throughout their range.

Natural history

Habitat selection and food habits

      As an order, the galliforms inhabit a wide variety of vegetational types, including dense and open forest, open grasslands, scrub and second-growth forest, and flooded riparian (river) forests. Megapodes (megapode) live in dense jungle, some appearing in the open only to lay eggs on sandy beaches. The mallee fowl (Leipoa ocellata) is an exception, inhabiting the eucalyptus thickets that characterize the arid interior of Australia. The majority of galliforms roost on elevated perches at night, even those species that spend the daylight hours foraging on the ground. Virtually the only ones that live in treeless regions are certain of the grouse, such as the tundra-inhabiting ptarmigan, the prairie chickens (prairie chicken) (Tympanuchus cupido and T. pallidicinctus), the sage grouse (Centrocercus urophasianus), and the sharp-tailed grouse (Pedioecetes phasianellus). The Eurasian black grouse (Tetrao tetrix) occurs in open country and in forested regions. Most of the true pheasant, including the peafowl, are residents of open forest with clearings. guinea fowl of the genera Guttera and Agelastes and quail of the New World genus Odontophorus inhabit dense tropical forest. The hoatzin is closely associated with water and is almost never found far from it.

      The food of galliforms is varied, most species being basically vegetarian, but they also take large numbers of insects, worms, and other invertebrates. Many use their feet to uncover food in leaf litter. The hoatzin feeds on leaves and fruit, especially those of arums (Araceae), rarely descending to the ground but occasionally entering the water to take small crabs or fish.

courtship and mating

      Gallinaceous birds (bird) vary considerably in reproductive behaviour: some exhibit monogamous pair formation with a pair bond lasting through the breeding season, while others show varying types of polygamy, usually with members of both sexes being more or less promiscuous. Many, if not most, quail and partridge are monogamous, as are ptarmigan, guinea fowl, the hoatzin, some pheasant, and those megapodes and cracids that have been studied. Polygamy is known to occur in many grouse, including the North American grassland species, and in peafowl (Pavo) and some other phasianids. In the social displays of the grouse, a number of males assemble in a special assembly area, called a court, dancing ground, arena, or lek. The dancing ground lies outside of all nesting territories, and the same dancing ground is used year after year. Each of the males, which may number up to several dozen, has his own area within the chosen dancing ground. There he struts and postures, producing strange calls (some of which are produced in special esophageal air sacs) to attract the females, who visit the dancing ground to select a male and copulate. Dominant males occupy central positions in the arena group and may copulate with several females. Some species, such as the black grouse and the capercaillie (Tetrao urogallus), are variable in the degree to which males gather for display purposes, populations in more open areas tending more toward social displays. Males of the North American ruffed grouse (Bonasa umbellus) and those of some phasianids, the peacock pheasant (Polyplectron) and argus pheasant being examples, live in isolation, displaying to and copulating with any receptive female.

Nesting

      The large majority of gallinaceous birds nest on the ground, the nest being only a shallow scrape lined with soft grass or leaves. Cracids, the hoatzin, and the horned pheasant (Tragopan) build nests in trees; those of the cracids are relatively small for the size of the birds.

      The nesting of megapodes is unique among birds. The eggs of all species are placed in sand or soil, the heat for incubation coming from solar radiation, fermentation of plant matter, or even volcanic steam. Young megapodes, extremely precocious at hatching, dig their way to the surface and forage for themselves. They are fully feathered when hatched and can fly when 24 hours old. The simplest form of nesting, found in the maleo (Macrocephalon maleo), Wallace's megapode (Eulipoa wallacei), and some individuals of Freycinet's megapode (Megapodius freycinet, called jungle fowl in Australia), consists of placing each egg in a hole dug in sand to a depth of up to one metre (about one yard) in a site chosen for receiving the appropriate amount of solar radiation. Once laid and buried, the egg receives no further attention from the parents.

      Some members of the genus Megapodius (including jungle-dwelling members of M. freycinet) build mounds of decaying vegetation up to 10 metres (33 feet) long and 5 metres (16 feet) high. Mounds built by the brush turkeys (Alectura, Aepypodius, and Talegalla) are smaller, up to about 4 metres (13 feet) in diameter and 1 metre (3 feet) high. The mallee fowl, the most-studied of the megapodes, uses a combination of solar radiation and fermentation to maintain the incubation temperature.

      In the mound-building species of megapodes, the male maintains the mound for much of the year. Prior to and during the incubation period, he opens the mound once or twice a day to control the generation, absorption, and radiation of heat. It is generally believed that the bird measures the temperature of the egg chamber when opening the mound. The temperature-sensing organ is not known, but the most likely organ is the tongue. The mound is maintained within a degree or two of 33 °C (91 °F) throughout the period of several months that there are eggs in it. Observations of one species of brush turkey, Alectura lathami, indicate that the frequent opening of the mound may be as important for ventilation as for temperature control.

      The eggs of most gallinaceous birds are solidly coloured in white, buff, or olive, but those of species nesting in open areas are usually protectively coloured with blotches of brown or black. There is great variation in the number of eggs laid. Most members of the order are indeterminate layers, the female producing eggs until a certain number has accumulated in the nest. This characteristic has allowed man to exploit galliforms (especially the domestic hen, of course) for egg production. If the domestic hen were a determinate layer, as are members of many other bird orders, each individual would produce her clutch of about a dozen eggs and, regardless of whether or not the eggs were removed, would discontinue laying for at least two months.

      The clutch size varies from two to about two dozen, the largest number occurring in megapodes. Partridge, quail, and the smaller pheasant lay from 12 to more than 20 eggs, but the larger pheasant, such as the crested argus (Rheinardia ocellata) and the great argus, normally lay only two eggs. The hoatzin and the cracids lay two or three eggs, three being the rule in the smaller cracids, such as chachalacas. Guinea fowl lay seven to 20 eggs, and turkeys eight to 18.

      Incubation is usually performed by the female alone. Males of some New World quail, including the Montezuma quail (Cyrtonyx montezumae), the bobwhite (Colinus virginianus), and other members of the genus Colinus, incubate for short periods. The female of the southern European red partridge (Alectoris rufa) has been reported to lay two clutches, one incubated by the male, the other concurrently by herself. This behaviour, unique among birds, is in need of further investigation.

      It is uncertain whether or not the male hoatzin incubates the eggs. Both sexes have been observed in nest building and taking care of the young, so it is possible that the male also assists in incubation.

Care of the young

      Young gallinaceous birds (except those of the hoatzin) are extremely precocious, walking and feeding within a few hours of hatching. Parental behaviour parallels mating behaviour: males of species in which a pair bond is formed usually assist in shepherding the young. Although clad in a protective coat of down and usually camouflaged with spots and streaks, the chicks suffer high mortality from predators and adverse weather. Mortality rates of 50 percent or more are reported to occur in the period of a few months between hatching and independence of the young. Partridge, quail, and grouse maintain family groups (coveys) of a dozen or more birds that remain together until the next breeding season.

      The hoatzin, unusual in many ways, also differs from other galliforms in that its nestlings are hatched practically naked and are fed by the parents in the nest. The young are fed regurgitated food from the crops of their parents. Although they have a long fledging period, young hoatzins scramble about in the branches around the nest, holding on with the clawed first and second digits of the wings (wing) and, in the manner of parrots, with their bills. The fact that the second digit of the wing (now considered to be digit III in the evolutionary sense) is free and bears a functional claw was once thought (erroneously) to indicate affinities with the reptilian forebears of birds. Most authorities now believe that the free digits are a secondary adaptation that allows the young hoatzin great mobility during its flightless period and allows it to climb out of the water, into which it may deliberately drop when danger threatens.

      Most gallinaceous birds reach sexual maturity at the age of at least one year. Some species, however, may be physiologically capable of reproduction at a much earlier age. The common quail (Coturnix coturnix), wild individuals of which normally breed at one year of age, matures to breeding condition in seven weeks in captivity. It is uncertain whether wild birds hatched in the spring actually do breed during the summer; environmental control factors, especially decreasing day length, probably prevent the attainment of breeding condition by two-month-old birds in natural situations. Even the Congo peacock (Afropavo congensis), a relatively large species, is able to reproduce at one year of age when reared in captivity.

Vocalizations

      Like many other birds that inhabit dense cover, forest galliformes are endowed with strong voices, ranging from musical whistles to harsh screams. The repertoire of most species includes alarm notes, food calls, “crowing” by males to advertise territories, and notes used to maintain social groups. Many species are especially vocal at daybreak; a few are reported to call at night. Megapodes give clucking and cackling calls during the day and mewing calls at night. Cracids produce loud calls that have immense carrying power. Males of some species have elongated, looped tracheae (windpipes), which are believed to add to the carrying power of the calls. Members of the social groups often cackle noisily together. Male peafowl utter a long mournful scream that sounds quite like a child in distress.

Form and function

 With the exception of the hoatzin, all galliforms have the same general body plan, being adapted for a primarily terrestrial existence. The feet (foot) and claws are large in all families, particularly so in the megapodes, reflecting their use for scratching and digging. The hind toe is larger and more functional in groups, such as the cracids, that spend much time in trees; it is smaller in the more terrestrial groups, but in none has it been lost, as it has in terrestrial birds of some other orders.

      The short, rounded wings (wing), powered by strong breast muscles (the white meat of the chicken), are indicative of the need for short, rapid bursts of flight, such as the escape from predators. Although no galliform is flightless, none is capable of long flights. The tail varies from extremely short (as in the painted quail) to strikingly long; in many male pheasant the tail may be more than two-thirds of the bird's total length. The tails of some pheasant and of most megapodes (megapode) are vaulted, having an inverted V-shape in cross section.

      Male ornamental plumage is often remarkable in shape and coloration, combining spots or bars of silver, green, or purple iridescence with areas of brilliant orange, yellow, or white. Sculptured, fleshy wattles on the faces of male pheasant and grouse are often coloured a bright blue or red. In some male pheasant, such as the firebacks (Lophura), impeyans (Lophophorus), and peafowl, the head is ornamented with a small tuft of modified plumes, forming a tiny fan. Many cracids have patches of bare skin on the face or throat, usually red, yellow, or blue. Males of many curassows (curassow) and guans possess head ornaments in the form of a brightly coloured fleshy knob or a bony casque (helmet) on the top of the head.

Evolution and paleontology

      Galliforms represent one of the oldest of all lineages of modern birds (bird), with roots in the Cretaceous Period. Megapodiidae, a family of specialized mound builders from Australasia, is the most primitive of the extant families. The second oldest family, Cracidae, originated in the early Paleogene. The first cracids were most likely from Central America and southern North America, which was tropical in climate at that time. It is also conceivable that the evolution of the major groups of cracids and the divergence of New World quail (Odontophoridae) from other galliform birds over 35 million years ago was influenced by the breakup of Gondwana.

Classification

Distinguishing taxonomic features

      The limits and interrelationships of galliform families have been determined on the basis of general body proportions, muscle and bone configurations, plumage, clutch size and egg characteristics, the appearance of the young, and some aspects of behaviour. Early 21st-century studies have utilized the biochemistry of DNA to indicate relationships.

Annotated classification

Order Galliformes

 

      Family Megapodiidae (megapodes (megapode) or mound builders)

 All use heat other than body heat to incubate their eggs. Large feet, small heads. Tail often vaulted. Medium to large birds, 25–65 cm (10–25 inches); sexes alike. 1 fossil, Chosornis praeteritus, from the upper Pleistocene; 7 extant genera, 12 species. Australia and East Indies to central Polynesia.

      Family Cracidae (chachalacas, guans, and curassows (curassow))

 Tail moderately long and broad. Plumage black or brown, duller in female. Most species with bare skin between eyes and beak (lores), some with fleshy wattles or other ornaments on face or crown. Medium to large; length 52–99 cm (20–39 inches). Lower Eocene to present; 11 fossil and 11 extant genera, 34 species. New World tropics and subtropics, from southern Texas to Paraguay.

      Family Tetraonidae ( grouse)

 Distinguished by having lower leg (tarsus) and sometimes feet at least partially feathered; nostrils also feathered. Tail strong; ornamental in some species. Many with brightly coloured bare skin over eye. Medium to large; 30–90 cm (12–36 inches). Lower Miocene to present; 3 fossil and 9 extant genera, 17 species. North America and northern Eurasia.

      Family Phasianidae (pheasants (pheasant), quail, chickens (chicken), partridges (partridge), turkeys (turkey), and relatives)

 Nostrils, feet, and (usually) tarsus unfeathered; esophageal air sacs lacking. Many species with spurs on the back of the tarsus. Plumage, especially of males, bright in many species, often with ornamental feathers; sexes usually different. Small to large; length 13–200 cm (5–80 inches). About 50 genera, approximately 200 extant species; virtually worldwide, except southern third of South America, northern Eurasia, and some oceanic islands.

      Family Numididae ( guinea fowl)

 Distinguished by presence of small whitish spots on dark bluish or slate body feathers; head and neck bare or slightly feathered, often brightly coloured. Legs and feet large; spurs present only in Phasidus and Agelastes. Medium to large; length 43–75 cm (17–30 inches). No fossil species; 5 genera, 7 species. Africa south of the Sahara and Madagascar.

Critical appraisal

      In most taxonomic systems, domestic chickens (chicken), turkeys (turkey), quail, and pheasant have been placed in the family Phasianidae. In some classifications, turkeys and New World quail are placed into their own families, Meleagrididae and Odontophoridae, respectively. Of the phasianids, the New World quail are genetically the most divergent natural group, whereas the turkeys are simply large pheasants.

      Descriptions of the enigmatic hoatzin are retained in this article only for convenience. It has been allied in some taxonomic systems to turacos (turaco) and cuckoos (cuckoo); however, early 21st-century DNA studies clearly refute the proposed cuckoo relationship, though they do not offer a strong alternative. Some authorities place the hoatzin in the family Opisthocomidae in the order Cuculiformes.

François Haverschmidt

Additional Reading

H.J. Frith, The Mallee-Fowl: The Bird That Builds An Incubator (1962), is an extensive study of the breeding habits of the megapode Leipoa ocellata. Alexander F. Skutch, “Habits of the Chestnut-winged Chachalaca,” Wilson Bulletin, 75:262–269 (1963), presents information on the breeding habits of Ortalis garrula. Charles Vaurie, “Taxonomy of the Cracidae (Aves),” Bulletin of the American Museum of Natural History, 138:133–259 (1968); and François Vuilleumier, “Relationships and Evolution Within the Cracidae (Aves, Galliformes),” Bulletin of the Museum of Comparative Zoology, Harvard University, 134:1–27 (1965), are two analyses of the taxonomic relationships within this family. See also Jean Delacour and Dean Amadon, Curassows and Related Birds (1973). Paul A. Johnsgard, The Grouse of the World (1983), includes a lengthy bibliography. Siegfried Klaus et al., Die Auerhühner, new ed. (1986), is a short monograph on the capercaille, and Die Birkhühner, new ed. (1990), is a study of the black grouse. George L. Girard, Life History, Habits, and Food of the Sage Grouse, Centrocercus urophasianus Bonaparte (1937), a brief work, provides insight into the natural history of this grassland species. Gardiner Bump et al., The Ruffed Grouse: Life History, Propagation, Management (1947, reprinted 1978), is a classic study of this woodland grouse. William Beebe, A Monograph of the Pheasants, 4 vol. (1918–22, reissued 4 vol. in 2, 1990), is a classic work on pheasants, magnificently illustrated with colour plates; revised, less technical editions were published with the title Pheasants: Their Lives and Homes, 2 vol. (1926), and 1 vol. (1936). Jean Delacour, The Pheasants of the World, 2nd ed. (1977), is a fully illustrated treatment of the group, leaning heavily toward aviculture. Philip Wayre, A Guide to the Pheasants of the World (1969), is an extensively illustrated work with emphasis on conservation and on breeding in captivity. Johnsgard's The Pheasants of the World (1986) and The Quails, Partridges, and Francolins of the World (1988) are companion volumes. Herbert L. Stoddard, The Bobwhite Quail: Its Habits, Preservation and Increase (1931, reprinted 1978), is an important book on this game species. Henry S. Mosby and Charles O. Handley, The Wild Turkey in Virginia (1943), is a broad study of the natural history of Meleagris gallopavo. A.W. Schorger, The Wild Turkey: Its History and Domestication (1966), is primarily a historical treatise. William Beebe, “A Contribution to the Ecology of the Adult Hoatzin,” Zoologica, 1:45–66 (1909), and Tropical Wildlife in British Guiana (1917), pp. 155–182, present information from the author's field observations of the hoatzin.François Haverschmidt Ed.

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