San Francisco State University
Department of Geography

Geography 316:  Biogeography

The Biogeography of California Quail (Callipepla californica)

by J. McIlvaine, student in Geography 316, Fall 2000


Quail Fig. 1: Male California quail in the Presidio, San Francisco (photo by J. McIlvaine)
 

Kingdom:  Animalia
Phylum:  Chordata
Class:  Aves
Order:  Galliformes
Family:  Odontophoridae
Genus:  Callipepla
Species:  Callipepla californica
 

Description of Species

The California quail, California’s state bird, is a 9-11 inch hen-like bird with a distinctive teardrop-shaped head plume called a top-knot. Their plump bodies vary from grayish to brown with scaly markings on the lower breast and abdomen. Males are particularly elegant with a black throat, chestnut patch on the belly, a bluish gray breast, white speckles on its flanks, and a white stripe on 
the forehead and around the neckline (Fig.1).

Females have a smaller top-knot and lack the male’s distinctive facial markings and black throat. 
Her crest is dark brown and her body is brown or gray with white speckles on the chest and belly. 
The marked sexual dimorphism is believed to play an important part in breeding displays. Juveniles resemble the female, but have shorter and lighter colored crests.

As ground dwelling birds, their short and powerful legs are well adapted for terrestrial locomotion. They can fly rapidly, but only for short distances. When alarmed they prefer to run, flying only as a last resort.

Habitat

California quail are best adapted to semiarid environments, ranging from sea level to 4000 feet and occasionally up to 8500 feet or higher (Sumner 1935). As long as there is abundant food, ground cover, and a dependable water source, quail are able to live in a variety of habitats including open woodlands, brushy foothills, desert washes, forest edge, chaparral, stream valleys, agricultural lands, and suburb areas. Cover is needed for roosting, resting, nesting, escaping from predators, and for protection from the weather (Sumner 1935, Leopold 1977).

Leopold (1977) separates California quail habitat areas into four major ecological zones (Fig. 2): arid ranges mostly in Southern California and Baja California, transitional ranges in the Sacramento Valley, humid forest ranges associated with the Coast and Cascade ranges, and interior Great Basin and Columbia Basin ranges. Of these the transitional ranges in the Sacramento Valley foothills provide the most stable quail habitat, characterized by mild winters, moderate rainfall, moderately dense ground vegetation, and generally adequate ground cover.


Quail Fig.2: Ecological zones of the California quail range. (Leopold 1977)

California quail are generalists and opportunists, so food intake varies by location and season. Their main food items are seeds produced by various species of broad-leafed annual plants, especially legumes. This includes plants such as lupine (Lupinus sp.), clover (Trifolium sp.), bur clover (Medicago sp.), and deer vetches (Lotus sp.) (Leopold 1977). Their bills are typical for seedeaters: serrated, short, stout, and slightly decurved.

Shields and Duncan (1966) studied California quail diet in the fall and winter during a dry year on the San Joaquin Experimental Range in the central Sierra Nevada foothills. They found that seeds comprised 82% of their diet, while green leafage contributed 18%. Duncan (1968) also studied quail diet in the same area and found that legume seeds were their most important food item. Quail also eat leafy materials, acorns, fruits and berries, crop residues, and some insects (Leopold 1977).

Natural History

During the fall and winter, California quail are highly gregarious birds, gathering into groups, called coveys. In most situations, covey size averages about 50 birds, but under intensive management and protection, coveys can get as large as 1000 birds (Leopold 1977). In the covey, the quail tend to imitate one another and exhibit cooperative behavior. For example, when one bird finds a good supply of food it often calls the others to it. Likewise, when a member of the covey perceives danger it will warn the group with the appropriate call (Sumner 1935). California quail communicate with 14 different calls (Leopold, 1977). This includes courtship, re-grouping, feeding, and warning calls. The most frequently heard location call has been described as “cu-ca-cow” or “chi-ca-go.”

At the start of nesting season in early spring the coveys break up, as quail pairs spread themselves out into different habitat areas to nest and rear their young. At the end of summer each new quail family rejoins the others to form a new covey where they will remain until the next breeding season.

Emlen (1939) observed this seasonal movement in his study of California quail on a 760-acre farm in the vicinity of Davis, California.  In the winter, four coveys, containing 21-46 birds, had home ranges of 17-45 acres, roughly one acre for each bird. The covey locations and range size depended on the amount of brush cover available. The four territories were separated by 350 yards to half a mile and contact between the coveys was infrequent. The members of a covey tended to feed and roost together in mid-winter, but occasionally they broke up into smaller units.  Winter movements were restricted with only 5 to 10 acres of an entire territory utilized by the covey on any one day. The same area would serve as a feeding ground for a few days to two or three weeks when the birds would move to another part of their territory.

The nesting season caused a major shift in the social organization and local distribution of the quail. Starting in late February the coveys began to break up into pairs and unattached males began to leave the covey, sometimes fighting to maintain territory in the vicinity of nesting pairs. Mated birds had rather small home ranges of only 12-25 acres prior to the start of nesting, and even smaller ranges of about 3-10 acres thereafter. In the fall, nine broods and 13 unattached stragglers merged into 4 new coveys.

California quail are monogamous, but usually pair up with new mates each spring (Genelly 1955). Females build their nests on the ground, well hidden under a bush or a brush pile. While the female feeds or constructs the nest, the male perches conspicuously above her where he can observe any potential threat to his mate. He stands motionless, sending out notes of either reassurance or warning (Sumner 1935).

Females lay 12-16 spotted cream-colored eggs and incubate them for 20-23 days and lay a second clutch on occasion (Johnsgard 1988). Once the chicks are hatched, both parents tend to the young. Chicks are precocious, feeding on their own shortly after hatching and the male acts as guardian while the young birds forage. The adult male tends to lose weight during this period, spending more time on the alert rather than feeding.

While most broods are reared by their parents alone, communal brooding in California quail populations has been observed. Lott and Sastrup (1999) conducted a study of California quail and found that 23 of 195 (12%) broods were reared communally. Parents of communal broods lived significantly longer (3.1 years) than parents of single broods (1.9 years) and hatched significantly more young during their lifetimes (36.3 vs. 15.7 young).

 The chicks grow rapidly, initially fledging at about two weeks of age and completing their juvenile plumage by about 11 weeks. By the age of 21 to 23 weeks all of the juvenile flight feathers except for the outer two are replaced by adult-like plumage (Raitt, 1961). Chicks are capable of short flights by the time they are a little over two weeks of age and are fully mature and capable of breeding at the age of ten months (Leopold 1977).

California quail are short lived with high mortality and high reproductive rates. The number of quail in a population is constantly undergoing change. The average rate of mortality is 74 percent. Mortality is highest in the first year of life. Only one bird in several thousand will live to be five years old. (Leopold 1977).

Evolution

California quail are part of a group of quail found only in the Americas called the New World quails. The Handbook of the Birds of the World (del Hoyo 1994) places the New World quails in their own family, Odontophoridae. The taxonomic status of the family has been debated for many years. They were more often considered part of the subfamily Phasianidae which includes the Old World quails, partridges, francolins, and pheasants (del Hoyo 1994). However, from DNA hybridization evidence it became surprisingly clear that New World quail are not closely related to Old World quail, turkeys or grouse (Sibley 1990). Sibley (1990) concluded that the Odontophoridae must be the descendants of an early divergence about 63 million years ago in South America during its isolation from North America.

The New World quail include nine genera with Dendrortyx (tree quails) as the earliest representative. Odontophorus (wood quails), Rhynchortyx (ex. Tawny-faced quail), Dactylortyx, and Cyrtonyx (ex. Montezuma quail) are genera predominantly adapted to the forest and are found in Central and South America. Colinus (bobwhite quails), Callipepla (ex. California quail), Oreortyx (ex. Mountain quail), and Philortyx (ex. Barred quail) are adapted to forest edge and are found primarily in North and Central America (del Hoyo 1994). Like the family taxonomic status, the genera of New World quail has also been debated over the years. Callipepla and Lophortyx have often been classified apart, but the differences between the two are considered too slight to be considered two genera. Instead the two forms are united in Callipepla.

Gutiérrez et al. (1983) through starch gel electrophoresis and fossil calibration, were able to determine that the earliest radiation of Oreortyx occurred 12.6 million years ago, followed by Colinus 7 million years ago, and most recently by Callipepla 2.8 million
years ago. They also suggest that Dendrortyx and Odontophorus diverged at least 16 million years ago (del Hoyo 1994).

According to Johnsgard (1973), the largest number of total quail species and the largest number of endemic quail species occur in Central and South America, whereas North America has the largest number of genera and endemic genera. The most primitive genera (Dentrortyx and Odontophyorus) are found in Mexico and further south, indicating the New World quail had their center of evolutionary history and speciation in tropical America.

Distribution

California quail have a continuous native distribution that extends roughly 1300 miles from Baja California to southern Oregon and about 300 miles from the California coast east into a small portion of western Nevada (Fig. 1). The species can be found in almost all of Baja California and California, except for high elevations of the Sierra Nevada and Cascade Mountains and in the Colorado and eastern Mojave Deserts (Leopold 1977).

While endemic in their native range, California quail have been widely transplanted throughout the West and now occur extensively in Oregon and Washington and are found throughout northern Nevada, western Idaho, and parts of British Columbia and Utah (Fig. 3). The species has also been successfully introduced to other parts of the world, including Hawaii, New Zealand, western Argentina, and central Chile (Leopold 1977).


Quail Fig. 3: Original and present range of California quail in North America. (Leopold 1977)
 

California quail are one of six species of native quail in the western United States. The others include Gambel’s (Callipepla gambeii), Scaled (Callipepla squamata), Bobwhite (Colinus virginianus), Mountain (Oreortyx pictus), and Montezuma quail (Cyrtonyx montesumae) (Fig. 4). California quail distribution overlaps with two of the species, Gambel’s and Mountain quail.


Quail Fig.4: Ranges of five species of western quail. (Leopold 1977)
 

Gambel’s quail are closely related to California quail and are so similar in size and coloration that they are often confused in the field. The native ranges of the two species are allopatric, slightly overlapping along their border in southern California. From the Panamint Range in Inyo County to the Santa Rosa Mountains in Riverside County, their populations exist together where they occasionally hybridize. Just to the east and west of this narrow border, however, the quail maintain distinct ranges. Johnsgard (1973) suggests that the Sierra Nevada Mountains provided an effective geographic barrier that allowed speciation to develop. Today, the major climatic differences on each of its slopes prevents extensive overlap of the two species.

While California and Gambel’s quail occur in two different geographic areas with only slight overlap, California and Mountain quail ranges overlap much more substantially. Like California quail, Mountain quail occur widely throughout California, from northern Baja California to Oregon, but the two species are geographically isolated for most of the year throughout their range. Mountain quail prefer conifer and oak forests, or dense chaparral, and can be found at elevations up to 10,000 feet (Scott 1995), while California quail are common in more open areas at low elevations. During the winter, however, a zone of sympatry occurs when snowfall forces Mountain quail to migrate to lower elevations (Gutiérrez 1980). In some areas, the two species are sympatric all year round. Gutiérrez (1980) studied one such area in Carmel Valley, California and found that the two species effectively partition their resources by utilizing different plant communities, allowing their populations to coexist with limited competition.

The California quail population has been divided into eight subspecies, occupying different parts of their range. Each subspecies has slight differences in coloration, size, and habit (Johnsgard 1973, del Hoyo 1994).

(1) C. c. californica, the Valley California quail, is the most widespread subspecies, found in most of central California extending from southern Oregon and western Nevada south to southern California and Los Coronados Islands of Baja California. They have been introduced into Oregon, eastern Washington, British Columbia, western Idaho, Utah and Colorado.

(2) C. c. orecta, the Great Basin California quail, occupies southeastern Oregon and extreme northeastern California.

(3) C. c. brunnescens, the Coast California quail, is found on the west slope of the California coast ranges from southern Santa Cruz County to Del Norte County and has been introduced to Vancouver Island in British Columbia.

(4) C. c. canfieldae, the Inyo California quail, inhabits the east slope of the southern Sierra Nevada Mountains of east central California.

(5) C. c. catalinensis, the Catalina Island California quail, is found on Santa Catalina Island in southern California and has been introduced on Santa Rosa and Santa Cruz Islands. This subspecies may have been introduced to Santa Catalina Island by Native Americans as a food source (Johnson 1972).

(6) C. c. plumbea, the San Quintín California quail, is found in southern California into northern Baja California.

(7) C. c. decoloratus, Grinnell California quail, occurs in central and southern Baja California.

(8) C. c. achrusterus, Cape California quail, can be found at the southern tip of Baja. This subspecies has also been introduced to Chile, Argentina, New Zealand, Australia, and Hawaii.

Other Interesting Issues

For Native Americans the California quail were highly esteemed as a source of food, and the “top knot” was used for decoration on clothing (del Hoyo 1994). European settlement, dramatic changes in the landscape, relocation, and interest in quails for sport and food have led to the distributions we see today. California quail are currently important in the business of hunting for sport and extensive management exists in some areas.

According to Johnsgard (1988) “with over two million California quail harvested each year by hunters in the United States, the population status of this species is clearly excellent, and does not warrant special attention.”  However, while the species is widespread and common, their populations have dwindled since 1960 (del Hoyo 1994). In San Francisco, the species is on the verge of becoming locally extinct.

In April 1999, California quail were included on the National Audubon Society’s list of threatened bird species (Martin, April 1999). In the summer of that year, in response to dwindling populations of California quail in San Francisco’s Golden Gate Park, the Golden Gate Audubon Society launched its “Save the Quail” campaign, spearheaded by its president at the time, Alan Hopkins. The main causes of the birds’ decline include habitat destruction and feral cats. With about 1,200 quail in the park at the turn of the century, the population declined to 12 in 1999. Hopkins estimated that the species would become extinct within the city in the next few years (Martin, July 1999).

The largest remaining population in San Francisco can be found in the Presidio and attempts are currently being made to ensure its survival. Eight road signs with the message “Quail in Area, Drive Carefully” have been installed and quail monitoring projects and habitat restoration activities are now underway. Saving the quail has become a citywide effort (St. John, June 2000). In July 2000, the California quail was designated San Francisco’s official city bird, illustrating the city’s committment to protecting and restoring quail habitat.


Quail Fig.5: Installing quail signs in the Presidio, San Francisco.
(Photo by J. McIlvaine)

Bibliography
del Hoyo, Josep, Andrew Elliot, and Jordi Sargatal, eds. 1994. Handbook of the Birds of the World. Vol. 2. New World Vultures to Guineafowl. Barcelona, Spain: Lynx Edicions.

Duncan, Don A. 1968. “Food of California quail on burned and unburned central California foothill rangeland.” California Fish and Game 54 (2): 123-127.

Duncan, Don A. and Paul W. Shields. 1966. “Fall and winter food of California quail in dry years.” California Fish and Game 52 (4): 275-282.

Emlen, John T. Jr. 1939. “Seasonal movements of a low-density valley quail population.” Journal of Wildlife Management 3 (2): 118-130.

Genelly, Richard E. 1955. “Annual cycle in a population of California quail.” The Condor 57: 263-285.

Gutiérrez, Ralph J. 1980. “Comparative ecology of the Mountain and California quail in the Carmel Valley, California.” Living Bird 18: 71-93.

Gutiérrez, R. J., R. M. Zink, and S.Y. Yang. 1983. “Genic variation, systematic, and biogeographic relationships of some galliform birds.” Auk 100: 33-47.

Johnsgard, P. 1988. The Quails, Partridges, and Francolins of the World. Oxford: Oxford University Press.

         1973. Grouse and Quail of North America. Lincoln, NE: University of Nebraska Press.

Johnson, N.K. 1972. “Origin and differentiation of the avifauna of the Channel Islands, California.” Condor 74: 295-315.

Leopold, A. Starker. 1977. The California Quail. Berkeley, CA: University of California Press.

Lott, Dale F. and Sonke N. A. Mastrup. 1999. “Facultative communal brood rearing in California quail.” The Condor 101: 678-681.

Martin, Glen. “A quail quandary: birds’ fans are trying to save the city’s falling population.” San Francisco Chronicle. 8 July 1999, sec.A, p.13.

         “Bird watch list released: group says California quail among threatened species.”  San Francisco Chronicle. 22 April 1999, sec. A, p. 17

Raitt, Ralph J., Jr. 1961. “Plumage development and molts of California quail.” The Condor 63: 294-303.

Scott, Shirley L. ed. 1995. Field Guide to the Birds of North America. Washington, D.C.: National Geographic Society.

Sibley, Charles G. and Jon E. Ahlquist. 1990. Phylogeny and Classification of Birds. New Haven, CT: Yale University.

St. John, Kelly. “Presidio takes quail under its wing: plans in place to help birds thrive.” San Francisco Chronicle. 29 June 2000, sec. A,  p.19.

Sumner, E.L. Jr. 1935. “A life history study of the California quail, with recommendations for its conservation and management. California Fish and Game 21: 167-253, 275-342.

Wilson, Yumi. “San Francisco Boardwatch”. San Francisco Chronicle 11 July 2000, sec. A,  p. 18.
 

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