New World quails

(Odontophoridae)

Class Aves

Order Galliformes

Suborder Phasiani

Family Odontophoridae

Thumbnail description
Plump, medium-sized birds with short, powerful wings and strong running and scratching legs and feet; all have a characteristic "toothed" bill and lack tarsal spurs in all species

Size
7–15 in (17–37 cm); 4–16 oz (125–465 g)

Number of genera, species
9 genera; 32 species

Habitat
Forest, woodlands, savanna, grasslands, and agricultural

Conservation status
Critically Endangered: 1 species; Vulnerable: 4 species; Near Threatened: 3 species

Distribution
Southern Canada, eastern United States, west coast and southwestern United States, much of Mexico and Central America, northern South America through the Amazon basin and Andes south to Bolivia, and east coast of Brazil and Uruguay; North American species introduced widely to Argentina, Chile, parts of Europe, and New Zealand

Evolution and systematics

There have been various groupings of the New World quail based on morphological characteristics, generally combining them with the pheasants as a subfamily within Phasianidae. However, DNA evidence suggests that family status is warranted and that the New World quail are not particularly closely related to the pheasants or Old World quail. Earliest fossils of quail-like birds have been found in Saskatchewan, Canada dating back 37 million years. Evidence appears to be pointing toward a divergence of the New World quail in South America 35–63 million years ago.

Within Odontophoridae there are 32 recognized species, although this number is subject to change as the lesser studied Latin American species are more thoroughly researched. There are nine genera including four (Oreortyx, Philortyx, Dactylortyx, and Rhynchortyx) containing only one species each. The remaining genera, Dendrortyx, Callipepla, Colinus, Cyrtonyx, and Odontophorus have three, four, four, two, and 15 species, respectively. Even taxonomy within these genera has varied especially in the Callipepla and Colinus. In the past, Callipepla has been split into two genera, Callipepla and Lophortyx, but it is now generally agreed that these species represent only a single genus. Among the better studied species, there is still confusion regarding taxonomy. The northern bobwhite (Colinus virginanus) may represent a super-species that can be split into several species. There is some suggestion that the masked bobwhite (C.v. ridgwayi) should be considered a distinct species. The Odontophorus wood-quails, which contains the largest number of species, are made up of several complexes of species. For example, the rusty-breasted complex and the black-throated complex are both found in the northern Andes. These nine species may be grouped or split in the future as more research is completed.

Physical characteristics

The New World quail are smallish Galliformes and, like most members of the order, have plump bodies and short wings.

Odontorphoridae are much less variable in size compared to the other families of Galliformes ranging from the smallest bobwhites (Colinus spp.), barred quail (Philortyx fasciatus), and tawny-faced quail (Rhynchortyx cinctus)—reported to be as little as 7 in (17 cm) in length and 4 oz (125 g)—to the long-tailed wood-partridge (Dendrotyx macroura) which may be 16 in (37 cm) in length and weigh 16 oz (454 g). Plumage tends to be more subtle than many of the other Galliformes. Also, sexual dimorphism which tends to be dramatic in the Phasianidae is much less distinct. Often there are slight size and plumage coloration differences between males and females. Some species appear to have no distinct external differences between males and females. A number of species have distinct crests ranging from small tufts to very long plumes. Many species have distinct, often red, fleshy rings around the eyes.

The serrated edge of the bill is a distinct characteristic of this family. None of the New World quail have tarsal spurs, unlike many of their Old World counterparts. In some species the legs and feet are very thick and strong for digging. All others still have strong legs and feet for running and scratching.

Distribution

Outside of human introductions, the New World quail are restricted to North, Central, and South America. The greatest number of genera and species are found in the vicinity of southern Mexico and Guatemala with the number of species decreasing outwards north and south. The genus Odontophorus is found mainly in southern Central America and northern South America. Colinus is distributed from eastern United States and Canada through Central America to Colombia, Venezuela, and the Guianas. Callipepla is restricted to western United States and Mexico. Oreortyx is restricted to western United States and just a small part of Mexico. Dendrortyx is restricted to Mexico and the vicinity of Honduras and Guatemala. Philortyx is restricted to Mexico. Cyrtonyx is found in southwestern United States, and Mexico, through to western Guatemala. Dactylortyx is found in southern Mexico through Honduras. Rhynchortyx is found along the eastern coast of Honduras to Panama where it is then found on the west coast down to Ecuador.

The northern bobwhite and California quail (Callipepla californica) have been widely introduced by humans. For the bob-white this includes established populations in the northwestern United States and British Columbia, Puerto Rico, Cuba, Hawaii, New Zealand, Italy, and Germany. The California quail has been introduced to Chile, Argentina, New Zealand, Australia, and Hawaii.

Habitat

The New World quail are ground birds inhabiting a wide range of tropical, subtropical, and temperate ecosystems. The bobwhites inhabit grassland, savannas, rangeland, and agricultural lands, but are also considered woodland edge species. The Callipepla species are mainly scrub to desert inhabitants. The two Cyrtonyx species are found in open pine or oak woodlands, but also inhabit scrub. The three Dendrortyx species inhabit montane and cloud forests. Oreortyx is found in mixed forest, forest edge, and chaparral. Philortyx inhabits xeric scrub and farmland. The wood-quails (Odontophorus) are found in a variety of tropical, subtropical, and montane forests. Rhynchortyx

is restricted to lowland tropical forest. Finally, Dactylortyx is found in a variety of montane forests, but some populations are found in lowland scrub and woodland edge.

Behavior

The most notable behavior of the New World quail is the covey, which has been reported in almost all species. It is interesting to note that these were formerly thought of as family groups, but now covey membership is thought to be much more complex. In some of the Odontophorus wood-quails there is some suggestion that coveys are family groups including adults pairs and helpers from previous clutches. Most species are diurnal and spend most of their time on the ground. Tree roosting has been observed in a number of the forest species. None of the New World quail are true migrants, although some appear to be altitudal migrants in mountainous regions.

Although the vocal repertoire of the New World quail is rather limited, most use a variety of calls and whistles to communicate. The bobwhites probably have the largest number of calls with at least 19 distinct calls. The Dendrortyx wood-partridges will give loud hooting calls; whereas the wood-quails exhibit loud guttural choruses.

Feeding ecology and diet

Most of the New World quail are gleaners and scratchers for seeds. Most eat a variety of seeds including those from grasses, forbs, shrubs, and trees. Many species will move into agricultural lands and eat waste grain seeds. Commonly reported crops eaten by various species include corn, wheat, sorghum, peanuts, and black beans. Some of the tropical forest wood-quails have been observed digging for fleshy roots. The two species of Cyrtonyx feed extensively on tubers and bulbs, especially the tubers of wood sorel (Oxalis spp). When studied, the diet of chicks is comprised mainly of invertebrates, and as they age they become more granivorous.

Reproductive biology

Breeding biology is not well studied in most species. Generally the New World quail were thought to be monogamous. However, data is mounting to the contrary. The northern bobwhite quail is now thought to have a flexible mating system that includes monogamy, polygyny, polyandry, and promiscuity. New genetic techniques combined with field methodologies are now being applied to this species to clarify its mating system. Anecdotal evidence suggests that some of the wood-quails may have adult pairs with helpers at nests and rearing young.

The temperate and grassland adapted species have the largest clutch sizes. For example, clutch size of the California quail ranges up to 17. Clutch size tends to decrease among the more tropical and forest adapted species. Where described, clutch sizes of three to five have been reported for some of the wood-quails. Nests are usually constructed on the ground forming a small bowl. Many species cover the nest with vegetation and form an "igloo-like" structure with an opening at one end. Incubation period is quite variable, although not well described in many species. Reports among the New World quail range from 16 to 30 days. Chicks are precocial and are capable of leaving the nest within hours of hatching. They grow rapidly and are capable of flight in less than two weeks. Mortality rates of nests and chicks is reported to be quite high in the better studied temperate and grassland species. Nest failure rates of 40–80% are quite common. Chick mortality rates of 20–50% are often reported. These species are persistent nesters with up to four nesting attempts reported in a breeding season. Although not reported for the more tropical and forest adapted species, the mortality rates of nests and chicks is probably lower.

Conservation status

Conservation status of the New World quail varies widely among species. The more temperate and grassland adapted species of Colinus and Callipepla tend to be very common. Some of these species are increasing in population and distribution as a result of human activity. Many of the forest adapted species seem to tolerate some human impact on their habitat, therefore seem to be maintaining reasonable populations. Most of the species with a conservation status are forest wood-quails in the genus Odontophorus. One species, the gorgeted wood-quail (Odontophorus strophium), found in oak forests in Colombia is considered critically endangered because almost all of its mid-elevational habitat in the Central Andes has been destroyed. The conservation status of many of the Latin American species is tentative because of lack of significant research to assess their status. For example, the bearded wood-partridge (Dendrortyx barbatus) was considered critically endangered in 1995. However, subsequent surveys of its montane forest habitat in the vicinity of Veracruz, Mexico identified a number of small and disjunct populations. This species is now considered to be Vulnerable because of continued threat to the remnant forest patches it inhabits.

Significance to humans

Like many of the Galliformes the New World quail are important to humans. Some species such as the northern bob-white quail are among the most studied birds in the world. This species is widely hunted and contributes greatly to local economies in parts of the United States and Mexico. Management for hunting in some areas has a significant impact on land use, oftentimes reducing the negative impact of grazing and farming on other wildlife. It is also widely raised in captivity to be released for hunting, as well as for the restaurant market. Most species are hunted either for sport or subsistence. This is done sustainably for a few species; however, the impact of hunting is not known for most of the Latin American species. There are a few cases of crop depredation by some species.

Species accounts

List of Species

Bearded wood-partridge

Dendrortyx barbatus

taxonomy

Dendrortyx barbatus Gould, 1846, Jalapa, Veracruz, Mexico. Monotypic.

other common names

English: Bearded partridge, Bearded tree-quail; French: Colin barbu; German: Bartwachtel; Spanish: Colin barbudo, Chivizcoyo.

physical characteristics

This is one of the larger species ranging 9–13 in (22–32 cm) in length and estimated weights of males about 16 oz (459 g) and for females about 14 oz (405 g). This bird has a grayish head with a small crest. Most notable are the bright red eye-ring, bill, and legs. The body has an overall reddish brown coloration with a darker rump and wings.

distribution

Restricted distribution to 14 fragmented populations ranging from west-central Veracruz to extreme northeast Queretaro, Mexico.

habitat

It is found in montane pine-oak, cloud, and older second growth forests. It is also associated with forest edges, shade coffee, and some agricultural land. Most of the forest, especially in Veracruz, Mexico is highly fragmented. The recently discovered population in Queretaro inhabits the most remote and intact forests.

behavior

Little is known. They form coveys like most of the other quails and have quite raucous chorus calls in the morning.

feeding ecology and diet

Little information available, but is known to feed on a variety of seeds and fruits. Will eat domestic crops such as black beans. Captive birds readily eat beans, corn, bananas, and grapes.

reproductive biology

Little is known, but the breeding season is suspected to be April–June. Broods of five have been seen in the wild. Captive birds constructed nests in shallow depressions in the ground and lined them with palms.

conservation status

Until 1995 this species was considered Critically Endangered. Surveys undertaken in late 1990s resulted in discovery of additional populations in remnant forests. Now downgraded to Vulnerable. However, rapid human growth and encroaching agriculture further threaten this species.

significance to humans

Because of its limited distribution and population, it is not an important species to humans. However, even in the remaining populations birds are still hunted for sport, and trapped as cage birds and for food. Local farmers have poisoned them when they are found depredating black bean fields.

Venezuelan wood-quail

Odontophorus columbianus

taxonomy

Odontophorus columbianus Gould, 1850, Caracas, Venezuela. Monotypic.

other common names

French: Tocro du Venezuela; German: Venezuelawachtel; Spanish: Perdiz montañera.

physical characteristics

This is a medium sized species ranging 11–12 in (28–30 cm) in length and with estimated weight of males about 12 oz (343 g) and 11 oz (336 g) for females. This species has a low brownish crest and general body color of reddish brown. The throat is white with black streaks. The breast is covered by distinct white teardrops outlined in black. The bill is black and the legs dark gray.

distribution

Restricted distribution in remnant forests in the northern coastal mountains west of Caracas, Venezuela and the northern-most tip of the Andes in northwestern Venezuela. Found in Henri Pittier and San Esteban National Parks.

habitat

Found in montane subtropical forests at altitudes of 2,950–7,900 ft (900–2,400 m). A recent study found that foraging habitat was usually associated with areas containing high

numbers of monocots, high vertical foliage density and low frequency of palms.

behavior

Like most of the other New World quails this species is found in coveys which are thought to be family groups. Group size in ranges from two to five individuals with one survey reporting an average group size of 4.5. Coveys are quite vocal and loud during early morning choruses. This might be for territory establishment and maintenance. When feeding they make a quiet, "güp-güp" call. Although almost always observed on the ground even when roosting, they have been observed roosting on palm fronds above the ground.

feeding ecology and diet

Little information available, but they have are suspected to feed on seeds, fruits, insects, and worms. They typically forage as a group working through the litter on the forest floor. They have been observed digging along the edges of surface roots and feeding on fleshy bits of the root.

reproductive biology

Little is known and only one nest has been described. Breeding season appears to run from March to July which corresponds to the wet season. The single nest was found at the base of palms and contained a roof of vegetation. It contained a clutch of six eggs and was incubated for 30 days.

conservation status

Considered Near Threatened because it occurs in several large national parks, most notably Henri Pittier National Park. However, populations and distribution have probably declined significantly as forests outside of the parks have been converted to agriculture and urban uses. Limited research suggests that there might be a decline in the population in the vicinity of Rancho Grande in Pittier National Park due to increasing tourism and vehicle traffic.

significance to humans

Because of its limited distribution and population, it is not an important species to humans. It may be opportunistically harvested by subsistence hunters who are hunting in remnant forest patches, and in the vicinity of the national parks.

Northern bobwhite quail

Colinus virginianus

taxonomy

Tetrao virginianus Linnaeus, 1758, America (=South Carolina). Twenty-two subspecies.

other common names

English: Bobwhite, northern bobwhite; French: Colin de virginie; German: Baumwachtel, Virginiawachtel, Wachtel; Spanish: Codorniz; Colín de virginia.

physical characteristics

Among the smaller Galliformes ranging in length from 8 to 10 in (20–25 cm) and 4 to 8 oz (129–233 g) in weight. Largest birds are found in the northern part of the range and the smallest in southern Mexico. Females are slightly smaller than males. Quite variable male plumage usually contains a combination of black, gray, and white. Males of the virginianus group usually have a white throat, as do the birds in the graysoni and pectoralis groups. Graysoni males tend to be more rufous colored;

whereas the pectoralis males tend to look most like the virginianus group, except the black collar is wider and more streaky on breast. Males of the coyolcos group have a solid black head and throat.

In females, the head and throat are mostly buff. The black parts of the head tend to be more chestnut in color.

distribution

Widespread distribution ranging from southern New England west through southern Ontario, Canada to southeastern Minnesota. The distribution continues southward to Florida in the east and extreme eastern Wyoming, western Kansas, and Oklahoma southward to Mexico in the west. There is a disjunct population in Sonora, Mexico and formerly Arizona. Restoration efforts are underway in Arizona. In Mexico, this species is found in the northeast state of Tamaulipas southward to the Isthmus of Tehuantepec. In central Mexico it reaches further west and in the south reaches the Pacific Ocean. It is replaced by the black-throated bobwhite (C. v. nigrogularis) on the Yucatan Peninsula. Distribution just reaches Guatemala. Introduced populations are found on many Caribbean Islands, Washington, Oregon, Hawaii, British Columbia, New Zealand, Italy, and Germany.

habitat

Found in a variety of habitats as long as some type of early successional habitat is present. Most closely associated with fire maintained pine savannas and forest openings or clearcuts in typically forested areas. Now most often found in farmland in those regions. Highest populations are found in grass and brush rangelands except where these are intensively managed for livestock. In the southeastern United States, best populations are found on "plantations" comprised of pine savanna ecosystems in southern Georgia and northern Florida. These areas have actually been maintained and managed for quail.

behavior

General behavior as in the family description. After the breeding season the birds go through an autumn "shuffle" to form larger coveys. This is time of greatest movement and some birds have been found to travel upwards of 60 mi (100 km). Home range sizes vary greatly. In better habitat might be 25–62 acres (10–25 ha) and much larger in poorer habitat.

feeding ecology and diet

Very well studied in this species. Shown to be primarily a seed eater with a wide range of seeds taken—a summary of studies identified 650 different types of seeds taken and 78 species that seemed to be more important. The types of seeds taken varies greatly with season and seems to be most related to seed abundance. Bobwhites will also take a variety of fruits and even large items such as oak (Quercus spp.) acorns. Chicks are primarily insectivorous, but begin consuming mainly seeds by six to eight weeks of age.

reproductive biology

The bobwhite quail was traditionally thought by lay people and scientists to form monogamous pairs with a great deal of parental care given by the males. More recent research is suggesting a much more complex social system where some individuals might be monogamous, but others are polygamous, including both polygyny and polyandry, and others appear to be promiscuous. Research is underway to try to clarify the bobwhite's mating system.

Unmated males give the "bob-white" call, which is a familiar spring call to anyone living within the distribution of this species. Pair formation is common, starting from January to March in the United States, with more northerly population beginning later. Like all the New World quails nests are built in a shallow depression on the ground, usually in dead grasses or other herbaceous vegetation. Clutch size averages 12–14 eggs, but ranges from seven to 28. Incubation is 23–24 days. The chicks are precocial and will leave the nest with the adults within hours of hatching. They can fly within two weeks. Mortality of nests, young, and adults is high. Hatching success ranges widely, but often from 20 to 40%. Hens are persistent renesters. Chick survival has been reported at 31% to one month of age. Annual survival is typically less than 30% and often less than 20%.

conservation status

The species is widespread and common. However, there have been significant declines in populations throughout the eastern United States. This is thought to be due to a combination of farmland abandonment and reforestation, loss of fire maintained pine savannas, and intensification of remaining agriculture. In many parts of the East, populations have declined by 70–90%, since the 1960s. Populations are less well know in Mexico. One subspecies, the masked bobwhite, which inhabits scrubland in Sonora, Mexico and formerly Arizona, is considered Endangered in the United States. Because of its importance as a harvested gamebird, there are management programs in several parts of its distribution to try to restore numbers.

significance to humans

This species is one of the most important Galliformes. It is widely hunted and reared for human consumption. Large amounts of money are spent for its conservation for hunting. In some areas of the Southern Plains management of rangeland has shifted to emphasizing bobwhite by using livestock because hunting has become more lucrative than raising livestock.

Resources

Books

Bonaccorso, Elisa. "Densidad y uso de habitat de perdíz Odontophorus columbianus en el Parque Nacional Henri Pittier." Trabajo Especial de Grado. Universidad Simón Bolívar Sartenejas.

Carroll, John P. "New World Quail." In Handbook of Birds of the World. Vol. 2. New World Vultures to Guineafowl, edited by Josep del Hoyo, Andrew Elliott, and Jordi Sargatal. Barcelona: BirdLife International and Lynx Edicions, 1994.

Johnsgard, Paul A. Quails, Partridges, and Francolins of the World. London: Oxford University Press, 1988.

Brennan, Leonard A. "Northern Bobwhite (Colinus virginanus)." In The Birds of North America, edited by Alan Poole and Frank Gill. No. 397. Philadelphia: The Birds of North America, Inc., 1999.

Fuller, Richard A., John P. Carroll, and Philip J.K. McGowan, eds. " Partridges, Quails, Francolins, Snowcocks, Guineafowl, and Turkeys. Status Survey and Conservation Action Plan 2000–2004." Gland, Switzerland; Cambridge, United Kingdom; and Reading, United Kingdom: WPA/BirdLife/SSC Partridge, Quail and Francolin Specialist Group, IUCN, and World Pheasant Association, 2000.

McGowan, Philip J.K., Simon D. Dowell, John P. Carroll, and Nicholas J. Aebischer, eds. "Partridges, Quails, Francolins, Snowcocks, and Guineafowl. Status Survey and Conservation Action Plan 1995–1999." Gland, Switzerland; Cambridge, United Kingdom; and Reading, United Kingdom: WPA/BirdLife/SSC Partridge, Quail and Francolin Specialist Group, IUCN, and World Pheasant Association, 1995.

Organizations

WPA/BirdLife/SSC Partridge, Quail, and Francolin Specialist Group. c/o World Pheasant Association, PO Box 5, Lower Basildon, Reading, RG8 9PF United Kingdom. Phone: +44 1 189 845 140. Fax: +118 9843369. E-mail: wpa@gn.apc.org Web site: <http:/www.pheasant.org.uk/>. PQF: <http://www.gct.org.uk/pqf/>

John Patrick Carroll, PhD