Fowls and pheasants

(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

List of Species

Wild turkey

Meleagris gallopavo

subfamily

Meleagridinae

taxonomy

Meleagris gallopavo Linnaeus, 1758, North America = Mexico. Six subspecies.

other common names

French: Dindon sauvage; German: Truthuhn; Spanish: Guajolote Gallipavo.

physical characteristics

Male 39–49 in (100–125 cm); female 30–37 in (76–95 cm); male 11.0–24.2 lb (5.0–11.0 kg); female 6.6–11.0 lb (3.0–5.0 kg). Males have a bare blue and pink head, red wattles, dark plumage with iridescent green and bronze highlights, white-barred flight feathers, a blackish breast tuft, and pinkish spurred legs. Females are duller in color and smaller than males.

distribution

Native of central and northern Mexico, throughout the United States ranging from southern Vermont to Florida, and west to Washington, Oregon, and California.

habitat

Prefers mix of hardwood forest, scrub, grass, and agricultural land, but tolerant of dry scrub and subtropical forest.

behavior

Gathers into large flocks of 50, exceptionally 500, in winter with males usually separate from females and young; no territorial defense; will run to cover, only flying routinely to reach communal roosts in trees.

feeding ecology and diet

Feeds mainly by picking at ground; takes leaves, shoots, small seeds, acorns, buds, fruits, as well as grasshoppers, crabs, and small vertebrates.

reproductive biology

Male display starts in February, and later takes up most of their time; successful males attract a group of four to five females for mating; nest in dense cover; clutch size usually 10–12; incubation by female alone, 27–28 days; female and young remain together until following spring.

conservation status

Until 1940s, over-hunting in the United States was reducing its range and population; following many successful translocations and systematic management of hunting, it is again common and widespread; it is local and much less common in Mexico.

significance to humans

A hunted game bird; the subject of turkey-calling contests; a flagship for its habitats.

Willow ptarmigan

Lagopus lagopus

subfamily

Tetraoninae

taxonomy

Tetrao lagopus Linnaeus, 1758, Swedish Lapland. Nineteen subspecies.

other common names

English: Red grouse, willow grouse; French: Lagopéde des saules; German: Moorschneehuhn; Spanish: Lagópodo Común.

physical characteristics

14–17 in (36–43 cm); male 0.9–1.8 lb (405–795 g); female 0.9–1.5 lb (405–700 g). Males have a rusty head and upper-parts, bright red eye combs, white underparts, and a black tail. Females are grayer, are more heavily barred on the breast and flanks, and lack the bright red eye combs. Both sexes are all white in winter except for black tail.

distribution

Circumpolar between 47° and 76°N.

habitat

Arctic tundra, sub-arctic scrub, and boreal forest edge; preferring moister areas with dwarf deciduous trees.

behavior

In large groups of variable sex ratio in winter; males highly territorial in spring through calling from landmarks and in flight.

feeding ecology and diet

Willow and birch buds and twigs in winter; invertebrates taken especially by young chicks in summer; berries in fall.

reproductive biology

Mostly monogamous. Pairs occupy exclusive territories; nesting starts April–June depending on latitude; clutch size eight to 11; incubation 22 days; male broods chicks; families remain together until fall.

conservation status

Not threatened. Locally common and widespread.

significance to humans

Locally managed to provide a substantial hunted surplus; keenly hunted in the United Kingdom, Scandinavian countries, Finland, and Russia, with 2.4 million birds taken annually in these areas together.

Capercaillie

Tetrao urogallus

subfamily

Tetraoninae

taxonomy

Tetrao urogallus Linnaeus, 1758, Sweden. Twelve subspecies.

other common names

French: Grand tétras; German: Auerhuhn; Spanish: Urogallo Común.

physical characteristics

Male 32–35 in (83–90 cm); female 23–25 in (59–64 cm); male 7.3–14.3 lb (3.3–6.5 kg); female 3.3–5.5 lb (1.5–2.5 kg). Males are mostly slate gray with a blackish head and neck, red eye combs, glossy greenish black breast, dark brown wings with white carpal patch, varying amounts of white on upper wings and underparts, and long, rounded tail. Females are mottled and barred in gray, buff, and black with a large rusty breast patch.

distribution

Northern Britain and Scandinavia to eastern Russia; more fragmented in eastern and southeastern Europe, the Alps; isolated populations in northern Spain and Pyrenees.

habitat

Mainly in old conifer forest with moderate understory usually of bilberry, interspersed with bogs; up to 6,600 ft (2,000 m) in Pyrenees.

behavior

Male usually alone, while females and young form wintering groups of up to 10; males gather loosely in lek areas to defend territories and attract females using calls, erect strutting, and tail-fanning displays.

feeding ecology and diet

Pine needles, holly leaves, birch buds, berries; leaves of heath plants; young chicks especially take invertebrates.

reproductive biology

Promiscuous. Laying in April–June; nest in thick cover in forest; clutch size six to nine; incubation 26 days; chicks able to fly after three weeks; males defer mating until third year.

conservation status

Small and fragmented populations are threatened and prone to extinction; continued hunting a particular threat in central and south Europe, where increased predator numbers, alpine tourism, and collisions with power lines and deer fences all cause problems.

significance to humans

Attracts trophy hunters; hunted for food mainly in fall.

Chinese grouse

Bonasa sewerzowi

subfamily

Tetraoninae

taxonomy

Tetrastes sewerzowi Przevalski, 1876, Gansu, China. Monotypic.

other common names

English: Severtzov's grouse, black-breasted hazel grouse; French: Gélinotte de Severtzov; German: China-haselhuhn; Spanish: Grévol Chino.

physical characteristics

13–14 in (33–36 cm); 0.6–0.7 lb (270–310 g). Brownish gray with black bars on upperparts, black chin bordered in white, a chestnut upper breast, and underparts spotted and barred in dark gray and ochre. Small erectile crest on head. Barred pattern on the tail distinguishes this species from B. bonasia.

distribution

China: central Gansu to southern Quinghai, eastern Tibet, northwestern Yunnan, and northern and western Sichuan.

habitat

Montane forests at 3,300–13,100 ft (1,000–4,000 m); conifer near treeline, birch and conifer below, willow thickets on riverbanks.

behavior

Forms flocks of up to 15 for fall-winter; spring dispersal for breeding; males repeat noisy display for most of day, and fight in treetops.

feeding ecology and diet

Forages on ground and in trees for buds and shoots of willow and birch, also taking various flowers, seeds, and berries.

reproductive biology

Monogamous. Nests on ledges and stumps in May–June; clutch size five to eight; incubation 25 days.

conservation status

Near Threatened as forest clearance and fragmentation causes local extinctions, with hunting and egg-collecting a problem outside protected areas.

significance to humans

A hunted resource locally.

Ruffed grouse

Bonasa umbellus

subfamily

Tetraoninae

taxonomy

Tetrao umbellatus Linnaeus, 1766, Pennsylvania, United States. Fourteen subspecies.

other common names

French: Gélinotte huppée; German: Kragenhuhn; Spanish: Grévol Engolado.

physical characteristics

17–19 in (43–48 cm); male 1.3–1.4 lb (600–650 g); female 1.1–1.3 lb (500–590 g). Cryptic plumage; gray and brown color morphs with gray commoner in northern parts of range and brown commoner in southern parts. Small crest on head, erectile black ruff on sides of neck, and fan-shaped tail with distinctive subterminal dark band.

distribution

North America from Alaska to Labrador and Nova Scotia, south to California and Utah in west and through Appalachians to northern Georgia in east; Nevada and Newfoundland.

habitat

Pacific Coast rainforest, boreal, and dry deciduous woodlands; prefers a mixed-age forest mosaic with aspen and brushwood clearings.

behavior

Roosts in conifers; drums with wings while in upright posture year round, but most intensely at dawn in March–June when males defend territories and sometimes form a loose lek.

feeding ecology and diet

Buds and twigs from aspens and other deciduous trees; herb flowers and catkins; berries and some invertebrates; newly hatched chicks depend on insects; also fungi and acorns.

reproductive biology

Promiscuous. Nests in May on forest floor; clutch size 10–12; incubation 23–24 days; chicks can fly at 10–12 days.

conservation status

Widespread and common in north of range; elsewhere range contractions have been reversed by restocking.

significance to humans

Most hunted grouse in North America.

Greater prairie chicken

Tympanuchus cupido

subfamily

Tetraoninae

taxonomy

Tetrao cupido Linnaeus, 1758, Virginia, United States. Two extant subspecies.

other common names

English: Pinnated grouse, prairie grouse; French: Tétras des prairies; German: Präriehuhn; Spanish: Gallo de las Praderas Grande.

physical characteristics

16–18 in (41–47 cm); male 2.2 lb (990 g); female 1.7 lb (770 g). Brown overall with extensive barring on both upperparts and underparts; short, rounded, blackish tail; elongated feathers on the sides of the neck (pinnae) are erect during courtship; golden yellow cervical sacs; yellow to orange eye combs. Females similar to males but with smaller pinnae and smaller, paler cervical sacs.

distribution

North America, mainly from Oklahoma to North Dakota.

habitat

Prairie remnants amid arable cropland.

behavior

Classic lek-forming species; spectacular display, audible over a mile or more; gather into large mixed-sex flocks in winter.

feeding ecology and diet

Takes acorns, smaller seeds, leaves, buds; cultivated grains (corn, soya); grasshoppers and other invertebrates.

reproductive biology

Promiscuous. Mating at leks, generally excluding yearling males; nests in thick grass cover in April–June; clutch size eight to 13; incubation 23–25 days.

conservation status

Not threatened overall, but the heath hen (T. c. cupido) of New England is Extinct, and Atwater's prairie chicken (T. c. atwateri) persists as two populations in southeastern Texas.

significance to humans

Enjoyed by bird-watchers for the spring spectacle and used as a flagship species for prairie conservation.

Tibetan snowcock

Tetraogallus tibetanus

subfamily

Phasianinae (Tribe Perdicini)

taxonomy

Tetraogallus tibetanus Gould, 1854, Tibet = Ladakh, India. Four subspecies.

other common names

French: Tetraogalle du Tibet; German: Tibetkönigshuhn; Spanish: Perdigallo Tibetano;.

physical characteristics

19–22 in (50–56 cm); male 3.3–4.0 lb (1.5–1.8 kg); female 2.6–3.5 lb (1.2–1.6 kg). Distinguished from other snowcocks by white underparts with heavy blackish streaks on flanks.

distribution

Tibetan plateau including northern India, Nepal, Bhutan, eastern Tajikistan, Tibet, and adjacent parts of Qinghai, Gansu, Sichuan, and Yunnan in China.

habitat

Mainly on bare and grassy slopes at 16,400–19,700 ft (5,000–6,000 m); stays above tree-line year round.

behavior

Gathered in groups of up to 50 out of breeding season; pairs formed by April and males call from vantage points at dawn; runs if disturbed, only taking flight reluctantly; roosts in rocky scree, flying downhill early to start feeding.

feeding ecology and diet

Roots, shoots, seeds, and berries; chicks consume legumes, but few invertebrates.

reproductive biology

Probably monogamous; nests late May concealed by shrub or boulder; clutch size four to seven.

conservation status

Not threatened, having an extensive range; relatively common in Nepal.

significance to humans

None known.

Red-necked francolin

Francolinus afer

subfamily

Phasianinae (Tribe Perdicini)

taxonomy

Tetrao afer Muller, 1776, Benguela, Angola. Seven subspecies.

other common names

English: Bare-throated francolin, red-necked spurfowl; French: Francolin à gorge rouge; German: Rotkehlfrankolin; Spanish: Francolín Gorgirrojo.

physical characteristics

10–15 in (25–38 cm); 1.0–1.7 lb (440–770 g). Distinguished from F. rufopictus and F. swainsonii by scarlet throat and red bill and legs.

distribution

Africa, from Angola, across Congo basin to coastal Kenya, and from Rift Valley to east coast, south to southeast South Africa.

habitat

Moist evergreen forest in west and north of range, but in drier grass and scrub elsewhere.

behavior

Reluctant to fly, runs if disturbed; males call early in the day and are territorial near nests; roost at night and midday in bushes and trees; single or in small parties, sometimes with other francolin species.

feeding ecology and diet

Tubers, roots, bulbs, shoots, berries, crops, and invertebrates, including ticks and termites.

reproductive biology

Most breeding late in rainy seasons with two clutches in some places; probably monogamous; clutch size three to nine; incubation 23 days; chicks fly at 10 days and fully grown in four months; families together for most of year.

conservation status

Not threatened and covering a large range; locally numerous but hunting may reduce numbers significantly in places.

significance to humans

None known.

Gray partridge

Perdix perdix

subfamily

Phasianinae (Tribe Perdicini)

taxonomy

Tetrao perdix Linnaeus, 1758, Sweden. Seven subspecies.

other common names

English: Common partridge; French: Perdrix grise; German: Rebhuhn; Spanish: Perdiz Pardilla.

physical characteristics

11–12 in (29–31 cm); male 0.7–1.3 lb (325–600 g); female 0.7–1.3 lb (310–570 g). Appearance varies especially in terms of grayer or browner plumage. Races in the western parts of the range tend to be more rufous brown, while those in the east are generally grayer and paler.

distribution

West to southeastern Europe, including south Scandinavia to Siberia and south to Kazakhstan and western Xinjiang.

habitat

Native of temperate grasslands and steppe, but now mainly in less intensively managed croplands; up to 8,500 ft (2,600 m) in Spain and Caucasus.

behavior

In parties of 5–25, consisting of one or more families, in over-lapping home ranges for winter; pairs form and live in more exclusive areas in spring.

feeding ecology and diet

Weed and cereal seeds, grass, and clover leaves, with chicks completely dependent on insects for first two weeks.

reproductive biology

Usually monogamous. Nests April–June depending on locality; clutch size usually 15–17; incubation 23–25 days; chicks fly at two weeks and reach adult weight in three months.

conservation status

Still widespread and locally abundant, but adversely affected by intensive farming because of removal of nesting cover, herbicide treatment for weeds, and reduction in insect availability through pesticide applications to crops; 80% reduction estimated overall.

significance to humans

An important game bird for hunting in Europe and United States; flagship for conservation in lowland agricultural landscapes.

King quail

Coturnix chinensis

subfamily

Phasianinae (Tribe Perdicini)

taxonomy

Tetrao chinesis Linnaeus, 1766, China and Philippines = Nanking, China. Ten subspecies.

other common names

English: Asian blue quail, painted quail, Chinese quail, blue-breasted quail; French: Caille peinte; German: Zwergwachtel; Spanish: Codorniz China.

physical characteristics

5–6 in (12–15 cm); male 1.2–1.7 oz (35–48 g); female 1.1–1.4 oz (31–41 g). Males are a dark brownish blue with a lighter bluish gray breast and chestnut belly; the face and throat are black and white. Females are a mottled brown overall and lack the black-and-white coloration on the face and throat.

distribution

Southwestern and northeastern India, Sri Lanka, eastern Myanmar, Indochina to Hainan and Taiwan, Malaysian peninsula, Philippines and Indonesia, northern and eastern coast of Australia.

habitat

Wet shrubland, swampy grassland, rice paddy, mainly in lowlands and coastal areas, but up to 6,600 ft (2,000 m) in Sri Lanka and India.

behavior

Crouches or runs rather than flying if disturbed; dust-bathes in open drier areas; usually in pairs or families.

feeding ecology and diet

Leaves, grass, seeds, and invertebrates, especially termites.

reproductive biology

Strong pair bond and assumed monogamous; nest often domed with grasses and sedges; clutch size four to eight; incubation 18–19 days by female; two broods per year in good conditions; chicks mature in eight weeks.

conservation status

Not threatened, widespread but cryptic; likely to be declining as favored swampy grasslands are drained for agriculture throughout range.

significance to humans

None known.

Udzungwa forest-partridge

Xenoperdix udzungwensis

subfamily

Phasianinae (Tribe Perdicini)

taxonomy

Xenoperdix udzungwensis Dinesen et al., 1994, Ndundulu Mountains, Tanzania. Monotypic.

other common names

English: Udzungwa partridge; French: Xénoperdrix de Tanzanie; German: Udzungwawachtel; Spanish: Perdiz de Udzungwa.

physical characteristics

11 in (29 cm); 8–9 oz (220–239 g); male slightly larger. Barred upperparts and blotched underparts; red bill.

distribution

First found in 1991 and known only from the eastern Udzungwa highlands and the Rubeho Mountains in southern Tanzania, Africa.

habitat

Montane and sub-montane evergreen forest with open understory at 4,400–6,200 ft (1,350–1,900 m).

behavior

Usually in small flocks of up to eight; roosts in trees and shrubs.

feeding ecology and diet

Forages for invertebrates and seeds by searching litter on forest floor.

reproductive biology

Adults with chicks seen in November–December. No other information is available.

conservation status

Vulnerable. Only known from four populations, but these populations appear to be stable.

significance to humans

None known.

Sichuan hill-partridge

Arborophila rufipectus

subfamily

Phasianinae (Tribe Perdicini)

taxonomy

Arboriphila rufipectus Boulton, 1932, west Sichuan, China. Monotypic.

other common names

English: Boulton's hill-partridge, Sichuan partridge; French: Torquéole de Boulton; German: Boultonbuschwachtel; Spanish: Arborófila de Sichuán.

physical characteristics

12 in (29–31 cm); male 14–17 oz (410–470 g); female 12–13 oz (350–380 g). Distinguished from the common hill-partridge by a white throat and russet breast patch.

distribution

China: Southern Sichuan and adjacent northern Yunnan within a fragmented range.

habitat

Primary subtropical broadleaf forest, and adjacent disturbed or broadleaf plantation areas, with relatively open understory, at 3,600–7,400 ft (1,100–2,250 m).

behavior

Pairs in spring, with family parties staying together only until late fall.

feeding ecology and diet

Seeds and fruits from forest floor and shrubs, some invertebrates.

reproductive biology

Breeding pairs widely separated, probably monogamous; nests concealed among tree roots in April–May; clutch size five to six.

conservation status

Endangered; threatened by clear-felling of primary forest, agricultural encroachment, bamboo-shoot collection and livestock browsing; only in one small protected area.

significance to humans

None known.

Crested wood-partridge

Rollulus rouloul

subfamily

Phasianinae (Tribe Perdicini)

taxonomy

Phasianus rouloul Scopoli, 1786, Malacca, Malaysia. Monotypic.

other common names

English: Crested partridge, Roulroul; French: Rouloul couronné, German: Straußwachtel; Spanish: Perdiz rulrul.

physical characteristics

10 in (25 cm); male 8–11 oz (225–300 g); female 8–10 oz (225–275 g). Males have a spectacular reddish crest, dark plumage, and bright red bare parts. Females are mostly green with a gray head.

distribution

Southern Myanmar and Thailand through Malaysian peninsula to Sumatra and Borneo.

habitat

Lowland tropical rainforest, including disturbed areas, up to 3,900 ft (1,200 m).

behavior

Encountered as singles, pairs, families, and larger groups of up to 15; scatters on foot if disturbed, then regroups using calls; paired birds remain close together calling in soft whistles; roosts in low shrubs and trees.

feeding ecology and diet

Seeds, fruits, and invertebrates.

reproductive biology

Breeds mainly in first half of year; nest concealed on forest floor under pile of leaves; clutch size five to six; incubation 18 days; unusually for Galliformes, both parents actively feed chicks.

conservation status

Near Threatened due to continuing degradation and clearance of lowland tropical forests in Southeast Asia.

significance to humans

None known.

Satyr tragopan

Tragopan satyra

subfamily

Phasianinae (Tribe Phasianini)

taxonomy

Meleagris satyra Linnaeus, 1829, Bengal = Sikkim, India. Mono-topic.

other common names

English: Crimson tragopan, Indian tragopan, crimson horned pheasant; French: Tragopan satyre; German: Satyrtragopan; Spanish: Tragopán Sátiro.

physical characteristics

Male 26–28 in (67–72 cm); female 22–23 in (57–59 cm); male 3.5–4.6 lb (1.6–2.1 kg); female 2.2–2.6 lb (1.0–1.2 kg). Males have deep red underparts, bare blue facial skin, and brown plumage on lower back and rump; upperwing-coverts also brown. Females are dull brown to rufous with bars and lance-shaped markings.

distribution

Himalayas from northern India through Nepal, Sikkim, Bhutan to western Arunachal Pradesh, including southeastern Tibet.

habitat

Temperate montane forest with dense understory at 5,900–14,100 ft (1,800–4,300 m).

behavior

Usually in pairs or singles early in year, and family parties in July–September; male's repeated wailing call at dawn from roost in April–June heralds breeding season.

feeding ecology and diet

Roots, bulbs, and invertebrates, and plucking leaves from cover and trees.

reproductive biology

Monogamous in captivity; usually crude stick nests are constructed up to 20 ft (6 m) in trees; clutch size two to three; incubation 28 days.

conservation status

Not threatened and still widespread, but hunted (except in Bhutan).

significance to humans

Used as a flagship for central Himalayan forest conservation campaigns.

Chinese monal

Lophophorus lhuysii

subfamily

Phasianinae (Tribe Phasianini)

taxonomy

Lophophorus lhuysii Geoffrey St. Hilaire and Verreaux, 1866, Moupin, China. Monotypic.

other common names

English: Chinese monal pheasant; French: Lophophore de Lhuys; German: Grünschwanzmonal; Spanish: Monal Coliverde.

physical characteristics

Male 30–31 in (76–80 cm); female 28–29 in (72–75 cm); 6.2–7.0 lb (2.8–3.2 kg). Larger and more heavily built than other pheasants in this genus.

distribution

China: centered on western Sichuan, but extending into southeastern Quinghai, southern Gansu, northeastern Tibet, and northwestern Yunnan.

habitat

Alpine meadows and sub-alpine scrub adjacent to highest conifer forests, mostly at 9,200–16,100 ft (2,800–4,900 m).

behavior

Groups of two to eight individuals common in winter, with single- and mixed-sex flocks being seen in spring; a vocal species at roost in spring and summer, and when alarmed.

feeding ecology and diet

Tubers and bulbs, but also takes moss, leaves, flowers, and some invertebrates.

reproductive biology

Breeding starts during March in snow; nests at 12,500–13,100 ft (3,800–4,000 m); clutch size three to five; incubation 28 days.

conservation status

Vulnerable and on CITES Appendix I, prohibiting trade in wild birds.

significance to humans

None known.

Red junglefowl

Gallus gallus

subfamily

Phasianinae (Tribe Phasianini)

taxonomy

Phasianus gallus Linnaeus, 1758, Poulo Condor, Vietnam. Five subspecies.

other common names

English: Wild junglefowl; French: Coq bankiva; German: Bankivahuhn; Spanish: Gallo Bankiva.

physical characteristics

Male 25–30 in (65–78 cm), female 16–18 in (42–46 cm); male 1.5–3.2 lb (0.7–1.5 kg); female 1.1–2.3 lb (0.5–1.1 kg).

distribution

Northern India, Nepal, and Bangladesh, southern Yunnan to Hainan Island, Southeast Asian peninsula, and Sumatra, Java, and Bali.

habitat

Woodland edge and secondary scrub in tropical and sub-tropical areas from sea level to 6,560 ft (2,000 m).

behavior

Often seen in groups consisting of one male, several females, and offspring; roosts socially in trees or bushes.

feeding ecology and diet

Seeds, including rice, and invertebrates, including eggs.

reproductive biology

Polygamous. Breeds almost year round in India and Malaysia; nests typically under bushes or in bamboo; clutch size five to six; incubation 18–21 days; chicks can fly at seven days.

conservation status

Widespread and locally common.

significance to humans

The supposed progenitor of all domestic fowl, and therefore arguably the most important bird species of all to humans.

Edwards's pheasant

Lophura edwardsi

subfamily

Phasianinae (Tribe Phasianini)

taxonomy

Gennaeus edwardsi Oustalet, 1896, Quangtri, Vietnam. Two subspecies.

other common names

English: Annam pheasant; French: Faisan d'Edwards; German: Edwardsfasan; Spanish: Faisán de Edwards.

physical characteristics

23–25.5 in (58–65 cm); 2.4 lb (1.1 kg). Males have black plumage with a blue sheen and metallic green fringes on upperwing-coverts; small white crest on head. Females are chestnut brown with darker flight feathers and tail and no crest.

distribution

Very restricted range in adjacent parts of Quangtri, Thua Thien, and Quang Binh provinces, east of the Annamese mountains in central Vietnam.

habitat

Primary and secondary forest on level lowlands below 1,300 ft (400 m).

behavior

Little known, but shy and fond of dense understory.

feeding ecology and diet

Nothing known.

reproductive biology

Clutch size four to seven; incubation 21–22 days in captivity.

conservation status

Endangered and on CITES Appendix I, prohibiting trade in wild birds.

significance to humans

None known.

Brown eared-pheasant

Crossoptilon mantchuricum

subfamily

Phasianinae

taxonomy

Crossoptilon mantchuricum Swinhoe, 1863, vicinity of Peking = Beijing, China. Monotypic.

other common names

English: Manchurian eared-pheasant; French: Hokki brun; German: Brauner ohrfasan; Spanish: Faisán Orejudo Pardo.

physical characteristics

37–39 in (96–100 cm); male 3.7–5.5 lb (1.7–2.5 kg); female 3.3–4.4 lb (1.5–2.0 kg). The only brownish member of its genus.

distribution

Formerly widespread in China; now mainly in Shanxi.

habitat

Mixed montane forest with shrub understory at 3,600–8,500 ft (1,100–2,600 m).

behavior

Encountered in large groups (10–30) moving through forest for much of the year.

feeding ecology and diet

Roots, bulbs, and tubers, but also a wide variety of leaves, some fungi, and invertebrates.

reproductive biology

Males issue long calls in April–June; nests on ground in conifer forest; clutch size usually five to eight; incubation 26–27 days.

conservation status

Vulnerable and on CITES Appendix I, prohibiting trade in wild birds.

significance to humans

Tail feathers have been worn on military uniforms since about 500 b.c., reflecting battles fought by males in the mating season.

Ring-necked pheasant

Phasianus colchicus

subfamily

Phasianinae (Tribe Phasianini)

taxonomy

Phasianus colchicus Linnaeus, 1758, Africa, Asia = Rion River. Thirty-one subspecies.

other common names

English: Common pheasant; French: Faisan de Colchide; German: Fasan; Spanish: Faisán Vulgar.

physical characteristics

Male: 29–35 in (75–89 cm); female 21–24 in (53–62 cm); male 2.6 lb (1.2 kg); female 2.0 lb (0.9 kg).

distribution

Japan, Taiwan, central and eastern China, with apparently isolated populations spread across central Asia to the Caucasus; possibly into southeastern Europe; introductions worldwide.

habitat

Mixed temperate scrub, riverine and woodland edge, adjacent to cultivation, avoiding dense forest, dry areas, and high mountains.

behavior

Often in large feeding groups with wide variation in sex ratio; tends to run for cover rather than fly if disturbed.

feeding ecology and diet

Leaves, cereal grains, tree seeds, buds, fruits and some invertebrates as adults; invertebrates essential to young for first month after hatching.

reproductive biology

Single males crow loudly in spring and some associate with a group of females, suggesting harem polygyny; nests in ground cover; clutch size eight to 14; incubation 22–25 days.

conservation status

The most widespread and common pheasant in the world, although some of its isolated western subspecies may be threatened.

significance to humans

Tens of millions of birds are artificially reared each year for release in sport hunting enterprises, especially in Europe and North America.

Palawan peacock-pheasant

Polyplectron emphanum

subfamily

Phasianinae (Tribe Phasianini)

taxonomy

Polyplectron emphanum Temminck, 1831, Sunda Islands or Moluccas (=error: Palawan Island, the Philippines). Monotypic.

other common names

English: Napoleon's peacock-pheasant; French: éperonnier napoléon; German: Napoleonfasan; Spanish: Espolonero de Palawan.

physical characteristics

Male 20 in (50 cm); female 16 in (40 cm); male 1.0 lb (0.4 kg); female 0.7 lb (0.3 kg). Male has a long, pointed crest on head, solid black underparts, shiny blue and green on mantle, and a distinctive black and white face pattern. Female is brown with buff markings.

distribution

Palawan Island, southwestern Philippines.

habitat

Previously typical of lowland coastal forest below 2,000 ft (600m); this now mostly logged and species found inhabiting montane forest and bamboo scrub at 4,900 ft (1,500 m) in 2000.

behavior

No information available.

feeding ecology and diet

No information available.

reproductive biology

Males call and keep small areas clear of leaves, for use as courtship display grounds; clutch size two; incubation 18–20 days.

conservation status

Vulnerable and on CITES Appendix I, prohibiting trade in wild birds.

significance to humans

None known.

Resources

Books

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.

Periodicals

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.

Organizations

Game Conservancy Trust. Fordingbridge, Hampshire SP6 1EF United Kingdom. Phone: +44 1425 652381. Fax: +44 1425651026. E-mail: [email protected] Web site: <http://www.gct.org.uk>

Ruffed Grouse Society. 451 McCormick Rd, Coraopolis, PA 15108. Phone: (888) 564-6747. Fax: (412) 262-9207. Web site: <http://www.ruffedgrousesociety.org>

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: [email protected] Web site: <http://www.pheasant.org.uk>

Peter Jeffery Garson, DPhil