Phasianidae

(vertebrate zoology) A family of game birds in the order Galliformes; typically, members are ground feeders, have bare tarsi and copious plumage, and lack feathers around the nostrils.

(Phasianidae)

Class: Aves

Order: Galliformes

Suborder: Phasiani

Family: Phasianidae

Thumbnail description
Plump, ground-based birds of a great size range, with short, broad wings and stout bills and feet; males of larger species are often heavier with striking plumage and elaborate displays

Size
6–49 in (15–125 cm); 1.5 oz–24.2 lb (43 g–11.0 kg)

Number of genera, species
46 genera; 179 species

Habitat
Forest, woodland, bogs, tundra, mountains, savanna, desert fringes

Conservation status
Critically Endangered: 3 species; Endangered: 9 species; Vulnerable: 38 species; Near Threatened: 21 species

Distribution
North America, Europe, Asia, Africa, and Australasia

Evolution and systematics

From studies based on DNA comparisons as well as traditional morphological work, it seems clear that the turkeys (Meleagridinae), grouse (Tetraoninae), and pheasants and Old World partridges (Phasianinae) form an assemblage distinct from the other Galliformes. The current DNA evidence confirms that grouse and turkeys are closely related and probably evolved alongside the other main types within this complex family. When more such data becomes available and the resulting pattern of relationships among these species stabilizes, a completely new taxonomy for the Phasianinae is likely to emerge.

Unraveling the taxonomic affinities of some individual species in this family has produced some real surprises. The DNA evidence shows that the Congo peafowl (Afropavo congensis) is not closely related to either the guineafowls or partridges with which it shares its continent, Africa, having the most in common with the other peafowls (Pavo spp.) in South and Southeast Asia. The Udzungwa forest-partridge (Xenoperdix udzungwensis), first discovered in 1991, appears to be more closely related to the Southeast Asian hill-partridges (Arborophila) than to any African species. The Gunnison sage grouse (Centrocercus minimus) was only recognized as a full species in 2000, and lives in just eight localities at the southern limit for this genus in southwest Colorado and southeast Utah.

Some long-established species have also come under renewed scrutiny, including Edwards's pheasant (Lophura edwardsi) and the imperial pheasant (L. imperialis) from central Vietnam. During 2000 and 2001, the Berlioz's silver pheasant (L. nycthemera berliozi) was deliberately crossed with Edwards's pheasant to produce birds that appear identical to imperial pheasants, thus suggesting that the few imperials recovered from the wild were rare interspecific hybrids. Yet another pheasant had been discovered in the same Annamese lowlands area in 1964, and was adopted as a new species, the Vietnamese pheasant (L. hatinhensis) in 1975. It differs from the Edwards's pheasant only in having white central tail feathers in the male.

Physical characteristics

The species in this family vary enormously in size, from tiny quails (Coturnix spp.) weighing less than 2 oz (43 g) to the wild turkey (Meleagris gallopavo) weighing up to 24.2 lb (11 kg). Their dominant common feature is a heavy rounded body, which is the result of extreme development of the flight muscles over the sternum. This in turn has evolved in parallel with their typical flight behavior: an explosively energetic take-off to gain height followed by a fast glide. Generally, the legs and neck are short, the head and tail small, although in some of the large species, longer necks and tails have evolved.

Facial adornments are many and various, again particularly in the larger species and especially in the males. Male turkeys have a naked red crop and the fleshy and flexible caruncle, which can change rapidly in color from red to blue and dangles beside the beak. The ring-necked pheasant (Phasianuscolchicus) has a bright red skin patch around the eye, while in the crested fireback (Lophura ignita) it is blue. Fleshy and brightly colored wattles are characteristic of Bulwer's pheasant (Lophura bulweri) and the junglefowls (Gallus spp.). Garishly colored bib-like air sacs and paired erectile horns are unique to the tragopans (Tragopan spp.). Other specialties include feathery ears in the eared-pheasants (Crossoptilon), head crests as in the koklass (Pucrasia macrolopha), monals (Lophophorus), and crested wood-partridge (Rollulus rouloul), and neck-ruffs in Lady Amherst's pheasant (Chrysolophus amherstiae) and the ruffed grouse (Bonasa umbellus).

Many grouse species have bright yellow to red fleshy combs above the eyes, which in males especially become engorged and more prominent in the mating season. In those species believed to have polygynous or promiscuous mating systems, the males are more extravagantly adorned and may be up to twice the size of the more dowdy but well-camouflaged females. In the prairie grouse (Tympanuchus spp.), males have a pair of large and brightly colored air sacs on their necks, which they inflate during their breeding season displays. Grouse are adapted to live in cold climates by having an exceptionally thick and heavy plumage, as well as feathering right down to the toes. The ptarmigans (Lagopus spp.) adopt a special white winter plumage so that they remain well camouflaged.

A number of species have long tails, which can be held in fans during displays. In grouse, these are plain, as in the blue grouse (Dendragapus obscurus) or barred, as in sage grouse (Centrocercus urophasianus). In the peafowls (Pavo spp.) and peacock-pheasants (Polyplectron spp.), they bear numerous eye-like ocelli.

Distribution

The relict distribution of the ocellated turkey (Meleagris ocellata) in the Yucatán peninsula of Central America is the southern-most point for this family in the New World, with the wild turkey (M. gallopavo) and prairie grouse (Tympanuchus spp.) originally occupying much of the United States between them. Other grouse species occur over large parts of Canada, including most of the islands in the extreme north. The range of the ruffed grouse (Bonasa umbellus) spans the continent, as does its counterpart in the Old World, the hazel grouse (B. bonasia). The willow grouse (Lagopus lagopus) and rock ptarmigan (L. mutus) have circumpolar distributions in the northern tundra.

Between them, the Old World partridges and pheasants occupy almost all of Europe, Africa, and Australasia. The genus Francolinus contains numerous African species, extending to almost every corner of that continent. The bush-quails (Perdicula spp.) and spurfowl (Galloperdix spp.) occur only in southern Asia, while the hill-partridges (Arborophila spp.) are Southeast Asian and Chinese in distribution.

Habitat

Most grouse species inhabit northern tundra or boreal forests, and the equivalent habitats in isolated mountainous regions further south, as in the case of the capercaillie (Tetrao urogallus) in the Cantabrian Mountains of northern Spain.

Several North American grouse species and the two turkeys have evolved to occupy relatively open temperate and subtropical habitats over much of the continent, the niche occupied by the bustards (Otidae) in the Old World.

Most pheasant species (Phasianini) are forest specialists, with exceptions such as the cheer (Catreus wallichi), which occupies open grass and scrub habitats in the western Himalayan foothills, and the Chinese monal (Lophophorus lhuysii), which lives in alpine scrub and grassland. The great argus (Argusianus argus) occurs in lowland tropical rainforests in Southeast Asia, while the koklass is a temperate forest species of the Himalayas and China.

Partridge (Perdicini) occur in all habitats except for the northern boreal forests and tundra, where they are replaced by grouse. Conversely, in the high alpine areas of central Asia, where there are no grouse, the snowcocks (Tetraogallus), snow partridge (Lerwa lerwa), and Tibetan partridge (Perdix hodgsoniae) occupy their niche. The temperate grasslands of Europe are home to the gray partridge (Perdix perdix) and several rock partridges (Alectoris spp.). Many species occur in tropical grasslands and savanna such as the yellow-necked francolin (Francolinus leucoscepus) in southeastern Africa and the painted francolin (F. pictus) in India. Philby's rock partridge (Alectoris philbyi) lives on rocky slopes in southwestern Arabia. Lowland tropical rainforest species include the crested wood-partridge (Rollulus rouloul) of Southeast Asia and Latham's francolin (F. lathami) from equatorial Africa. Montane forests harbor the Hainan hill-partridge (Arborophila ardens) from China and the red-billed hill-partridge (A. rubirostris) of Sumatra. The swamp francolin (F. gularis) and Manipur bush-quail (Perdicula manipurensis) inhabit wet grasslands south of the Himalayas.

Behavior

Out of the breeding season, open country species such as the wild turkey, snowcocks, African savanna francolins, and the alpine Tibetan eared-pheasant (Crossoptilon harmani) form groups of 20–100, presumably as a protection against predation. Forest species are in general much less gregarious. However, in some species, female gregariousness in the prebreeding period has apparently enabled males to defend small groups of them and thereby acquire a harem, as in the wild turkey and the red junglefowl (Gallus gallus). An alternative tactic is to defend territories in strategically important habitats for females: thus males of the introduced ring-necked pheasant (Phasianus colchicus) in Britain defend territories on woodland edge in farmland where groups of females nest.

Studies of territorial behavior in various grouse species reveal that loud and often repeated calling by the males is the routine method for establishing and maintaining territories. Fighting is also frequent, particularly if there is a large surplus of birds still seeking territories in a limited area of habitat, as for the red grouse (L. l. scoticus) in the British uplands.

The daily routine of birds in this family is rather universal. They emerge from their roosts at dawn for an intensive period of feeding activity, in order to refill their crops following the night fast. After an hour or two, birds retreat to cover, presumably avoiding inclement weather and predators. In winter, arctic grouse need to feed in the open for as little time as possible to avoid excessive heat loss. They have an unusually large crop in which to store rapidly gathered food for later digestion in warmth and safety. Towards the end of the day, there is usually another burst of feeding activity, sometimes followed by calling as families or larger groups gather to roost for the night.

Migration is not a prominent feature of these species. Despite their small size, quails are unusual among them in making long-distance spring and autumn movements between wintering and breeding grounds. The only other species that migrate regularly are the northern-most populations of the rock and willow ptarmigan (Lagopus mutus) that move several hundred miles/kilometers south in fall to escape the worst winter conditions. Some montane pheasant species such as the Himalayan monal (Lophophorus impejanus) have been found to mimic this kind of movement by undertaking altitudinal migrations of up to 4,900 ft (1,500 m) in fall, returning to breed in the sub-alpine scrub in spring after the snow has receded.

Feeding ecology and diet

High-montane snowcocks feed almost exclusively on vegetation. Boreal forest species such as the spruce grouse (Falcipennis canadensis) subsist on the oily buds and needles of conifers throughout the winter months. Grouse also have exceptionally long caecae, the blind-ended tubes in the gut where symbiotic bacteria digest cellulose into sugars. Forest pheasants such as tragopans thoroughly dig over areas of litter and soil with their feet, the monals also using their stout beaks, to depths of at least 12 in (30 cm) in order to feed on tubers, bulbs, and roots.

Tropical forest species take a huge range of items as food, with ants, termites, and other invertebrates of the forest floor and understory being prominent alongside fruits, seeds, and leaves. Prairie chickens also take a significant amount of insect food as adults, mainly in the form of grasshoppers in summer on the North American plains. In the dry grasslands of southern Africa, the gray-winged fracolin (F. africanus) takes a varied mixture of roots and bulbs, and seeds, fruits, and invertebrates. All these open-country species, as well as the wild turkey and the quails, have been able to adapt well to agricultural expansion by feeding in fields of crops and on seeds left after harvesting.

Newly hatched chicks in almost all species rely completely on a protein-rich diet of invertebrates, although they rapidly switch to a less exclusive diet within their first month. An exception is provided by snowcock chicks, which take legumes as a major part of the diet.

Reproductive biology

Nests are usually simple scrapes on the ground, lined with only a little vegetation, and camouflaged by grasses, shrubs, or rocky overhangs. Clutch size in some open-country perdicines and the ring-necked pheasant can be a high as 15–20. In contrast, the Malaysian peacock-pheasant (Polyplectron malacense) lays just one egg.

Incubation is usually carried out entirely by the female. Chicks immediately leave the nest at hatching and feed themselves. The care provided by one or both parents is normally limited to protecting chicks from predators, and sheltering them under their wings in inclement weather and at night. Rudimentary flight is achieved in just 7–10 days in many grouse and partridges, although families may stay together for two to three months.

Females generally come into breeding condition each year including their first, although wild turkeys defer breeding for a year if their fat reserves are low in spring. Males in many larger species do not mate in their first season. Male silver pheasants and tragopans only molt into full adult colors in their second fall, while the male great argus does not achieve adult plumage until the third year. The train of the Indian peafowl takes four years to develop fully.

The promiscuous grouse and pheasants species gather at collective display and mating grounds known as leks. Here, 5–50 males defend territories just a few yards wide, with a very small percentage of the males obtaining the vast majority of copulations. Males with the largest, most decorated, or least damaged tails are most successful. Males with these characteristics also father high-quality chicks, are in better body condition, carry lower parasite loads, and survive better themselves. This suggests that sexual selection via both male-male competition and female choice can cause the evolution of elaborate plumage, bizarre displays, and the much greater body size of the males in such species.

Conservation status

In 2000, of the 179 extant species in the Phasianidae, 50 (28%) were included on the IUCN Red List as being threatened with extinction, a proportion nearly three times that for all birds (11%). A further 21 (12%) were classified Near

Threatened. Of the 108 species native to Asia, 70% of the forest specialists are threatened, compared to only 18% of those living in open habitats, implying that forest degradation and fragmentation are important threats. However, being ground-based and often either large or gregarious, these species are universally harvested as a source of food. Over 90% of the threatened species in Asia are suspected of being over-hunted.

Only one species in this family has become Extinct relatively recently: the New Zealand quail (Coturnix novaezelandiae) was common on both North and South Islands in 1850, but was last seen in 1876; it was probably wiped out by a disease carried by an introduced bird species. One of the Critically Endangered species is the Djibouti francolin (Francolinus ochropectus), which is only known from one small area of highly disturbed juniper forest in this politically unstable corner of Africa. The other two species in this highest threat category are the Himalayan quail (Ophrysia superciliosa) and the gorgeted wood-quail (Odontophorus strophium).

The nine Endangered species are: the newly-described Gunnison sage grouse (Centrocercus minimus); Edwards's pheasant (Lophura edwardsi); Vietnamese pheasant (L. hatinhensis); Bornean peacock-pheasant (Polyplectron schleiermacheri); Nahan's francolin (Francolinus nahani) on the border between Uganda and the Democratic Republic of Congo; Mount Cameroon francolin (F. camerunensis); Sichuan hill-partridge (Arborophila rufipectus) from central southern China; orange-necked hill-partridge (A. davidi) from south Vietnam; and chestnut-headed hill-partridge (A. cambodiana).

Significance to humans

It is easy to suggest that this family of birds is of greater importance to the human race than any other, as it contains the wild ancestors of both domestic chickens and turkeys, as well as many species that have been hunted in the wild for food over millennia. The precise origins of the domestic chicken are uncertain, but there is a common consensus that the progenitor is the red junglefowl. Archaeological investigations at the sites of cities dating from the third millennium B.C. in the Indus valley of south Asia indicate that their sophisticated inhabitants kept domesticated fowl as well as a variety of hoofed livestock. By 1500 B.C., the chicken was being used in China, Egypt, and northwest Europe. It subsequently achieved global distribution, even reaching many islands in the South Pacific where it became feral.

Bones found during investigations of pre-Columbian settlements in North America suggest that the wild turkey was an important source of meat in the diet of Native Americans. By around A.D. 500–700, it was being kept as a domesticated bird by people living in northern New Mexico and Arizona. The domestic turkey arrived in Europe as a result of the 1519 Cortés expedition to Mexico by the Spaniards, and in 1607 it was taken back to the New World by the first European settlers. It now takes a place of pride on tables at both Thanksgiving in the United States and Christmas throughout the Western world.

It has been judged that at the end of the 1970s around 8.5 million grouse were hunted each year in North America. One species takes the brunt of this harvest: the ruffed grouse (Bonasa umbellus) at around six million per year, but more than half a million sharp-tailed grouse and sage grouse are also taken. In the Old World the willow grouse is cropped at a rate of about eight million per year, mainly in Russia, and in Fenno-Scandinavia the annual total for all the species is more than half a million birds. Since the late nineteenth century in Britain, the habitat of the red grouse has been nurtured in order to provide sport hunting of the highest caliber. In most species, it is believed that hunting has little effect on populations, because a high proportion of the birds shot would die through some other cause.

In the European Alps, hunting revenues are sufficient to support the costs of habitat preservation and improvement for capercaillie. Clear felling of old open woodland and its replacement by forests in which the trees are much closer together, reduces or obliterates the understory layer on which this species depends for food and nesting cover. Efforts to reverse these effects for capercaillie have had a wider impact on the utility of the habitat for other wildlife: areas where these grouse occur have more woodpeckers and a greater number of songbird species than places not yet re-colonized.

Species accounts

Resources

del Hoyo, J., A. Elliot, and J. Sargatal, eds. Handbook of the Birds of the World. Vol. 2. New World Vultures to Guineafowl. Barcelona: Lynx Edicions, 1994.

Delacour, J. The Pheasants of the World. 2nd edition. Hindhead: Spur/Saiga/World Pheasant Association, 1977.

Fuller, R. A., J. P. Carroll, and P. J. K. McGowan, eds. Partridges, Quails, Francolins, Snowcocks, Guineafowl, and Turkeys. Status Survey and Conservation Action Plan 2000-04. WPA/BidLife/SSC Partridge, Quail and Francolin Specialist Group. Gland and Cambridge: IUCN/Reading: World Pheasant Association, 2000.

Fuller, R. A. and P. J. Garson, eds. Pheasants. Status Survey and Conservation Action Plan 2000-04. WPA/BidLife/SSC Pheasant Specialist Group. Gland and Cambridge: IUCN/Reading: World Pheasant Association, 2000.

Hill, D. A., and P. A. Robertson. The Pheasant. Oxford: BSP Professional Books, 1988.

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

Johnsgard, P. A. The Pheasants of the World. 2nd ed. Washington, DC: Smithsonian Institution Press, 1999.

Johnsgard, P. A. The Quails, Patridges and Francolins of the World. London: Oxford University Press, 1986.

Madge, S., and P. McGowan. "Pheasants, Partridges, and Grouse." Helm Identification Guides. London: Christopher Helm, 2002.

Potts, G. R. The Partridge. London: Collins, 1986.

Storch, I., ed. Grouse. Status Survey and Conservation Action Plan 2000-04. WPA/BidLife/SSC Grouse Specialist Group. Gland and Cambridge: IUCN/Reading: World Pheasant Association, 2000.

Dinesen, L., T. Lehmberg, J. O. Svendsen, L. A. Hansen, and J. Fjeldså. "A New Genus and Species of Perdicine Bird (Phasianidae, Perdicini) from Tanzania: A Relict Form with Indo-Malayan Affinities." Ibis 136 (1994): 2–11.

Kimball, R. T., E. L. Braun, and J. D. Ligon. "Resolution of the Phylogenetic Position of the Congo Peafowl, Afropavo congensis: A Biogeographic and Evolutionary Enigma." Proceedings of the Royal Society of London Series B 264 (1997): 1517–1523.

Kimball, R. T., E. L. Braun, P. W. Zwartes, T. M. Crowe, and J. D. Ligon. "A Molecular Phylogeny of the Pheasants and Partridges Suggests that these Lineages are Not Monophyletic." Molecular Phylogenetics and Evolution 11(1999): 38–54.

Young, J. R., C. E. Braun, S. J. Oyler-McCance, T. W. Quinn, and J. W. Hupp. "A New Species of Sage Grouse (Phasianidae: Centrocercus) from Southwestern Colorado, USA." Wilson Bulletin 112 (2000): 445.

Game Conservancy Trust. Fordingbridge, Hampshire SP6 1EF United Kingdom. Phone: +44 1425 652381. Fax: +44 1425651026. E-mail: info@gct.org.uk Web site:

Ruffed Grouse Society. 451 McCormick Rd, Coraopolis, PA 15108. Phone: (888) 564-6747. Fax: (412) 262-9207. Web site:

World Pheasant Association. P.O. Box 5, Lower Basildon, Reading RG8 9PF United Kingdom. Phone: +44 1 189 845140. Fax: +44 118 963369. E-mail: office@pheasant.org.uk Web site:

[Article by: Peter Jeffery Garson, DPhil]

The noun has one meaning:

Meaning #1: pheasants; quails; partridges
  Synonym: family Phasianidae