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Broad-Scale Population Declines in Four
Species of North American Quail: An
Examination of Possible Causes
Leonard A. Brennan 1
Abstract - Christmas Bird Count (CBC) data from 1960-1989 indicate that
California quail (Cal/ipepla califomica) , northern bobwhite (Colinus
virginian us) , and scaled quail (Callipepla squamata) populations have
experienced significant declines in major portions of their geographic ranges.
Additionally, surveys and hunter bag returns during the past 50 years
indicate that mountain quail (Oreortyx pictus) populations have eXperienced
a series of local extinctions across broad areas (several thousand km2) in
Idaho and Nevada. Although changing land uses can be related to these
declines, no single factor can be linked to all species. For northern
bobwhites, clean farming methods in agricultural environments and
intensive, high-density pine-dominated silviculture seem to be the two major
reasons for broad-scale population declines, especially in the southeastern
states.
For mountain quail, regional extinctions in Idaho and Nevada are apparently
related to two factors: (1) intensive agriculture and associated hydro-power
reservoir impoundments along the Snake River corridor, and (2) disruption
of key habitat resources along secondary riparian corridors by excessive
cattle grazing. Factors responsible for declines in California quail and scaled
quail populations are at present unknown, but are apparently related to
abuses associated with excessive grazing of western rangelands.
Management strategies that can be used to sustain quail populations in
wildland environments are summarized in an ecological context.
INTRODUCTION
agriculture and forestIy have, however, called into question what
were once symbiotic relationships between people, quail,
farming and forestIy.
During the past decade, reports indicated that northern
bobwhite populations (Colinus virginian us) had declined at
many locations (Rosebeny and Klimstra 1984, Droege and
Sauer 1990). TIlls downward trend of one of the most common
and widely distributed game birds in North America surprised
many people. Further analyses revealed that northern bobwhites
had indeed declined on both continental, regional and statewide
scales (Brennan 1991, Brennan and Jacobson 1992).
The extent and magnitude of the bobwhite decline resulted
in a Strategic Planning Workshop for Quail Management and
Research in the United States that was held at the Third National
Quail Symposium in 1992 (Brennan 1993a, 1993b). This
woIkshop was the ftrst attempt to develop a comprehensive
Historically, populations of New World quail
(Odontophorinae) have been considered a sustainable by-product
of many agricultural and silvicultural activities (Stoddard 1931,
Leopold 1933, Rosene 1969, Leopold et al. 1981).
Abundant quail populations in rural and wildland
environments improved the quality of life for people by
providing recreational opportunities, economic returns from
leasing lands for hunting, and other positive social values that
resulted from a consumptive connection with wild vertebrate
resources (Leopold 1933). Changing patterns of land use in
1 Leonard A. Brennan is Director of Research, Tall Timbers
Research Station, located in Tallahassee, FL.
44
Independent analyses of Breeding Bird SUlVey (BBS) data
collected by the U.S. Fish and Wildlife SelVice (Sauer et aI.
1993) corroborated the patterns shown by the CBC data.
Although the mountain quail (Oreortyx pictus) showed no
evidence of decline based on CBC data (Figure 1), game
biologist sUlVeys, hunter bag returns, and comprehensive field
SUlVeyS have indicated that this species has undergone nearly a
statewide, regional extinction in Idaho (Figure 2) and Nevada
(Brennan 1993a).
strategy for quail management and research in the United States.
It followed a regional strategic planning effort for upland game
birds that was developed for western states by the Bureau of
Land Management (Sands and Smurthwaite 1992).
My objectives in this paper are to (1) summarize long-tenn
trends of quail populations at the continental scale in the United
States and evaluate evidence of declines, (2) identify real and
possible causes for obselVed declines and geographic range
contractions, and (3) summarize strategies for management and
research that might be used to sustain quail populations in an
ecological context. My overall purpose is to use quail
populations as an example of what .happens when relationships
between seemingly abundant vertebrate populations and land use
practices are taken for granted.
Hopefully, these case histories will raise awareness of
problems facing this unique, and often overlooked group of
native avifauna
POSSIBLE ICAUSES OF POPULATION
DECLINES
Characteristics of Declining Quail Populations
Populations decline when rates of birth and/or immigration
are less than rates of death and/or emigration (Begon and
Mortimer 1986). With respect to species such as quail, normal
annual mortality rates can be as high as 80-90% (Rosene 1969,
Leopold 1977, Roseberry and Klimstra 1984). Throughout
evolutionmy time scales, quail have evolved characteristics such
as large clutch sizes (Leopold et al. 1981) and indeterminate
egg-laying (Welty 1975) which selVe as reproductive strategies
that can potentially offset such high mortality rates. However,
the high percentage of annual turnover that most quail
populations experience means that when habitat components, or
other key resources needed for survival are eliminated,
populations can decline and disappear at an extremely rapid rate.
EVIDENCE OF DECLINES
Brennan (1993a) summarized population trends for 6 species
of quail in the United States from 1960-1989 based on Christmas
Bird Count (CBC) data.
Three of the 6 species of quail in this study (California quail,
Callipepla squamata~ northern bobwhite~ and scaled quail, C.
squamata) showed statistically significant evidence of declines
(Figure 1). None of the species in this study showed evidence
of increasing populations (Figure 1).
5~--------
A.
4
__________
CALIFORNIA QUAIL
r- -0.43, P- 0.01
~
3.5
0.4
C.
3
0.3
2.5
a:
::>
3
2
2
1.5
0.2
oJ:1
0.1
~
a: o~·~~~~~~~~
~
a:
w
a..
-I
«
::>
0
MONTEZUMA QUAil
r.. -0.09, p .. 0.60
60 62 64 6668 70 72 74 76 78 80 82 84 86 88
0.3
D.
MOUNTAIN QUAIL
r- -0.15, P- 0.94
0.2
o~~~~~~~~~~~
Q ...
60 62 &4 66 68 70 72 74 76 78 80 82 84 86 88
0.25
GAM BEL'S QUAIL
r- 0.22, P- 0.24
0.5
E.
0.8
60 62 64 66 68 70 72 74 76 78 80 82 84 86 88
F.
NORTHERN BOBWHITE
r- -0.85, P< 0.001
SCALED QUAil
r- -0.6, P< 0.001
0.6
0.15
2
0.4
0.1
0.2
0.05
o~~~~~~~~~~
60 62 64 66 68 70 72 74 76 78 80 82 84 86 88
o~~~~~~~~~~~
60 82 64 66 68 70 72 74 76 78
eo 82
84 86 88
o~~~~~~~~~~~
60 62 64 66 68 70 72 74 76 78 80 82 84 86 88
YEAR
Figure 1. - Quail population trends in the United States based on 31 years of Christmas Bird Count data. Statistics are correlation
coefficients (r) and probability that the slope of the regreSSion line is significantly different from zero. Data from Brennan (1993a),
45
1938
1965
1989
Figure 2. - Changes in the geographic distribution of mountain quail in Idaho during the past 50 years. 1938 map based on data from
Murray (1938). 1965 map based on Ormiston (1966). 1989 map compiled by Idaho Fish and Game Department and other surveys.
challenged interpretations made about the lack of widespread,
direct antagonistic relationships between fue ants and quail
(Brennan 1991), they have yet to present experimental or
circumstantial evidence that the presence of fire ants limits quail
population productivity.
Issues such as fue ants, coyotes, global warming, and other
such potential epiphenomena are, in many ways, red herrings
that threaten to steer us off the trnck of the real problems that
are at the root of the bobwhite decline (Brennan 1993c). These
problems relate to changing land use in agriculture and forestry,
and in the ever-increasing utbanization that eliminates bobwhite
habitat, and/or erodes its quality on a broad scale.
With bobwhites, changing land uses have clearly had a broad
and largely negative impact on populations (Klimstra 1982,
Brennan 1991). In agriculture, the herbicides may indirectly
reduce or eliminate arthropod resources needed by growing
chicks. Elimination of native weedy plants which provide
substrntes that produce abundant insects has broad, negative
impacts on partridge (potts 1986). This relationship may vety
well hold true for bobwhites and other quail, but it needs to be
tested. In forestty, the widespread proliferation of high-density
pine plantations, and reduction in use of prescribed fire has
eliminated hundreds of thousands of acres of old-field habitats
that once produced quail (Brennan 1991). In rural social
contexts, the collapse of the tenant farming system in the
southern U.S. and a broad-scale move from an agrarian to a
setVice-based economy (Winter 1988, Bradshaw and Blakeley
1982) has apparently had devastating effects on quail (Brennan
1991).
The linkage between declining bobwhite populations and
changing land use becomes clear when local case histories are
examined in light of good quail management and habitat is either
improved or maintained. For example, case histories in
Mississippi (Brennan et al. 1991, Brennan 1992a, Brennan
1993c, Brennan 1993d) point to a dramatic increase in bobwhite
numbers when habitat conditions are improved, but other effects
If habitat or other limiting conditions do not become favorable
within a relatively short time period, local and regional
extinctions can occur. Where a limiting factor is abiotic (such
as water from sporndic annual rninfall in the Rio Grnnde Valley
of Texas), annual bobwhite population fluctuations can be
dramatic (Lehmann 1984). In this situation, biotic habitat
components remain relatively stable, and alternating wet and city
periods, which often persist across multiple years, are the
primaty cause of the fluctuations. However, when biotic habitat
components are degrnded through changing land use, application
of agrochemicals, or other factors, populations of small
galliformes such as quail or partridge (Perdix perdix) often
decline and disappear quickly and thus undergo local or regional
extinctions. With quail, such extinction processes may occur so
quickly and at such a broad scale that recovery in a
metapopulation context (Hanski 1991, Rolstad 1991) may not
be possible.
Northern Bobwhite
Numerous factors have been attributed as being responsible
for the broad-scale declines that northern bobwhites have
experienced during the past 30 years. These factors range from
the geogrnphic expansion of the coyote (CaniS latrans) in the
south, to broad-scale increases in hawk and owl populations, to
the invasion of the imported fue ant (Solenopsis spp.).
Experimental evidence linking factors such as these to
bobwhite declines does not exist. In some situations,
circumstantial evidence of rnptor predation may be compelling
in the absence of changing land use and lack of agrochemicals.
However, linking factors such as coyotes and fire ants to the
broad-scale bobwhite decline are myths that must be eliminated
through education (Brennan 1991). Study of coyote foods in the
southeast indicated that bobwhites are the least-common dietaIy
item of coyotes (Wagner 1993). Although Allen et al. (1993)
46
likely to change. Modifying the way cattle are managed is clearly
the most significant opportunity for restoration of this quail in
portions of its fonner range in Idaho.
(such as predators and fire ants) are kept constant. Conversely,
when habitat conditions are allowed to erode, bobwhite numbers
will decline concomitantly (Dimmick 1992). Furthennore, the
vast area (200,000 hal of private lands managed for bobwhites
in the Red Hills region of southern Georgia and northern Florida
continues to produce abtmdant quail populations at the same
time bobwhite numbers continue to decline elsewhere in the
southeastern coastal plain. The linkage between land use and
bobwhites is an issue that has been raised on a regular basis for
over 60 years (Stoddard 1931, Rosene 1969, Rosebeny and
Klimstra 1984), yet, often seems to be neglected in favor of
some other more easily identifiable villain such as predators
(Mueller 1989) or fire ants (Allen et al. 1993).
Scaled Quail
Evidence of the scaled quail decline swprised many of the
participants at the Third National Quail Symposium last year.
Mechanisms responsible for the decline in scaled quail
populations are not asl well understood as the factors behind the
northern bobwhite and mountain quail declines. There are,
however, some potential relationships between excessive grazing
and this decline that should be explored. Scaled quail clearly
have an affinity for desert grasslands with sparsely scattered
shrubs (Schemnitz 1961, Brown 1989). Homogenous grasslands
without a shrub component are usually unsuitable for scaled
quail (Schemnitz 1961). Excessive grazing by cattle removes or
reduces grasses and forbs and tends to result in an increase in
woody and shrub vegetation Good range stewardship that
allows residual grasses and forbs to persist through the winter
results in lower scaled quail mortality and an increase in local
populations (Brown 1989). Whether this is the
across broad
portions of this birds' range and whether such a management
strategy can be used to sustain scaled quail populations remains
to be tested.
Mountain Quail
Mountain quail clearly represent a classic example of how
quail populations can be sustaine<L.or eliminated as a function
of land use. In the montaine areas of northern California,
mountain quail populations are apparently stable, and can persist
at densities of up to 30 birds per 100 ha (Brennan and Block
1986). Conversely, populations have undergone broad regional
and local extinctions in Idaho as a result of key wintering and
breeding habitats being eliminated as a result of anthropogenic
changes to key aspect of their habitat.
In contrast to California where extensive areas of chaparml
vegetation provide good~uality mountain quail habitat across
large regions (Brennan et al. 1987), the local restriction of
mountain quail to linear arrangements of creekside and riparian
brush communities in Idaho has apparently made them
vulnerable to elimination of wintering habitat from
hydro-electric dams along the Snake River conidor and its
tributaries. When mountain quail migrate from high elevation
breeding habitats to low elevation wintering habitats, they can
encounter a variety of risks, not the least of which are reservoirs
that eliminate vast areas of wintering habitat. Furthennore,
excessive grazing simplifies the floristic composition of the
creekside brush communities on which these birds rely, and
decreases their suitability as mountain quail habitat (Brennan
1992a).
We can also gain insight into factors that limit mountain quail
in Idaho by looking at the characteristics of the places where
they continue to persist. The remnant populations that are
apparently self-sustaining are located in steep, isolated portions
of the Snake and Salmon River Canyons in areas inaccessible
to cattle. Although this infonnation is circumstantial and not
experimental, it provides strong inferential evidence that
rangeland abuses from grazing may be responsible, at least in
part, for the declines mountain quail have experienced in Idaho.
It also offers evidence of an opportunity for improving habitat
for this species by practicing good rangeland stewardship. The
hydroelectric impoundments and intensive agriculture are clearly
established fixtures along the Snake River corridor, and are not
case
California Quail
Leopold (1977) provides a comprehensive overview of
California quail biology and ecology in the context of land use.
Although grazing can be used to improve the conditions of some
environments for California quail, abuses of this practice can
seriously degrade the quality of the habitat for this species.
Leopold (1977:158) states "To increase usefulness of brush
stands for quail, there must be cover at the ground level as well
as overhead...often the most effective way to achieve this end
is to exclude livestock from portions of the brush." Brush
conversion projects and other so-called rangeland improvements
have been known to have deleterious effects on California quail
for years, and caused Leopold (1977:160) to state, "I am
increasingly distressed at the progressive 'cleaning up' of field
borders ... [in] modem, slick, mechanical fanning." Fifteen years
later, these trends continue, and so too does the erosion of
California quail numbers.
The widespread human population increase in California
during recent years is not apparently directly responsible for
broad-scale declines in California quail. CBC count circles from
urban and suburban locations did not show a significantly greater
than expected number of declines in California quail.
47
1983). The recent uproar at the proposal to lower subsidies for
public land grazing fees that encourage overgrazing and
associated abuses and link grazing fees on public lands with fair
matket values is a classic example of this recalcitrant attitude.
Whether these complex, wicked problems (Allen and Gould
1986) of public land management can be solved remains to be
STRATEGIES FOR SUSTAINING QUAIL
IN AN ECOSYSTEM CONTEXT
We need to begin with the modest assumptions that (1) quail
are renewable resources, and (2) they can be sustained in the
context of contemporary land use prnctices. Emerging trends in
agriculture (Robinson 1990) and forest management (Sharitz et
al. 1992) indicate that there is some promise and hope for
stopping the broad-scale declines that many quail populations
have been experiencing. Howevet; whether the mainstream
managers in foresny and agriculture adopt these philosophical
changes remains to be seen .
In agriculture, the direct and indirect roles of agrochemicals
with respect to quail (especially northern bobwhite) need to be
assessed. The ConselVation Headlands approach to partridge
management in agricultural environments in England (potts
1986) appears to have profound implications for integrating
northern bobwhites in modem production agriculture. This
approach entails reduction of Iilemicide application around field
perimeters so that weedy foms and phytophagus insects can
grow and provide food resources for growing partridge chicks.
In foresny, considerntion needs to be given to uneven-aged
management strntegies that emphasize long rotation and single
tree selection. Such foresny prnctices, when combined with
frequent, annual burning, have sustained abundant huntable
populations of northern bobwhites in the Red Hills plantation
counny of southern Georgia and northern Florida for over 60
years. Such land use prnctices can clearly selVe as a model for
habitat management in other parts of the northern bobwhite's
range, especially on public lands where multiple uses are
mandated. Another such model is the relationship between the
endangered red-cockaded woodpecker (Picoides borealis) and
northern bobwhites in pine forests of the southeastern coastal
plain (Brennan and Fuller 1993). Brennan (1991) and Brennan
et al. (1993) obselVed a significant, positive response of northern
bobwhites to habitat management for the red-cockaded
woodpecker at Noxubee National Wildlife Refuge in east-central
Mississippi. Conversely, the private plantations that have been
managed for bobwhites in the Red Hills region of Georgia and
Florida support the largest extant population of red-cockaded
woodpeckers on private lands (T. Engstrom, personal
communication, Tall Tunbers Research Station). In northern
California, Block et al. (1991) obselVed that mountain quail were
loosely affiliated with a guild of approximately 8 species of
birds that shared an affinity for brushy and chaparral-dominated
vegetation Identifying similar linkages that establish positive
relationships between management for species of quail and other
terrestrial vertebrntes (or vice versa) is clearly needed.
In contrast to some of the recent potentially positive
conceptual developments for integrating quail with other wildlife
resources in forest and agricultural environments, similar
relationships in rangeland environments have apparently not
been established. Range managers seem to be uncooperative
when it comes to implementing comprehensive stewardship and
adopting a pay-as-you-go philosophy (Ferguson and Ferguson
seen.
The fate of all quail, and many other vertebrates as well, are
clearly linked to the ways that we farm our land, graze our grass,
and manage our forests. Focusing on strntegies that maintain the
integrity and functional processes of ecosystems (Regier 1993)
would clearly be the most effective way to sustain populations
of wild quail. Maintaining system integrity with an ecosystem
approach allows managers the opportunity to provide for the
annual cycle needs of the birds.
Consider the alternatives. There have been vast amounts of
resources poured into recovety efforts aimed at the endangered
masked bobwhite (Co!inus virginian us ridgwayi). Recovety
efforts were continually met with failure until a large tract of
land (Buenos Aires Ranch) was purchased and managed as a
refuge (Brown 1989). Even today, quantitative descriptions of
masked bobwhite habitat components are not available, and
habitat management on Buenos Aires is largely based on the
"best guess" approach, because reliable information has not
been compiled (W. Kuvlesky, personal communication, Buenos
Aires National Wildlife Refuge).
The masked bobwhite, lesser prairie chicken (Tympanuchus
pallidicinctus) and other once common game birds have been
driven to the brink of extinction by changing land use practices.
If contemporary trends in land use continue, and an ecosystem
approach to sustaining quail and other wildlife resources is
ignored, then we will most likely add other species of once
common gallifonnes to this list. To the naysayers who doubt
that birds as common as quail can be potential candidates for
extinction, I offer the example of the passenger pigeon
I
(Ectopistes migratorious).
ACKNOWLEDGMENTS
The ideas in this paper developed through the course of my
work on habitat and population ecology of quail that I began as
a graduate student in 1982. R.J. Gutierrez of Humboldt State
University was instrumental in providing academic opportunities
that allowed me to develop many of these ideas. Support for
work on mountain quail in California and Idaho was provided
by the California Department of Forestry, the now-defunct
International Quail Foundation, and the still extant Chukar
Foundation Many of my ideas about problems facing northern
bobwhites evolved from 1990-1993 when I was supported by
the Mississippi Department of Wildlife, Fisheries and Parks, and
worked in the Department of Wildlife and Fisheries at
Mississippi State University. George Hurst deselVes exceptional
credit for helping me see in a single afternoon how 20 years of
plant succession can influence bobwhites. I thank Teresa Pruden
48
woodpecker habitat management on non-target forest
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Nacogdoches, TX. (in press).
Brennan, L. A. and R. S. Fuller 1993, Bobwhites and
red-cockaded woodpeckers: endangered species management
helps quail too! Quail Unlimited Magazine 12(3): 16-20.
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need for proactive lmanagement. Giber Faune Sauvage 9(4).
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W. Strickland. 1991. Bobwhite quail management at the
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for reading an earlier draft of this paper and helping me improve
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me to attend this symposium, and Bill Block for inviting me to
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