Galliformes

(vertebrate zoology) An order of birds that includes important domestic and game birds, such as turkeys, pheasants, and quails.

Family: Moundbuilders
Family: Curassows, Guans, and Chachalacas
Family: Guineafowl
Family: Fowls and Pheasants
Family: New World Quails

(Chicken-like birds)

Class: Aves

Order: Galliformes

Number of families: 5

Number of genera, species: 77 genera; 281 species

Evolution and systematics

Few birds have such a long relationship with people as the Galliformes, but their own history is even older. Fossils show that their predecessors date back to the Eocene period (50 to 60 million years ago), when northern latitudes were tropical. The earliest known cracid ancestor was found in the United States in Wyoming, although the megapodes are probably more primitive. All the Galliformes are of a similar, standard design, perfected for a terrestrial lifestyle that has been little modified by millions of years of evolution.

There are two tribes: the Craci (the megapodes, chachalacas, guans, and curassows) and the Phasiani (the turkeys, grouse, New World quails, pheasants, partridges, and guinea fowls). The two are distant in evolutionary terms, and no examples are known of a bird from one tribe hybridizing with one from the other. They are distinguished by the hallux, the hind toe, which in the Craci is in line with the other toes, but in the Phasiani is above the others.

The Phasianidae is numerically dominant, accounting for 155 species. Work on the mitochondrial DNA of birds in the late twentieth century has resulted in the New World quails (Odontophoridae) being split from the pheasants (Phasianidae). Discoveries continue to change our understanding. For example, the Udzungwa forest-partridge (Xenoperdix udzungwensis), discovered in southern Tanzania in 1991, is more closely related to the Asian hill-partridge (Arborophila torqueola).

Physical characteristics

Several characteristics are common to the Galliformes, all of which can be seen in the domestic chicken, derived from the red junglefowl (Gallus gallus) of Southeast Asia. Most gallinaceous species are medium to large in size, with a stocky body, small head, and short wings. The Old World quails are the smallest, the most diminutive being the Asian blue (Coturnix chinensis) at just 5–6 in (12–15 cm) and weighing less than 1 oz (20 g). By contrast, the wild turkey (Meleagris gallopavo) weighs 17–22 lb (8–10 kg); only the domesticated forms destined for the table can attain 44 lb (20 kg); while a large, male green peafowl (Pavo muticus) is up to 98 in (250 cm) long, although its immense tail accounts for more than half of this. Pheasants in particular show a significant size difference between males and females, with the tail often responsible for one-third of the total.

In many cases, males and females are mottled brown or black, adapted to camouflage in the forest or scrub. In a few species, however, males are colorful, with iridescent colors that have long made them attractive to humans. The male Indian peafowl (Pavo cristatus), the "peacock" of art and the movies, whose fanned tail has hundreds of "eyes" on the tips, is perhaps the most well known.

Feeding ecology and diet

Galliformes are terrestrial, spending their day foraging for food in grasslands or the understory of the forests. Birds have a short, often downcurved bill, used to peck plant material from the ground or from short vegetation, though several species in northern latitudes, such as the capercaillie (Tetrao urogallus), depend on the stiff needles of coniferous trees (Pinus) to see them through the long winter when snow covers the ground. They also have large, strong feet, a crucial attribute that allows them to expose seeds and roots that are inaccessible to most other animals (the name "megapode" is derived from the Greek words for "big foot"). These feet are capable of moving heavy branches or stones; the orange-footed scrubfowl (Megapodius

reinwardt) can move a stone up to eight times its own weight. Their heavy build indicates a diet based on bulky, vegetable matter, although the chicks of many species depend on insects and larvae during their first few weeks.

Gallinaceous birds have a roomy, flexible crop, which can be extended to cache food before beginning to digest it. They also have a very strong gizzard, used to grind down the hard exterior of seeds and nuts, and the tough fibers in green vegetation. To aid digestion, birds regularly swallow small stones. Even the most secretive forest Galliformes visit roads and tracks early in the morning in search of grit before their day's feeding. Some make only occasional visits to water, even during dry periods, but a few species visit salt licks, where they ingest claylike soil to supplement their diet with minerals.

Reproductive biology

Galliformes display a wide variety of breeding strategies. In general, species with the least sexual dimorphism in size and color are monogamous, and those in which the male has more resplendent plumage are polygynous. In many grouse species, males display at communal leks, seeking to be the dominant male to attract a harem of females.

Male Galliformes have a range of adornments to attract females: bright colors, crests, unusually shaped tail feathers, or markings. Some have additional modifications, such as long, pendant wattles, dewlaps, combs, or "eyebrows." Most species display one of these "badges," or white patches on the wings or tail, although the curassows are the most highly evolved family, with colorful knobs, or ramphothecae, on their bill, which grow larger as the bird ages.

Unlike many nonpasseriform birds, calls play an important part in display and territorial ownership, and also simply to keep in touch with a mate. This is not surprising, since many species are solitary, living deep in scrub or forest, and crepuscular, being most active at dawn and dusk. In tropical areas, the wailing calls of tinamous and guans travel across the forest in the fading daylight, up to 4 mi (6.4 km) in some species. This is possible because a modification in the length of the trachea and a loop between the skin and the pectoral muscles enables some cracids to produce calls at a lower pitch and a higher volume than most other species, although swans (Cygnus) and cranes (Grus) have a similar modification.

The breeding strategy of the megapodes, which do not use body heat to incubate their eggs, is unique in the avian world, although it does not demonstrate a link to reptiles, as some have suggested. The male builds a huge mound of sand or plant material or constructs a burrow, invites a female to lay her eggs, then tends to the nest, regulating the temperature for many weeks until the young hatch. Indeed, some male megapodes are attached to their mound for 11 months of the year. In monogamous species, however, both birds help to rear the young, maintaining the pair bond through mutual preening or activities such as wing-drumming.

In most species, the young are precocious, able to feed semi-independently within a few hours of hatching. Generally, the first downy feathers are subdued in color to reduce the risk of a predator seeing them.

Distribution and habitat

Gallinaceous birds are found in a wide variety of habitats, in semideserts, steppes, savannas, forests, mountains, and farmland. The cracids are the most arboreal family, with most species spending at least part of the time in the forest canopy, but even some of the chachalacas feed in more open habitats. Members of other families are more specialized; for example, the British race of the red grouse (Lagopus lagopus scoticus) lives only on upland Calluna heather moorlands, devoid of any trees, whereas the other 18 races live around dwarf trees!

The Galliformes occur on every continent except Antarctica, with some families found on a single continent—megapodes in Australasia, cracids in Central and South America, turkeys in North America, New World quails in North and South America, and guinea fowl in sub-Saharan Africa. Only two families are spread across more than one continent—grouse in North America and Eurasia, and pheasants and partridges in Africa, Eurasia, and Australasia. There is relatively little geographic overlap between the families, perhaps not surprising given their sedentary nature; most birds moving only a few miles from where they hatched.

Behavior

The social behavior of the Galliformes is complex, and the commoner species have been the subject of many studies by ornithological and hunting interests. Many species are solitary or spend the year in pairs; the males being strongly territorial, charging intruders with their neck raised and wings spread open. In some grouse species, this has developed into a mating display, in which males demonstrate their defensive prowess to females gathered nearby, a behavior known as "lekking." Outside the breeding season, some species, such as chachalacas, brush-turkeys, and pheasants, feed communally where there is a good supply. Some will also roost communally, flying to the tree canopy where they are safer from predatory ground mammals.

Many Galliformes have cryptic plumages. Birds sit tight in thick vegetation, hoping not to be noticed, and only when the threat is almost upon a bird will it move. A few species, such as the Nicobar scrubfowl (Megapodius nicobariensis), evade predation by running swiftly away, but most species explode into the air in a rush of wings. This is possible because Galliformes have strong breast muscles and strong legs, enabling a near vertical take-off. In flight, many are bulletlike, especially the partridges, their wings beating rapidly, although only over short distances. Birds fly close to the ground, although this brings its own problems. Many capercaillies and black grouse (Tetrao tetrix) are killed against high deer fences around European forestry plantations.

Most gallinaceous birds bathe, often visiting the same sites repeatedly, squatting in a shallow pit and beating their wings to shower sand or dust across their plumage to maintain the feathers and remove parasites. In tropical species, bathing and preening usually takes place during the middle of the day when birds are resting, whereas birds are more active, displaying and feeding, during the three hours around sunrise and sunset.

Most species are sedentary, but a few are altitudinal migrants, moving down the mountainsides outside the breeding season; four Old World quails (Coturnix) are true long-distance migrants, traveling from breeding grounds in Eurasia to sub-Saharan Africa.

Conservation status

Of the 281 species, 104 are Threatened or Near Threatened, far above the average of 10% for all bird species. The pheasants and partridges are under the greatest threat, with 71 listed by the IUCN as Critically Endangered, Endangered, Vulnerable, or Near Threatened; one, possibly two, pheasant species have become extinct since 1600.

Gaps in our knowledge constrain the development of conservation measures for many of the species in the remotest habitats. There are many species, such as the Bruijn's brushturkey (Aepypodius bruijnii), for which further research is crucial to their conservation.

Hunting of adults or the collection of eggs for food remains a problem for several of these "game" birds. Historically, cracids were an important sustainable source of protein for native Amerindians, but rapid human colonization since 1492 has led to over-exploitation of birds, as well as the destruction of their tropical rainforest habitat. Some species are now on the brink of extinction, with the Alagoas curassow (Mitu mitu) only known in captivity since the 1980s, the horned guan (Oreophasis derbianus) limited to a few isolated mountain ranges in Mexico and Guatemala, and the endemic Trinidad piping-guan (Pipile pipile) now restricted to a few square miles of montane forest on a single island. It is doubtful whether the harvesting of eggs from some species is sustainable, and over-exploitation has probably caused the extinction of several megapode species on Pacific Islands. Such gathering continues today, with, for example, an estimated five million eggs taken every year from a single site in New Britain where Melanesian scrubfowl (Megapodius eremita) gather to breed.

However, habitat destruction is the principal threat. Galliformes that depend on primary, tropical forests are under the greatest threat, with logging of timber or intensive burning to clear the land for agriculture being major problems, especially in Southeast Asia. As if the removal of the old forest was not enough, herbivorous livestock, such as cattle, sheep, and goats, compete with Galliformes for seeds and vegetation. The Galliformes' dependence on certain habitats makes them good indicators of environmental change. As consumers of a large biomass of seeds and roots, they also play a critical role in dispersing seeds, especially in tropical forests.

Our fascination with these amazing birds provides a potential, sustainable solution to the need of local people to earn an income. In parts of South America, for example, ecotourism can be more important to the local economy than logging or beef production, and is certainly better for the Galliformes that birdwatchers come to see.

Significance to humans

We can only guess when Homo sapiens first discovered that some Galliformes were relatively easy to catch and that their meat tasted good and was high in protein. Some time later, as people moved from being hunter-gatherers to farmers, they learned to domesticate several species, including turkeys, chickens, and guinea fowl, which remain part of human diet across the world. Galliform eggs are much sought after, with a high yolk content that provides a rich source of protein. The word "fowl," which is applied generically to game and domesticated birds, has its origins in the Old English fugol, the Old Norse fogl, and the modern German vogel.

During the last 200 years, many gallinaceous species have been moved between countries and continents for decoration or for shooting on vast, private estates. Introductions, whether deliberate or accidental, are the third most serious threat to global avifauna, after habitat destruction and degradation. Introductions are most problematic on islands, and although Galliformes are not known to be an special threat, they have been widely transferred. At least 45 of the 281 species have been introduced to two archipelagoes, Hawaii and New Zealand, though only two-thirds colonized successfully, the rest failing either because of poor stock or an inability to deal with predators that they were not used to facing.

The sedentary nature of most Galliformes makes them popular for shooting. Even in countries where strict legislation makes it illegal to shoot birds, many Galliformes are excluded. At the height of the British Empire, aristocrats and civil servants spent their leisure time in Africa and Asia shooting small game (Galliformes), and brought some of the most numerous back to Europe, most notably the ring-necked pheasant (Phasianus colchicus). Each year, tens of millions are reared and released for shooting. These and European game species, such as the gray partridge (Perdix perdix) and rock partridge (Alectoris graeca), were subsequently introduced into North America for sport.

Resources

BirdLife International. Threatened Birds of the World. Cambridge: BirdLife International, 2000.

Brooks, D. M., and S. D. Strahl, eds. Curassows, Guans and Chachalacas: Status Survey and Conservation Action Plan for Cracids 2000–2004. Cambridge: International Union for Conservation of Nature and Natural Resources (IUCN), 2000. del Hoyo, J., A. Elliott, and J. Sargatal, eds. Handbook of the Birds of the World. Vol. 2, New World Vultures to Guineafowl. Barcelona: Lynx Edicions, 1994.

Hudson, P. J., and M. R. W. Rands. Ecology and Management of Gamebirds. Oxford: BSP Professional Books, 1988.

Johnsgard, P. A. The Grouse of the World. Lincoln: University of Nebraska, 1983.

Sibley, C. G., and B. L. Monroe, Jr. Distribution and Taxonomy of Birds of the World. New Haven: Yale University Press, 1990.

Snow, D. W., and C. M. Perrins. Birds of the Western Palearctic: Concise Edition. Vol. 1, Non-Passerines. Oxford: Oxford University Press, 1998.

Storche, I., ed. Grouse: Status Survey and Conservation Action Plan 2000–2004. Cambridge: IUCN and the World Pheasant Association, 2000.

Urban, E. K., H. C. Fry, and S. Keith, eds. Birds of Africa Vol. 2, Gamebirds to Pigeons. London: Academic Press, 1986.

Buchholz, R. "Older Males Have Bigger Knobs: Correlates of Ornamentation in Two Species of Curassow." Auk 108(1991): 153–160.

Vaurie, C. "Taxonomy of the Cracidae." Bulletin of the American Museum of Natural History 138 (1968): 135–243.

Webre, A., and J. Webre. "Ecotourism and the Plain Chachalaca Ortalis vetula in Texas." Bulletin of the IUCN/Birdlife/WPA Cracid Specialist Group. 6 (1998): 13–14.

BirdLife International. Wellbrook Court, Girton Road, Cambridge, Cambridgeshire CB3 0NA United Kingdom. Phone: +44 1 223 277 318. Fax: +44-1-223-277-200. E-mail: birdlife@birdlife.org.uk Web site:

IUCN Species Survival Commission. Rue Mauverney 28, Gland, 1196 Switzerland. Phone: +41 22 999 01 53. Fax: +41 22 999 00 15. E-mail: lwh@hq.iucn.org Web site:

World Pheasant Association. P. O. Box 5, Lower Basildon St, Reading, RG8 9PF United Kingdom. Phone: +44 1 189 845140. Fax: +44 118 984 3369. E-mail: office@pheasant.org.uk Web site:

[Article by: Julian Hughes]

A large order of birds containing the gallinaceous or chickenlike birds. They are found worldwide, although the several subgroups have a more limited distribution. Part of the superorder Neognathae, the galliforms are closely related to the anseriforms and include the most important domesticated birds, the chicken (Gallus gallus) and the turkey (Meleagris gallopavo). See also Anseriformes.

The order Galliformes is divided into two suborders with two families, as follows.

Order Galliformes

     Suborder Cracoidea

          Family: Megapodiidae (megapodes or brush turkeys; 7 species)

                    Cracidae (curassows, guans, chachalacas; 49 species)

     Suborder Phasianoidea

          Family: Phasianidae (grouse, pheasant, quail, guinea fowl, turkey; 212 species)

               Subfamily: Tetraonidae (grouse)

                    Phasianiaen (pheasant, peafowl, chicken)

                    Odontophorinae (New World quail)

                    Numidinae (guinea fowl)

                    Meleagrinae (turkey)

The galliforms have been one of the most economically important group of birds to humans. Almost all species have been or are still being hunted for food. Many species of galliforms have become reduced in numbers because of overhunting and habitat destruction. Some species, especially the forest pheasants of Asia, are seriously endangered. See also Aves.

The noun has one meaning:

Meaning #1: pheasants; turkeys; grouse; partridges; quails; chickens; brush turkeys; curassows; hoatzins
  Synonym: order Galliformes

Galliformes
Male Grey Junglefowl, Gallus sonneratii
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Aves
Subclass:	Neornithes
Infraclass:	Galloanserae
Order:	Galliformes Temminck, 1820
Families
Megapodiidae Cracidae Odontophoridae Numididae Phasianidae and see text
Synonyms
Craciformes Gallomorphae

Galliformes are an order of birds containing turkeys, grouse, chickens, quails, and pheasants. Common names are gamefowl or gamebirds, landfowl, gallinaceous birds or galliforms. "Wildfowl" or just "fowl" are also often used for Galliformes, but usually these terms also refer to waterfowl (Anseriformes), and occasionally to other commonly-hunted birds.

This group has more than 250 living species, one or more of which are found in essentially every part of the world's continents (except for the innermost deserts and perpetual ice). They are more rare on islands, and in contrast to the closely-related waterfowl are essentially absent from oceanic islands – except if introduced there by humans. Several species have been domesticated during their long and extensive relationship with Homo sapiens.

Contents 1 Description 2 Ecology 2.1 Food and feeding 2.2 Reproduction 3 Systematics and evolution 3.1 Evolution 3.2 List of major taxa 4 Footnotes 5 References 6 External links

Description

Female (left) and male Common Pheasant. Sexual dimorphism is conspicuous in this species, one of the most apomorphic gamefowl.

These birds vary in size from the diminutive Asian Blue Quail (Coturnix chinensis) at 12.5 centimetres (5 in) long and weighing 28 to 40 grams (1 to 1.4 oz) to the largest extant galliform species, the North American Wild Turkey (Meleagris gallopavo), which may weigh as much as 14 kilograms (about 30.5 lb) and may exceed 120 centimetres (47 in).

The galliform bird species with the largest wing-span and largest overall length (including a train of over 6 feet) is most likely the Green Peafowl (Pavo muticus). Most galliform genera are plump-bodied with thick necks and moderately long legs, and have rounded and rather short wings. Grouse, pheasants, francolins, and partridges are typical in their outwardly corpulent silhouettes. While most galliforms are rather weak-flying, flightless forms are unknown among the living members of the order. The Sylviornis, a huge prehistorically extinct mound-builder relative of New Caledonia, was flightless, but as opposed to most other flightless birds like ratites or island rails which become flightless due to arrested development of their flight apparatus and subsequently evolve to larger size, the Sylviornis seems to have become flightless simply due to its bulk, with the wing reduction following a consequence, not the reason for its flightlessness. There are a number of prehistorically extinct mound-builders from Pacific islands, and these seem to have arrived at flightlessness in the more conventional way.

Adult males of many galliform birds have one to several sharp horny spurs on the back of each leg, which they use for fighting. In several lineages, there is pronounced sexual dimorphism, and among each galliform clade, the more apomorphic ("advanced") lineages tend to be more sexually dimorphic.

Ecology

Most of the galliform birds are more or less resident, but some of the smaller temperate species (such as quail) do migrate over considerable distances. Altitudinal migration is evidently quite common amongst montane species and a few species of subtropical and subarctic regions must reach their watering and/or foraging areas through sustained flight. Species known to make extensive flights include the ptarmigans, sage-grouse (Centrocercus), Crested Wood-partridge, Green Peafowl, Crested Argus, Mountain Peacock-pheasant (Polyplectron inopinatum), Koklass (Pucrasia macrolopha), Reeves's Pheasant (Syrmaticus reevesii) and Green Junglefowl (Gallus varius)^{[citation needed]}. Other species—for example, most of the New World quails (also known as the toothed quails), the enigmatic Stone Partridge (Ptilopachus petrosus) of Africa, guineafowl and eared-pheasants (Crossoptilon)—are all notable for their daily excursions on foot which may take them many miles in a given day.

Some Galliformes are adapted to grassland habitat, and these genera are remarkable for their long, thin necks, long legs and large, wide wings. Fairly unrelated species like the Crested Fireback (Lophura ignita), Vulturine Guineafowl (Acryllium vulturinum) and Malleefowl (Leipoa ocellata) are outwardly similar in their body types (see also convergent evolution).

Most species that show only limited sexual dimorphism are notable for the great amount of locomotion required to find food throughout the majority of the year. Those species that are highly sedentary but with marked ecological transformations over seasons exhibit marked distinct differences between the sexes in size and/or appearance. Eared-pheasants, guineafowls, toothed quails and the Snow Partridge (Lerwa lerwa) are examples of limited sexual differences and requirements for traveling over wide terrain to forage.

The Bronze-tailed Peacock-pheasant (Polyplectron chalcurum), Snow Partridge, Painted Spurfowl (Galloperdix lunulata) and the Crimson-headed Partridge (Haematortyx sanguiniceps) are notable in their habit of moving around as pairs not only on foot but also in the air.^{[citation needed]}

Food and feeding

A flock of adult and young Helmeted Guineafowl foraging

Herbivorous to slightly omnivorous galliforms, comprising the majority of the group, are typically stoutly built and have short thick bills primarily adapted for foraging on the ground for rootlets or the consumption of other plant material such as heather shoots. The young birds will also take insects.

Peafowl, junglefowl and most of the subtropical pheasant genera have very different nutritional requirements from typical Palearctic genera. The Himalayan Monal (Lophophorus impejanus) has been observed digging in the rotting wood of deadfall in a similar manner to woodpeckers to extract invertebrates, even bracing itself with aid of its squared tail. The Cheer Pheasant (Catreus wallichi), Crested Argus (Rheinardia ocellata), the Crested Wood-partridge (Rollulus roulroul) and the Crested Guineafowl (Guttera pucherani) are similar ecologically to the Himalayan Monal in that they too forage in rotting wood for termites, ant and beetle larvae, molluscs, crustaceans and young rodents.

Typical peafowl (Pavo), most of the peacock-pheasants (Polyplectron), the Bulwer's Pheasant (Lophura bulweri), the ruffed pheasants (Chrysolophus) and the hill partridges (Arborophila) have narrow, relatively delicate bills, poorly suited for digging. These Galliform genera prefer instead to capture live invertebrates in leaf litter, in sand and in shallow pools or along stream banks. These genera are also outwardly similar in that they each have exceptionally long, delicate legs and toes and the tendency to frequent seasonally wet habitats to forage, especially during chick-rearing. The Blue Peafowl (Pavo cristatus) is famed in its native India for its appetite for snakes – even poisonous cobras – which it dispatches with its strong feet and sharp bill. The Lady Amherst's Pheasant (Chrysolophus amherstiae), Green Peafowl (Pavo muticus), Bulwer's Pheasant and the Crestless Fireback (Lophura erythrophthalma) are notable for their aptitude to forage for crustaceans such as crayfish and other aquatic small animals in shallow streams and amongst rushes in much the same manner as some members of the rail family (Rallidae).

During mating season, the male Capercaille feeds mainly on bilberry leaves, which are toxic to most herbivores

The tragopans (Tragopan), Mikado Pheasant (Syrmaticus mikado) and several species of grouse and ptarmigan are exceptional in their largely vegetarian and arboreal foraging habitats; grouse are especially notable for being able to feed on plants rich in terpenes and quinones – such as sagebrush or conifers –, which are often avoided by other herbivores. But many species of moderate altitudes—for example the long-tailed pheasants of the genus Syrmaticus—also find a great deal of their daily nutritional reqirements in the tree canopies, especially during the snowy and rainy periods when foraging on the ground is dangerous and less than fruitful for a variety of reasons. Although members of the genus Syrmaticus are capable of subsisting almost entirely on vegetarian materials for months at a time, this is not true for many of the subtropical genera. For example, the Great Argus (Argusianus argus) and Crested Argus may do most of their foraging during rainy months in the canopy of the jungle as well. There they are known to forage on slugs, snails, ants and amphibians to the exclusion of plant material. How they forage in the forest canopy during the rainy months is unknown but is a compelling issue for future investigations.

Reproduction

Most galliforms are very prolific, with clutches regularly exceeding 10 eggs in many species. In contrast to most birds which are – at least for a particular breeding season – monogamous, galliforms are often polygynous or polygamous. Such species can be recognized by their pronounced sexual dimorphism.

Galliform young are very precocious and will roam their habitat with their mothers – or both parents in monogamous species – mere hours after hatching. The most extreme case are the Megapodidae, where the adults do not brood but leave incubation to mounds of rotting vegetation, volcanic ash or hot sand. The young have to dig their way out of the nest mounds after hatching, but they emerge from the eggs fully feathered and as soon as they leave the mound, they are able to fly for considerable distances.

Systematics and evolution

Despite its distinct appearance, the Wild Turkey is actually a very close relative of pheasants

The living Galliformes were long divided into some 7 or more families. More recently, it is generally believed that the quite distincly-appearing grouse and turkeys do not warrant separation as families due to their quite recent origin from partridge- or pheasant-like birds. As it seems, the turkeys became huge after their ancestors colonized temperate and subtropical North America where pheasant-sized competitors were absent. The ancestors of grouse, on the other hand, adapted to harsh climate and thus grouse could colonize subarctic regions. Consequently the Phasianidae are expanded in current treatments to include the former Tetraonidae and Meleagrididae as subfamilies.^[1]

The Anseriformes (waterfowl) and the Galliformes together make up the Galloanserae. They are basal among the living neognathous birds, and normally follow the Paleognathae (ratites and tinamous) in modern bird classification systems. This was first proposed in the Sibley-Ahlquist taxonomy and has been the one major change of that proposed scheme that was almost universally adopted. On the other hand, the Galliformes as they were traditionally delimited are called Gallomorphae in the Sibley-Ahlquist taxonomy, which splits the Cracidae and Megapodidae as an order "Craciformes". This is not a natural group however, but rather an erroneous result of the now-obsolete phenetic methodology employed in the Sibley-Ahlquist taxonomy.^[2] Phenetic studies do not distinguish between plesiomorphic and apomorphic characters, which leads to basal lineages appearing as monophyletic groups.

Historically, the buttonquails (Turnicidae), mesites (Mesitornithidae) and the Hoatzin (Opisthocomus hoazin) were placed in the Galliformes too. The former are now known to be shorebirds adapted to an inland lifestyle, whereas the mesites are probably related to at least some of the birds traditionally in the "Gruiformes" assemblage. The relationships of the Hoatzin are entirely obscure, and it is usually treated as a monotypic order Opisthocomiformes to signify this.

Evolution

Given that the oldest known waterfowl, Vegavis iaai, dates from the Late Cretaceous, galliform ancestors must also have roamed the Earth contemporaneously with animals such as Tyrannosaurus rex.^[3]

Indeed, there exist a few fragmentary fossils of putative galliforms from the Cretaceous, of which the most interesting fossil taxon is Austinornis. Formerly referred to as Ichthyornis lentus, Graculavus lentus or Pedioecetes lentus, its partial left tarsometatarsus was found in the Late Cretaceous Austin Chalk near Fort McKinney, Texas. This bird was quite certainly closely related to Galliformes, but whether it was a part of these or belongs elsewhere in the little-known galliform branch of Galloanserae is not clear.^[4]

Other Mesozoic fossils are referred to the Galliformes with considerably less certainty: The very enigmatic Gallornis living in today's France some 140–130 Ma (million years ago) was as far as anyone can tell an ornithuran. But although the possibility that it was a very early galliform cannot be positively excluded, its ancient age makes this seem not particularly likely. The case of Austinornis shows that it is certainly possible to confuse a putative galliform with more primitive ornithurans like Ichthyornis.

Specimen PVPH 237 from the Late Cretaceous Portezuelo Formation (Turonian-Coniacian, about 90 Ma) in the Sierra de Portezuelo (Argentina) makes a far more plausible galliform candidate. This is a partial coracoid of a neornithine bird, which in its general shape and particularly the wide and deep attachment for the muscle joining the coracoid and the humerus (upper arm) bone resembles the more basal lineages of galliforms.^[5]

The Paleogene had several galliforms of now-extinct families, namely the Gallinuloididae, Paraortygidae and Quercymegapodiidae. In the early Cenozoic, there are some additional birds that may or may not be early Galliformes, though even if they are, it is rather unlikely that these belong to extant families:

• †Argillipes (London Clay Early Eocene of England)
• †Coturnipes (Early Eocene of England, and Virginia, USA?)
• †Paleophasianus (Willwood Early Eocene of Bighorn County, USA)
• †Percolinus (London Clay Early Eocene of England)
• †Amitabha (Bridger middle Eocene of Forbidden City, USA) – phasianid?
• †"Palaeorallus" alienus (middle Oligocene of Tatal-Gol, Mongolia)
• †Anisolornis (Santa Cruz Middle Miocene of Karaihen, Argentina)

From the mid-Eocene onwards – about 45 Ma or so –, modern-type galliforms are known, and these completely replace their older relatives in the early Neogene. Since the earliest representatives of living galliform families apparently belong to the Phasianidae – the youngest family of galliforms -, the other families of Galliformes must be at least of Early Eocene origin but might even be as old as the Late Cretaceous. The ichnotaxon Tristraguloolithus cracioides is based on fossil eggshell fragments from the Late Cretaceous Oldman Formation of southern Alberta, Canada which are similar to chachalaca eggs,^[6] but in the absence of bone material their relationships cannot be determined except that they are apparently not from a non-avian dinosaur.

Modern genera of phasianids start appearing around the Oligo-/Miocene boundary, roughly 25–20 Ma. It is not well known whether the living genera of the other, older, galliform families originated around the same time or earlier, though at least in the New World quails, pre-Neogene forms seem to belong to genera that became entirely extinct later on.

A number of Paleogene to mid-Neogene fossils are quite certainly Galliformes, but their exact relationships in the order cannot be determined:

Palaeortyx skeleton, Muséum national d'histoire naturelle, Paris

• †Austinornis (Austin Chalk Late Cretaceous of Fort McKinney, USA) – tentatively placed here, formerly Graculavus/Ichthyornis/Pedioecetes lentus
• †Procrax (middle Eocene? – Early Oligocene) – cracid? gallinuloidid?
• †Palaeortyx (middle Eocene -? Early Pliocene) – phasianid or odontophorid
• †Palaeonossax (Brule Late Oligocene of South Dakota, USA) – cracid?
• †Taoperdix (Late Oligocene) – gallinuloidid? Includes "Tetrao" pessieti
• †Archaealectrornis (Oligocene) – phasianid?
• †Galliformes gen. et sp. indet. (Oligocene) – formerly in Gallinuloides; phasianid?^[7]
• †Archaeophasianus (Oligocene? – Late Miocene) – phasianid? (tetraonine?)
• †Palaealectoris (Agate Fossil Beds Early Miocene of Sioux County, USA) – tetraonine?
• †Linquornis (middle Miocene)
• †Palaeoalectoris (Xiacaowan middle Miocene of Sihong, China)
• †Shandongornis (middle Miocene)
• †"Cyrtonyx" tedfordi (Barstow Late Miocene of Barstow, USA)

List of major taxa

For a long time, the pheasants, partridges and relatives were indiscriminately lumped in the Phasianidae, variously including or excluding turkeys, grouse, New World quails and guineafowl, and divided into two subfamilies – the Phasianinae (pheasant-like forms) and the Perdicinae (partridge-like forms). This crude arrangement was long considered to be in serious need of revision, but even with modern DNA sequence analyses and cladistic methods, the phylogeny of the Phasianidae has resisted complete resolution.^[8]

A tentative list of the higher-level galliform taxa, listed in evolutionary sequence, is:^[8]

• Family †Paraortygidae
• Family †Quercymegapodiidae
• Family †Sylviornithidae – Sylviornis
• Family Megapodidae – mound-builders and scrubfowl, or megapodes
• Family Cracidae – chachalacas, guans and curassows
• Superfamily Phasianoidea
  • Family †Gallinuloididae – tentatively placed here
  • Family Odontophoridae – New World quail
  • Family Numididae – guineafowl
  • Family Phasianidae – pheasants, partridges and relatives
    • Subfamily Arborophilinae – jungle- and wood-partridges
    • Subfamily Coturnicinae – Old World quails, scrub-partridges and spurfowl
    • Subfamily Pavoninae – peafowl and ocellated pheasants
    • Subfamily Gallininae – francolins and junglefowl
    • Subfamily Meleagridinae – turkeys
    • Subfamily Perdicinae – grey partridges (probably belong in either Meleagridinae or Phasianinae)
    • Subfamily Tetraoninae – grouse
    • Subfamily Phasianinae – true pheasants

The relationships of many pheasants and partridges are still very badly resolved and much confounded by adaptive radiation (in the former) and convergent evolution (in the latter).^[9] Thus, the bulk of the Phasianidae can alternatively be treated as a single subfamily Phasianinae. The grouse, turkeys, true pheasants etc would then become tribes of this subfamily, similar of how the Coturnicinae are commonly split into a quail and a spurfowl tribe.^[10]

Note that the taxon Perdicinae is tentatively limited to the genus Perdix and perhaps one or two others. That "the" partridge of Europe is not closely related to other partridge-like Galliformes is already indicated by its sexually dimorphic coloration and numerous (more than 14) rectrices, traits it shares with the other advanced phasianids. However, among these its relationships are obscure; it is entirely unclear whether it is closer to the turkeys or to certain short-tailed pheasants like Ithaginis, Lophophorus, Pucrasia and Tragopan.^[11]

Footnotes

1. ^ Kimball et al. (1999), Dyke et al. (2003), Smith et al. (2005), Crowe et al. (2006a,b)
2. ^ Smith et al. (2005), Crowe et al. (2006a,b)
3. ^ Though not necessarily the same part of the Earth as that particular dinosaur.
4. ^ Clarke (2004)
5. ^ Agnolin et al. (2006)
6. ^ Zelenitsky et al. (1996)
7. ^ Specimen MCZ 342506. A proximal humerus of a bird larger than Gallinuloides: Mayr & Weidig (2004)
8. ^ ^{a b} Kimball et al. (1999, 2001), Crowe et al. (2006a,b)
9. ^ Dyke et al. (2003)
10. ^ See e.g. the phylogenies in Kimball et al. (2006) and Crowe et al. (2006a,b)
11. ^ Kimball et al. (1999, 2001), Smith et al. (2005), Crowe et al. (2006a,b)

References

• Agnolin, Federico L.; Novas, Fernando E. & Lio, Gabriel (2006): Neornithine bird coracoid from the Upper Cretaceous of Patagonia. Ameghiniana 43(1): 245–248. HTML fulltext
• Clarke, Julia A. (2004): Morphology, Phylogenetic Taxonomy, and Systematics of Ichthyornis and Apatornis (Avialae: Ornithurae). Bulletin of the American Museum of Natural History 286: 1–179 PDF fulltext
• Crowe, Timothy M.; Bloomer, Paulette; Randi, Ettore; Lucchini, Vittorio; Kimball, Rebecca T.; Braun, Edward L. & Groth, Jeffrey G. (2006a): Supra-generic cladistics of landfowl (Order Galliformes). Acta Zoologica Sinica 52(Supplement): 358–361. PDF fulltext
• Crowe, Timothy M.; Bowie, Rauri C.K.; Bloomer, Paulette; Mandiwana, Tshifhiwa G.; Hedderson, Terry A.J.; Randi, Ettore; Pereira, Sergio L. & Wakeling, Julia (2006b): Phylogenetics, biogeography and classification of, and character evolution in, gamebirds (Aves: Galliformes): effects of character exclusion, data partitioning and missing data. Cladistics 22(6): 495–532. doi:10.1111/j.1096-0031.2006.00120.x PDF fulltext
• Dyke, Gareth J; Gulas, Bonnie E. & Crowe, Timothy M. (2003): Suprageneric relationships of galliform birds (Aves, Galliformes): a cladistic analysis of morphological characters. Zool. J. Linn. Soc. 137(2): 227–244. doi:10.1046/j.1096-3642.2003.00048.x PDF fulltext
• Kimball, Rebecca T.; Braun, Edward L.; Zwartjes, P.W.; Crowe, Timothy M. & Ligon, J. David (1999): A molecular phylogeny of the pheasants and partridges suggests that these lineages are not monophyletic. Mol. Phylogenet. Evol. 11(1): 38–54. doi:10.1006/mpev.1998.0562 PDF fulltext
• Kimball, Rebecca T.; Braun, Edward L.; Ligon, J. David; Lucchini, Vittorio & Randi, Ettore (2001): A molecular phylogeny of the peacock-pheasants (Galliformes: Polyplectron spp.) indicates loss and reduction of ornamental traits and display behaviours. Biol. J. Linn. Soc. 73(2): 187–198. doi:10.1006/bijl.2001.0536 PDF fulltext
• Kimball, Rebecca T.; Braun, Edward L.; Ligon, J. David; Randi, Ettore & Lucchini, Vittorio (2006): Using molecular phylogenetics to interpret evolutionary changes in morphology and behavior in the Phasianidae. Acta Zoologica Sinica 52(Supplement): 362–365. PDF fulltext
• Mayr, Gerald & Weidig, Ilka (2004): The Early Eocene bird Gallinuloides wyomingensis – a stem group representative of Galliformes. Acta Palaeontologica Polonica 49(2): 211–217. PDF fulltext
• Smith, Edward J.; Shi, Li & Tu, Zhijian (2005): Gallus gallus aggrecan gene-based phylogenetic analysis of selected avian taxonomic groups. Genetica 124(1): 23–32. doi:10.1111/j.1095-8312.2001.tb01356.x (HTML abstract)
• Zelenitsky, Darla K.; Hills, L.V. & Currie, Philip J. (1996): Parataxonomic classification of ornithoid eggshell fragments from the Oldman Formation (Judith River Group; Upper Cretaceous), Southern Alberta. Canadian Journal of Earth Sciences 33(12): 1655–1667. PDF fulltext

External links

Wikimedia Commons has media related to: Galliformes

• The World Pheasant Association - The ultimate authority on Galliformes
• Gamebird Journal - An Online Magazine Devoted To The Gamebird Enthusiast
• Galliformes - Conservation and aviculture

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