California Quail
Callipepla
californica
A very readable
book that still stands as the definitive work on California
quail natural history: A. Starker
Leopold. 1977. The California Quail. University of
California Press, Berkeley, CA. The 1985
reprint is available online at the Barnes & Noble and
Amazon.com.
Below are some nontechnical articles.
California Quail
By Lee Franks
The California
Quail, Callipepla
californica, is the state
bird of California and the subject of A. Starker
Leopold’s 1977 book, The California Quail, which
combines nearly a century’s worth of published and
unpublished research into a single text.
This attractive game bird inhabits broken scrubby habitat
(perennial shrubs broken by spaces with annuals) where it
has access to cover and to annual food species, mainly
legumes (members of the pea family). Quails also like fruit
and seeds from buckbrush and poison oak. They live in
coveys, or flocks, that move widely throughout the Park
during the non-breeding season. During the breeding season
the covey breaks up and individual pairs spread across the
covey range to nest and raise their young.
The California Quail is distinguished from other quail
species by its unique plumage pattern and the presence of a
forward facing comma-shaped black plume that makes them
look like a flapper from the 1920’s. The adult male
has a boldly patterned black-and-white face with a
buffy-yellow forehead, gray breast, black
“scaling” on the belly, and a chestnut patch at
the center of the belly. The adult female is similar but
duller and browner overall, with markings on the side of
the neck and upper back dark brown instead of black. The
head is entirely grayish, without black and white markings.
Behavior
This is a highly gregarious species, moving around in
coveys that average 20-25 birds. They tend to run rather
than fly, but will fly short distances to avoid predators.
The birds usually depart from night roosting sites
(generally off ground in oaks and laurel) between first
light and sunrise to forage. The covey departure is
initiated by Assembly Calls. During foraging one individual
(usually male) often acts as sentinel, sitting on a high
perch and giving Contact and Aerial Alarm Calls when it
observes danger.
Adults eat seeds, leaves, and flowers from grasses, shrubs,
and trees. They also will consume berries and small amounts
of insect food, especially when there is a limited amount
of water in their habitat. They seem to require nearby
cover from perennial plants while foraging on annual
vegetation. Foraging is primarily on the ground, although
they will occasionally climb trees and pull off berries and
flowers. During foraging bouts, the covey stays together
through Contact Calls.
Coveys break down during the breeding season as
intra-sexual aggressive behavior increases. Pair bonds
generally form between birds from the same covey.
Approximately 2 months elapse between covey breakup in
March and complete segregation of birds into pair bonds. If
both individuals of a pair survive until the next year,
they show a tendency to re-mate. Older birds generally mate
earlier than younger birds, and adult females generally
mate with adult males rather than yearling males. The
primary manifestation of courtship by both sexes is
courtship feeding.
Breeding
Females lead in the selection of a nest site and the
building of nests. The nest is on the ground and well
concealed, often in dry grass, weeds, and dead brush. Hens
make the nest by lining a protected depression in the
ground with grass and weed stems. Egg laying generally
occurs in late April or early May. Females lay 3 eggs every
4 days. Average clutch size ranges from 11 to 17 eggs.
Incubation starts after the entire clutch is laid, and
lasts for 22 to 23 days. The female does all of the
incubating. The male acts as sentinel while his mate
incubates.
Young birds hatch with eyes open, covered with down, and
capable of moving around on their own. They usually leave
the nest within 2 days, and trail after their parents who
show them how to find food. For the first two weeks,
however, the chicks are not capable of adequate temperature
regulation, and the female broods them at night and in
early morning to prevent chilling and overheating. A
brooding female gathers chicks under her and fluffs her
feathers over them.
The rate of quail reproduction is closely related to the
amount of rainfall. Those years with enough rain to produce
spectacular displays of wild flowers also tend to be good
years for the reproduction of the quail. The rain seems to
regulate the breeding of the quail by influencing the
chemistry of the plants that they eat.
Food
Habits
The best foraging habitat occurs in broken brush. During
the first few weeks of life, chicks are vulnerable to
predation and forage close to cover. Adults will forage at
distances of 100 meters from cover in the absence of aerial
predators. This distance will shrink to 15 meters under
pressure from raptors. Cooper’s hawks, however, are
known to hunt quail by their calls.
Feeding techniques include scratching for seeds, jumping
for flowers and buds, pecking at ground, and shelling of
acorns. During the non-breeding season, quail feed twice a
day, in the morning just after dawn and again in late
afternoon. During storms they feed sporadically throughout
the day during breaks in the storm. If frightened by a
Cooper’s hawk, they may forgo the second feeding.
Sounds
Quail do not sing, but they have a wide array of calls that
they use when alarmed, aggressive, advertising, maintaining
contact with others, and assembling the covey. It has 3
syllables, with emphasis on the second syllable
(cu-CA-cow), and is given when an individual is separated
from the group or a mate, and before and during collective
covey movement. The Assembly Call is usually loud and may
be repeated 10 or more times.
Unmated males give the Advertisement (of courtship desires)
Call early in the breeding season. This is a single
syllable, (cow) , similar to the final note in the Assembly
Call, but with a longer duration. It is given from a high
position, where the calling male stands erect, head
elevated and thrown back at each note.
Males also give Aggressive Calls during the breeding
season. This call is a series of (squill) syllables given
with the head thrown back for each syllable. It is usually
given in encounters between males during the establishment
of dominance relationships. Males have dominance
hierarchies which function in mate selection, inter-covey
social relationships, and the movement of broods. All adult
males and some immature males participate. Only individuals
in the hierarchy acquire mates. Dominant birds call more
often than subordinate birds. In general, subordinate males
do not call after the dominance relationship is
established.
References
The Birds of North America; Calkins, J.D., Hagelin, J.C.,
Lott, D.F.: No. 473, 1999.
The Birder’s Handbook; Ehrlich, P.R., Dobkin, D.S.,
Wheye, D.
The Biogeography of California Quail
by J. McIlvaine
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.
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: 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.
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.
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).
Bibliography
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1994. Handbook of the Birds of the World. Vol. 2. New World
Vultures to Guineafowl. Barcelona, Spain: Lynx Edicions.
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unburned central California foothill rangeland. California
Fish and Game 54 (2): 123-127.
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food of California quail in dry years. California Fish and
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