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Cougar Management in North America: Canada

Authors:
  • Golder Associates Ltd
From the moment Europeans established the
original colonies that would become the United
States, predators were viewed as a threat, not only
to livestock but also to the settlers themselves and to the
other wild animals settlers relied on for food. Since then
three phases of cougar (Puma concolor) management have
evolved: attempted eradication, followed by agency man-
agement to sustain sport hunting and address depredation
concerns, and more recently an effort to sustain viable cou-
gar populations as part of the ecological community.
The fi rst phase of cougar management emerged as an
agricultural ethic that focused on eradicating “undesirable
species” that potentially threatened livestock, game animals,
or the settlers themselves. One of these undesirables was
the cougar. This attitude of eradication dominated until the
middle of the twentieth century, when states and provinces
began to assume management authority for the species, ini-
tiating the second phase of cougar management.
Along with this new authority to manage cougars came
the responsibility to sustain cougar populations. This phase
is characterized by the hunting of cougars to provide recre-
ational opportunity, while continuing to address livestock
concerns. Complexity of cougar management accelerated
rapidly from this point as a more biologically based ethic
began to develop within agencies and the public, and new
stakeholders entered the management arena. This increasing
complexity has forced some agencies into a third phase in
which they have begun to examine management approaches
more critically. State and provincial management plans
illustrate these evaluations by addressing cougar ecology
and sociopolitical aspects of cougar management. They
also incorporate stakeholder input and recently acquired
knowledge in order to develop management programs more
acceptable to the public at large. In the fi rst two sections of
this chapter, Charles Anderson Jr. and Frederick Lindzey
discuss the phases of cougar management in North America,
primarily in the United States. They describe the evolution
from eradication to supporting a sport harvest, addressing
cougar predation on livestock, and sustaining viable cougar
populations for ecological and recreational purposes. In the
third and fi nal section, Kyle Knopff, Martin Jalkotzy, and
Mark Boyce identify how management in Canada differs
from that in the United States.
Earliest bounties paid by settlements, colonial govern-
ments, and later fl edgling states were directed primarily at
controlling wolves (Canis lupus) . As wolf numbers declined
and density of human settlement increased, bounties were
also paid for other predators, such as cougars (Young and
Goldman 1944, 1946a). From colonial times to the 1960s,
the goal was to eradicate predators, a philosophy that moved
westward across North America with European settlement.
The result of eradication, along with habitat losses associ-
ated with human development, was that by the twentieth
century cougars were extirpated from North America east
of the Rocky Mountains, except for a remnant population
in Florida (Nowak 1976; see also Range Map, p. vii).
Although wolves were initially the focus of predator
control efforts in the West as they had been in the East,
other predators were taken incidentally during wolf control
campaigns. Later these species, including the cougar, became
prime targets as well. Livestock associations often hired hunt-
ers and trappers to kill cougars in areas with depredation
Chapter 4 Cougar Management
in North America
United States: Charles R. Anderson Jr.
and Frederick Lindzey
Canada: Kyle H. Knopff, Martin G. Jalkotzy,
and Mark S. Boyce
a species accompanied management authority. For example,
Wyoming statutes charge the Wyoming Game and Fish
Commission and Department with providing an adequate
and fl exible system for the control, propagation, manage-
ment, protection, and regulation of all Wyoming wildlife.
The department is the only entity of Wyoming state govern-
ment charged with managing wildlife and conserving it for
future generations (Wyoming Game and Fish Department
2006b, Department and Program-Level Strategic Plans).
Real or perceived livestock depredation and wildlife preda-
tion concerns did not go away simply because the cougar
was decreed a game animal. At last, however, the attitude of
some members of the public and of many professional biolo-
gists supported the notion of a balanced approach in which
“positive” as well as “negative” aspects of the species would
be considered in management planning.
Legal protection of the cougar as a game species sig-
naled the entry of new stakeholders attempting to infl uence
cougar management (Chapter 14, 15). Agencies faced the
unenviable task of trying to achieve management decisions
that would serve to maintain biological integrity while also
balancing the demands of confl icting interests. Approaches
and techniques learned from decades of managing other
game species were often of little use when dealing with
an obligate carnivore that occurred in comparatively low
densities and traveled over large areas. The general history
accounts (i.e., Young and Goldman 1946a) and the few
more specifi c reports available in the early 1960s on breed-
ing, food habits, and natural history (e.g., Connolly 1949;
Gashwiler and Robinette 1957; Robinette et al. 1959,
1961) provided limited support for, or help in, develop-
ing management plans in what was certain to be a hostile
management environment.
Regulation Begins
Typically, the fi rst years of wildlife agency responsibility for
cougars saw little more than the setting of bag limits. Hunt-
ing regulations then became progressively more restrictive
as season lengths were shortened and timing shifted. Man-
agement areas were delineated to control distribution of the
hunting harvest. In some areas, quotas were set to limit total
kill and /or the harvest of male and female cougars. Over
time, regulations protecting spotted juveniles and females
with kittens have also been adopted by most management
agencies (Table 4.1; management status reports in Becker
et al. 2003). At the same time, most states and provinces
continued to include liberal provisions for livestock owners
to respond to cougar depredation problems (Roberson and
Lindzey 1984, 207).
problems. Bounties were commonly used to reward such
hunters and direct their efforts to selected areas, a practice
that continued in the western states until the mid-twentieth
century. Young and Goldman (1946a, 166) found nine west-
ern states offering cougar bounties in 1937, with payments
ranging from $50 per animal in Colorado to $2 in Nebraska.
Poisons were widely used for predator eradication and likely
killed cougars as well as their canid targets. As noted by
Young and Goldman (1946, 167), young cougars may have
been most vulnerable to poisoning from consuming poisoned
carrion baits set for wolves and coyotes, Canis latrans. But
poisoning the carcasses of animals killed by cougars can be
effective in targeting the cats, because they commonly return
to feed on these carcasses (Anderson 1983).
Federal involvement began in 1907 with U.S. Forest Ser-
vice wolf reductions on national forests. The federal gov-
ernment formally entered the predator control business in
earnest in 1914 with passage of legislation providing money
for the Department of Agriculture to fund “experiments
and demonstrations in destroying wolves, prairie dogs
and other animals injurious to agriculture and animal hus-
bandry” (Young and Goldman 1946a, 383). The animal
damage control program within the Department of Agri-
culture’s Bureau of Biological Survey was later moved to
the Department of Interior, and then eventually returned to
Department of Agriculture (see Chapter 1 for fuller discus-
sion of predator control).
Bounties for cougars were generally phased out by the
early 1960s (Cougar Management Guidelines Working
Group 2005). Indiscriminate use of poisons on public lands
was terminated by presidential proclamation in 1972, and
most western states and provinces assumed management
authority for cougars between 1965 and 1973. Current
Environmental Protection Agency regulations prohibit the
use of poisons on public lands unless a memorandum of
understanding is developed with the federal land manage-
ment agency. In these cases, only target-specifi c methods
(M-44s or 10-80 collars) can be employed. Colorado and
Nevada reclassifi ed the cougar as a game animal in 1965;
Washington in 1966; Oregon and Utah in 1967; California
in 1969; Arizona, Montana, and New Mexico followed
suit in 1971; Idaho in 1972; Wyoming in 1973 (Rober-
son and Lindzey 1984, Smith 1989, Dawn 2002); North
Dakota in 1991 (with closed season until 2005); and South
Dakota in 2003. Except for Florida, the remaining states
and provinces have not implemented active cougar man-
agement programs because viable cougar populations are
not evident.
State and provincial agencies now faced the task of man-
aging an animal that had been persecuted for two centuries to
protect livestock and wild prey. Responsibility for cougars as
42 Charles R. Anderson Jr., Frederick Lindzey, Kyle H. Knopff, Martin G. Jalkotzy, and Mark S. Boyce
Lindzey 1984, 97). Seasons typically end in the spring,
before ungulate peak birthing periods, in order to reduce
the potential of disturbing ungulate neonates ( Table 4.1).
In northern states, timing of seasons in this manner often
did little to reduce cougar hunting opportunities because
hound hunting was normally limited to periods of optimal
opportunity, when snow was on the ground. Additionally,
most hound hunters chose to avoid situations where their
dogs might be shot (e.g., during big game seasons). Season
timing has also been used to reduce the likelihood of kittens
being killed by dogs (e.g., Utah, Colorado). Because cougars
give birth year-round, however, timing of seasons does not
protect all kittens; patterns of cougar litter production are
given in Chapter 5 (see Figures 5.25.6). When harvest quo-
tas are used, seasons end when the quota is met.
Regulating Hunter Numbers and Distribution
Regulation of sport hunting for cougars typically follows
one of three harvest strategies: general seasons, limited entry,
and harvest quota systems (see Table 4.1; Cougar Manage-
ment Guidelines Working Group 2005). General seasons
allow unlimited hunting of cougars of either sex and have
the least control over harvest levels, the only restrictions
being the number of licenses issued per hunter (typically
one per season) and the timing and length of the hunting
season. General seasons provide the most hunting opportu-
nity and have the least control over harvest levels. Also they
may result in uneven hunting pressure—accessible areas are
hunted more heavily than inaccessible areas, limiting con-
trol over the harvest level, composition, and distribution.
Limited entry programs may restrict the number of hunt-
ers per management unit through limited license allocation,
using either a fi rst-come, fi rst-served basis or lottery license
sales. This approach may be the most limiting in terms of
hunter opportunity but can be useful to disperse hunting
pressure and control harvest levels, and it may increase the
opportunity for hunters to be selective (increasing the male
harvest) in areas where hunting pressure is low. For hunters
willing to travel, limited entry may continue to allow simi-
lar hunting opportunities if the number of permits allocated
is high.
Harvest quota management requires setting a limit on the
total harvest and/or the number of female or male cougars
harvested from an area. Quotas are not goals but allowable
harvest limits set to achieve specifi c population-level objec-
tives. The hunting season is closed for the area once the
quota has been met. Advantages of the quota approach are
that hunting opportunity remains high and harvest distribu-
tion and level can be regulated. The quota system requires an
agency to develop a method that allows hunters to monitor
Agencies have used a combination of approaches to man-
age cougars, often with only a limited understanding of the
effectiveness of their management prescriptions. Working
with imperfect knowledge is nothing new to biologists, but
it is a politically risky affair when the focus is an animal
over which people are dramatically separated in their views.
When faced with limited information, wildlife managers
tend to be conservative in setting harvest objectives. Because
knowledge of the effectiveness of various actions is slow
to accumulate, state- or provincewide management strate-
gies usually also change slowly. Political pressures, however,
sometimes cause sudden and drastic shifts in cougar man-
agement programs (discussed in Chapter 15).
Data Analysis Units
Although most states and provinces began cougar manage-
ment on a province- or statewide basis, many have subse-
quently delineated their cougar habitat based on vegetation
and topography, and presumably similar cougar densities,
to refi ne management efforts (i.e., cougar management
units). These areas are typically large enough to support
population-level analyses, although it is generally recognized
that they rarely contain isolated cougar populations. Man-
agement units, in turn, are frequently broken into smaller
contiguous areas or hunt areas that share “social” char-
acteristics (Wyoming Game and Fish Department 2006b),
such as cougar depredation history, land ownership, pub-
lic access, and hunting history, where unique management
strategies can be applied to address localized issues (e.g.,
historic depredation incidents, cougar-human interactions).
This division allows agencies to direct management actions
to address the sociopolitical aspects of cougar management.
Traditionally, social issues revolved around cougar depre-
dation of livestock or potential predation impacts on game
species (e.g., deer and elk). More recently, human-cougar
interactions have become of increasing concern as recre-
ational activities have increased in, or human development
has encroached on, cougar habitat.
Seasons
Earliest regulations typically provided year-round hunting,
but as management evolved, seasons were often shortened
and timed to refl ect specifi c objectives within management
or hunt areas. For example, year-round hunting has been
used to direct hunting effort into areas experiencing cougar
depredation problems. Seasons are often set to begin after
hunting seasons for other game species are over to avoid
confl icts among different kinds of hunters, particularly in
areas where cougar hunting involves dogs (Roberson and
Cougar Management in North America 43
the harvest so that they know when the quota has been met.
This is accomplished using a toll-free hotline that is continu-
ally updated as cougars are harvested; hunters are expected
to monitor the harvest by accessing the hotline. Occasion-
ally, quotas are exceeded because there is often a lag in the
reporting of kills and their entry on the hotline. This should
be recognized and adjusted for in the development of harvest
quotas. Female subquotas can be used to support a manage-
ment objective of sustaining harvest levels by limiting female
harvest levels and reducing impact on the cougar popula-
tion. Potential disadvantages of harvest quotas are that the
number of hunters per management unit is unlimited until
quotas are fi lled, and quotas may be exceeded if several cou-
gars are taken toward the end of the season but before the
harvest is recorded on the quota hotline.
All human-caused cougar deaths (including depredation
control removals and known accidental deaths, such as from
vehicle collisions) may or may not be counted against the
quota; Wyoming, for example, recently moved to include all
such deaths in its quota for fuller accountability. Counting
all human-caused mortalities toward management quotas is
a desirable management strategy because mortality factors
other than hunting likely contribute to cougar population
dynamics (Laundré et al. 2007).
Table 4.1 Cougar population statusa and characteristics of management programs in the western United States and Canadian provinces in 2008.b
State or
Province
Population
Size/Trend
Legal
Statusc
Season Dates
(Bag Limit)d
Season
Structuree
Dogs
Allowed
Female &
Cub laws
Pursuit
Seasons
Mandatory
Inspection
Depredation
Compensation
Alberta 800–1200/I Big game 12/1–2/28(1) FQ, MQ Yes Yes No Yes Yes
Arizona 1,500–2,500/Unk Big game 9/1–5/31(1+) Gen Yes Yes No Yes No
British
Columbia
4,000–6,000/S Big game 9/8–6/30(2)fGen, FQ Yes Yes Yes Yes No
California 4,000–6,000/S Protected NA NA No NA No NA No
Colorado 3,000–3,600/Unk Big game 11/19–3/31(1) TQ Yes Yes No Yes Yes
Idaho 2,000/D Big game 8/30–3/31(1–2)fGen, FQ Yes Yes Yes Yes Yes
Montana Unk/Unk Big game 10/21–4/14(1) LE, FQ, MQ Yes Yes Yes Yes No
Nevada 2,500–3,000/S Big game YR(2) TQ Yes Yes No Yes No
New Mexico 2,000–3,000/Unk Big game 10/1–3/31(1–2)gTQ Yes Yes No Yes No
North Dakota 27–101 adults/I Furbearer 9/1–3/11(1) TQ Yes Yes No Yes No
Oregon 5,700/I Big game 8/1–5/31(1–2)gGen, TQ No Yes No Yes No
South Dakota 200–225/I Big game 11/1–12/31(1) TQ, FQ No Yes No Yes No
Texas Unk/S Non-game YR/unlimited Gen Yes No No No No
Utah 2,528–3,936/Unk Big game 11/21–6/1(1)gLE, TQ Yes Yes Yes Yes Yes
Washington 1,000–2,500/D
21 counties
6 counties
Big game
9/1–3/15(2)
12/1–3/31(2)
Gen
LE, TQ, FQ
No
Yes
Yes
No
Yes
No
Yes
No
Wyoming Unk/S Trophy game 9/1–3/31(1)gTQ, FQ Yes Yes No Yes Yes
aPopulation size and trend based on subjective information such as harvest data, sightings, nuisance incidents, extrapolation of localized fi eld research, and/or literature-based
density estimates extrapolated to suitable cougar habitat. Trend: I = increase; S = stable; Unk = unknown; D = decrease. Population size and trend information reported from
most recent management summaries (Becker et al. 2003 or Martorello and Beausoleil 2005) if available, information accessible from agency websites, or Beausoleil et al. 2008.
bInformation accessed from management agency Web sites.
cLegal status change from predator to game animal: Colorado and Nevada in 1965; British Columbia and Washington in 1966; Oregon and Utah in 1967; California in 1969;
Alberta, Arizona, Montana, and New Mexico in 1971; Idaho in 1972; and Wyoming in 1973. Legal status in California changed from game animal to specially protected mammal
in 1990, and from protected to game animal in South Dakota in 2003 and North Dakota in 1991 (with a closed season until 2005).
dBag limit = maximum number of cougars harvested/hunter/year except in Arizona where some management areas allow for 1 cougar harvested/hunter/day.
YR = cougar hunting seasons open year-round.
eSeason structure: Gen = general; LE = limited entry; TQ = total quota; FQ = female quota or female subquota when used in combination with TQ; and
MQ = male quota.
fSeason dates vary among management areas within interval reported.
gSome management areas are open to cougar hunting year-round.
44 Charles R. Anderson Jr., Frederick Lindzey, Kyle H. Knopff, Martin G. Jalkotzy, and Mark S. Boyce
Hunting Methods
Methods of hunting cougars include opportunistic spot-and-
stalk hunting, calling cougars using predator calls, and hound
hunting—tracking and baying cougars using trained hunt-
ing dogs. Most western states and provinces allow hound
hunting, which has traditionally been the most common
and effective method for hunting cougars. However, some
stakeholders dislike the idea of pursing wildlife with dogs. As
a result, Oregon and Washington have banned hound hunt-
ing. Where hound hunting is not allowed, predator calling
and opportunistic cougar hunting during big game seasons
appear to be comparably successful, based on harvest levels
observed in Washington (Beausoleil et al. 2005) and South
Dakota Department of Game, Fish and Parks. “South
Dakota Mountain Lion Hunting Season,” http://www
.sdgfp.info/Wildlife/MountainLions/MtLionhuntingseason
.htm (accessed 2007). This may be a function of the increased
number of cougar hunting permits issued and longer sea-
sons in place of the more effective hunting method of using
hounds.
Results from Washington (Martorello and Beausoleil
2003) suggest that opportunistic cougar hunting is less
selective than hound hunting. This has made female cou-
gars more vulnerable: relative female harvest levels increased
from 42 percent to 59 percent after hound hunting was
banned in Washington. Harvest data from western states
(management status reports in Becker et al. 2003, Beausoleil
and Martorello 2005) suggest that hound hunting results
in the higher harvest of males than females. Presumably,
hound hunters have better opportunity than other hunt-
ers to identify females because they can often distinguish
size differences when they encounter a track. They also can
spend time observing the animal once it is treed. Addition-
ally, they are more likely to encounter males while tracking,
because males travel distances that average more than twice
the average distances for females (Anderson 2003). On the
other hand, opportunistic hunters who are not tracking cou-
gars are more likely to encounter the more abundant sex
(females; Logan and Sweanor 2001, Laundré et al. 2007).
Bag Limits/Permits. Agencies have offered “sportsman
packages” that included a cougar permit with the purchase
of other game tags, but most states and provinces typically
require the purchase of a separate cougar permit. Bag limits
are most commonly set at one per season, but larger bag
limits have been used to raise harvest levels in specifi c areas
(e.g., where depredation incidents are high). Sportsman
packages are more common where hound hunting is not
allowed, in an attempt to increase the number of licensed
hunters afi eld and thus maintain similar cougar harvest with
less effective methods.
Achieving Desired Cougar Sex and Age Classes of Harvest. It
is illegal in most jurisdictions (with the exception of Texas;
see Table 4.1) to kill spotted juveniles or females with young
at their side (prohibitions known as cub laws; management
status reports in Becker et al. 2003, Beausoleil and Martorello
2005). Although there are birth pulses, typically May–October
( Figures 5.25.6; Cougar Management Guidelines Working
Group 2005, 53), cougars may have young at any time of the
year. Consequently, because kittens do not always accompany
their mother, particularly when very young (Barnhurst and
Lindzey 1989), hunters may unknowingly kill females that
have young. Based on the proportion of reproductive-age
females killed by hunters each year, evaluations in Wyoming
(Wyoming Game and Fish Department 2006b) suggest that
juvenile loss resulting when females with young are killed
average about twenty-two per year. Although undesirable, this
loss should have limited effects on the statewide population.
Female subquotas, cub laws, and the statutory timing of
seasons to exclude summers are intended to offer juveniles
and females with young some protection. Aiming in part to
help hunters identify females and look for signs that they
may have young, Colorado and Washington have recently
introduced mandatory cougar hunter education programs,
and New Mexico, Utah, and Wyoming have similar voluntary
programs.
Pursuit Seasons. Currently, four states and one Canadian
province provide special seasons during which cougars can
be pursued by dogs and treed but not killed (Table 4.1).
Some jurisdictions prohibit pursuit during ungulate seasons
(Roberson and Lindzey 1984, 87). Pursuit seasons allow hound
hunters increased opportunity to work their dogs by baying
and releasing cougars during the pursuit season. Although
cougar populations are ostensibly not directly impacted by
this practice, unintentional kitten loss, potential for stress-
related mortality (Harlow et al. 1992), and increased illegal
take may result.
Dealing with Depredation. States and provinces have used
various approaches to deal with livestock depredation.
Generally, livestock owners are given a fair amount of
latitude in dealing with problem cougars. Typically, they can
kill the offending cougar without fi rst obtaining a permit
from the state. They are required to report any cougars killed
and to justify their actions. Agencies have also responded to
depredation problems by employing professional hunters,
structuring hunting regulations to increase sport harvest in
problem areas, reimbursing livestock owners for animals
killed by cougars (four states and one Canadian province,
Table 4.1), and contributing monies to federal animal damage
control programs (Roberson and Lindzey 1984, 102, 204).
Cougar Management in North America 45
Refi ning the Procedures
Documenting the harvest level and composition is a required
component of management. Hunter questionnaires have
been used to estimate total harvest, location of kill, sex and
age of the cougar taken, and effort expended. More com-
monly, agencies require that harvested cougars be checked
by agency personnel, when biological data and hunter infor-
mation are recorded. Cougars taken under depredation pro-
visions are subject to similar reporting requirements, except
in Texas, where cougar take is unregulated and mortality
reports are voluntary (Young 2003).
Management is the attempt to achieve desired objec-
tives. Success is measured by how closely the results of the
management prescription match the desired outcome. This
implies that results can be measured, but cougar densities
are rarely known or measured due to the cryptic nature
of this solitary large carnivore and the rugged terrain it
occupies. Hunting is the most controversial component of
cougar management. Besides providing recreational oppor-
tunity, hunting is also used to alter populations in an effort
to achieve specifi c objectives, such as reducing livestock
depredation or predation on other animals, both of which
are assumed to be related to cougar density. Although the
relationship between cougar density and ungulate predation
or depredation level is assumed to be linear—that is, a given
percentage reduction of the cougar population will result in
a proportionate reduction in predation or depredation—this
has not been demonstrated (Cougar Management Guide-
lines Working Group 2005). Specifi cally, we do not know
what percentage of the cougar population must be removed,
if any, or what seasonal conditions, prey number regimes,
and or husbandry practices result in a given level of preda-
tion or depredation reduction.
Population Assessments
Direct measures of cougar population characteristics (den-
sity, sex, and age composition) are needed to evaluate success
of management programs, but estimating cougar numbers
and documenting composition are diffi cult. The best density
estimates have come from long-term studies of relatively
small areas where most cougars were captured and dynam-
ics of the populations were monitored (e.g., Hemker et al.
1984; Ross and Jalkotzy 1992; Logan and Sweanor 2001,
Laundré et al. 2007). Such studies are costly, and they often
yield data from central habitats where circumstances may
differ from those in more peripheral parts of the species’
range. Nevertheless, these density estimates provide a start-
ing point for management programs.
Track surveys (Smallwood and Fitzhugh 1995) can pro-
vide estimates of relative cougar abundance in areas where
tracking substrates are suitable, abundant, and suffi ciently
spaced to support a proper sampling design. Line intersect
probability sampling, proposed by Van Sickle and Lindzey
(1991), and later evaluated and expanded on by C. R. Ander-
son (2003), can yield precise density estimates and can detect
1530 percent changes in population size with intensive
sampling of moderate to high density populations that pro-
vide reasonable sample sizes (at least ten cougar tracks per
survey). This approach, however, is expensive and geograph-
ically limited in application, because it involves helicopters
and requires specifi c snow conditions.
Indexes or indicators of population trend are valuable
tools for agencies, but if population size is unknown, the
actual relationship between the index (e.g., cougars har-
vested, depredation events, sightings) and population size is
also unknown. Thus, where rigorous documentation of pop-
ulation status is still needed, indexes have limited use alone or
even in combination. Anderson and Lindzey (2005) used the
relative vulnerability of cougar sex and age classes to hunting,
as described by Barnhurst (1986), and monitored changes in
composition of a hunted population as it declined and recov-
ered, in order to document compositional shifts that could
be used to index population status. Various methods that use
genetic identifi cation with sight-resight analyses have been
proposed (e.g., Ernest et al. 2002) and may provide useful
population estimators in the future. Regardless of the moni-
toring methods available, multiple indicators should be used
if the best evaluation of cougar management programs is to
be achieved.
Cougar Management Plans
Most western states (except for Arizona, Montana, and
Texas) periodically develop plans that, in effect, set the
policy for cougar management and attempt to balance bio-
logical and social aspects. These plans typically include syn-
theses of available knowledge about ecology of the cougar
and about management practices, provide for information
and education efforts on how to avoid confl icts, and are
intended to refl ect stakeholder values incorporated after
often lengthy public input sessions. Broad parameters are
set for how objectives will be determined for hunt areas and
at times management areas, how hunts are structured to
accomplish objectives, and how accomplishment of objec-
tives will be measured.
Development of cougar management plans took hold
primarily in the 1990s in response to several factors, includ-
ing an accumulation of management and biological informa-
tion from the Mountain Lion Workshops (see Chapter 2,
Table 2.2), new agency personnel having a broader eco-
logical background, the reported increase in cougar popu-
lations regionwide (management status reports in Becker
46 Charles R. Anderson Jr., Frederick Lindzey, Kyle H. Knopff, Martin G. Jalkotzy, and Mark S. Boyce
et al. 2003), increasing cougar-human interactions (e.g.,
Fitzhugh et al. 2003), and growing pressure from more
diverse stakeholders challenging traditional cougar manage-
ment programs. For the fi rst time, cougar management began
to include the human factor more explicitly. Approaches were
developed to educate people living in and using cougar habi-
tat on how to avoid confl icts (see Beausoleil et al. 2008).
In addition, the management planning process began
to acknowledge the value of cougars beyond recreational
hunting, recognize potential threats to cougar populations
from factors other than hunting (e.g., habitat loss), and
accept the need for justifi cation of hunting as a manage-
ment tool and for transparency in the management planning
process to ensure that all stakeholders are included. These
are ostensibly major shifts, but results are variable; some
of the newer participants contend that even where broader
public input is effectively gathered, it is nevertheless often
ignored (Chapter 14).
Evaluation of Management
Experience using various combinations of seasons, permits,
and sex and age restrictions to achieve management goals
has provided managers insight into the effectiveness of the
different approaches. Dawn (2002) conducted the fi rst
broad-scale evaluation of cougar management strategies
and, despite the apparent trend of progressively greater
restrictions on hunters, results showed that the number of
cougars taken by sport hunting increased (see Figure 4.1;
Appendix 2). She noted that increases in harvests after states
and provinces assumed management authority did not nec-
essarily (but could) mean that cougar numbers had grown
since the presumed lows in the mid-to late 1960s (Nowak
1976). Other factors might also have bearing, such as num-
ber and effectiveness of hunters and liberalization of allow-
able harvest limits. Further, Dawn’s analyses suggested that,
among the season types, general seasons yielded the lowest
harvest rates, a result she noted possibly refl ected the fact
that general seasons were most common during the early
agency management period when cougar numbers were
presumably low. The lowest percentages of females in the
harvest occurred when female subquotas were imposed.
Ross and colleagues (1996) also found that the harvest
increased under the quota system in Alberta. However, they
noted that the increase may have been due in part to con-
current increases in season length. The quota system with a
female subquota resulted in a reduction of the proportion
of females taken by sport hunters from 43 to 29 percent,
with the increased harvest being composed primarily of
males. Comparisons among various harvest methods over
time can be tenuous because cougar numbers and hunter
effectiveness may change. For example, less restrictive
management strategies—that is, general seasons—are more
common where cougar hunting is less effective, namely, in
Figure 4.1 Reported number of cougars killed by sport hunters in the western United States, 1980–2007. Calendar year reported from Montana, Arizona
and Oregon, and harvest year (fall of the previous year to spring of reported year) reported from Idaho, Utah, Colorado, Washington, New Mexico, Nevada, and
Wyoming. The overall reported harvest increased threefold from the early 1980s to the late 1990s.
Cougar Management in North America 47
areas lacking snowfall for tracking or areas where hound
hunting is not allowed. Moreover, most states have shifted
their management strategies in similar ways, broadly from
general seasons to setting quotas (Dawn 2002, 25), during
the period when cougar populations were considered to be
increasing (management status reports in Becker et al. 2003
and Beausoleil and Martorello 2005). Further evaluations
of the effectiveness of type and timing of seasons relative to
cougar densities will increase the confi dence of managers in
their application. State and provincial agencies are uniquely
positioned to test the prescriptions in their management
plans, if they so choose, using designs built into an adaptive
framework (see also Adaptive Management on page 49).
Trends in harvest level should also be interpreted cau-
tiously and in context. Harvest levels in western North
America show remarkably similar trends, increasing during
the early to mid-1990s, then leveling during the late 1990s
and early 2000s, and exhibiting recent declines in some
states (see Figure 4.1; Appendix 2). Although annual fl uc-
tuations in cougar harvests are in part related to chang-
ing harvest quotas/management strategies, some observers
interpreted increasing harvests as endangering cougar pop-
ulations, while others viewed the numbers as an indication
that populations were growing at least suffi ciently to sup-
port this increased harvest. States and provinces generally
felt they were dealing with increasing populations (man-
agement status reports in Becker et al. 2003; Beausoleil and
Martorello 2005) during the 1990s, even though evidence
to support this conclusion was largely anecdotal, based on
crude indices or a few long-term, localized studies. Four
research efforts occurring at different but overlapping time
spans and diverse locations (from New Mexico to Alberta)
documented increasing cougar densities beginning in the
mid-1980s and continuing through the mid-1990s (Ross
and Jalkotzy 1992; Murphy 1998; Logan and Sweanor
2001; Laundré et al. 2007). Whether these localized obser-
vations refl ected regionwide cougar population trends,
however, is unknown. Three factors suggest that cougar
populations were stationary, if not increasing, during the
rst three decades of agency management: (1) there was
no consistent trend toward increasing adult females in
the harvests, which would indicate that populations were
being affected (Anderson and Lindzey 2005), (2) cougar
populations were reestablishing along the species’ eastern
range (i.e., South Dakota, North Dakota; see Range Map,
p. vii) and (3) cougar observations have come from even
farther east and in Canada (Chapter 12; see also Canadian
section below).
The recent downturn in harvests from some western states,
however, may warrant attention. We do not know whether
this trend refl ects a reduction in cougar numbers and, if
so, whether the reduction results from habitat conditions,
prey densities (e.g., Laundré et al. 2007), increasing harvest
levels (Dawn 2002), hunter participation, or a combina-
tion of these factors. Management summaries from the
2005 Mountain Lion Workshop Proceedings (Beausoleil
and Martorello 2005) are confl icting relative to recent
trends, suggesting a decline in cougar populations in British
Columbia, Idaho, and Washington; increasing populations
in Oregon and South Dakota; and stable populations in
California and Nevada. If western cougar populations have
indeed increased over the past thirty years and are reaching
equilibrium, expansion eastward should be expected to con-
tinue and to present new management challenges in areas
being recolonized.
These population questions are controversial, and par-
ticipants in the cougar debates tend to have very specifi c
objectives. Livestock interests want management to lower
depredation levels, and it has generally been assumed that
the level of depredation is directly related to cougar den-
sity. Yet, the actual form of the relationship between cougar
density and level of livestock depredation is not known.
Addressing depredation simply by increasing the cougar
harvest may fi nd favor among livestock interests and deer
and elk hunters who perceive a benefi t from reduced cougar
densities, but it garners less enthusiasm from the broader
public and some cougar hunters, whose main interest is to
ensure healthy/abundant populations. Thus, the challenge
to management agencies is not merely to devise a biology-
based approach that balances different interests but also to
sell that approach to everyone and keep learning as we go
along. While it may be relatively easy to document changes
in livestock depredation or hunter opportunity, it is much
more diffi cult, but equally necessary, to document concur-
rent changes in the cougar population.
Among the population characteristics that have bearing
here is the fact that cougar populations tend to be genetically
and physically connected over large areas (see Chapter 3;
Culver et al. 2000a; Sinclair et al. 2001; Anderson et al.
2004), with segments of each population’s overall area vary-
ing in suitability as cougar habitat and in terms of hunter
access. The source-sink thesis described in Chapter 5 sug-
gests that areas with low cougar survival (whether because
of hunting and or other deaths) are supported by immi-
gration from adjacent source areas, where survival and
reproduction are higher. Logan and Sweanor (2001), and
later Laundré and Clark (2003), used the source-sink con-
cept to suggest a zone management and a metapopulation
approach, respectively, whereby refuge areas are formally
delineated within a state or province to guarantee sources.
Abundance within these refuges of higher survival, and
dispersal from them, should then ensure a supply of immi-
grants for exploited areas and assure continuation of viable
populations.
48 Charles R. Anderson Jr., Frederick Lindzey, Kyle H. Knopff, Martin G. Jalkotzy, and Mark S. Boyce
Managers have long been aware that some areas of
suitable cougar habitat are not hunted, or are seldom
hunted; hunt areas were often initially delineated based on
access, relative cougar density, and/or prevalence of dep-
redation problems. Review of harvest records providing
mortality density (number of human-caused deaths/area
of cougar habitat) and sex and age of harvested cougars
(Wyoming Game and Fish Department 2006b) would help
confi rm initial assignment of hunt areas to source or sink
status. Sex and age composition of harvests are best inter-
preted on the basis of whether the harvest is from a source
or sink area. Harvests composed primarily of young and
male cougars might be expected from both source and sink
areas. For a source area, a preponderance of young male
animals refl ects harvest of the most vulnerable age class,
with reproductive-age females remaining relatively secure.
For a sink area, such a harvest may simply refl ect the main
category of cougars present (Anderson and Lindzey 2005).
Past cougar mortality records providing changes in harvest
sex-age composition, human-caused cougar mortality den-
sities, and age estimates showing changes in age structure
can assist in determining initial source or sink status of
cougar populations. Monitoring changes in demographics
of harvested cougars over time should allow detection of
declines or increases in that population (Wyoming Game
and Fish Department 2006b).
Adaptive Management
State and provincial agencies do have the framework to
design and conduct experiments to answer questions about
the effectiveness of approaches such as using hunting to
reduce livestock depredation, reduce predation on other
desired wildlife, or reduce human-cougar interactions.
Whether and how agencies use that framework depends on
levels of interest, budget, and public pressure. The Cougar
Management Guidelines (2005, 9) suggest that “adaptive
management is characterized by the continual monitoring of
indicators that measure progress toward the achievement of
management goals and objectives, changing of management
practices when new information indicates that better alter-
natives are available, monitoring relevant stakeholder values
and interests, and the monitoring of natural environmental
changes that may affect cougar management results.” While
this statement seems to fi t the cougar management approach
of most agencies, learning from experiments requires that
careful thought be devoted to their design and that agency
and stakeholder support be secured.
Management actions should be advanced as questions to
be asked or hypotheses to be tested, with specifi c predictions
made as to the outcome. For example, if an agency wished to
test the question of whether sport hunting of cougars could
reduce depredation on domestic sheep, the design would
need to include means to measure level of cougar removal
by sport hunting and level of sheep losses to cougars over
the time of the experiment. Obviously, many other vari-
ables could act to infl uence the results. Sheep numbers and
husbandry practices would ideally be held constant during
the experiment, as would level of cougar harvest (this is
where agency and stakeholder buy-in becomes essential).
Weather patterns and general trends in cougar prey avail-
ability should be monitored as well.
The results of such an experiment conducted in an area
of contiguous cougar habitat might well differ from results
in a more isolated area, indicating the need to replicate the
experiment in other locales in order to predict better when
and where sport hunting may be an effective tool to reduce
sheep depredation problems. Results of properly designed
experiments could be of value to managers throughout
the species’ range, and if they were replicated with similar
methods by a number of states and provinces, the cost and
effort of gaining information could be shared.
Recent Changes in Management Status
Most western states and provinces have management
authority over cougars, with the exception of Texas, where
the species is not classifi ed as a game animal and harvest
is unregulated (Haverson et al. 1997; see also government
mandates and jurisdictions discussion in Chapter 15). Man-
agement fl exibility has been curtailed in some states and has
more recently been relaxed in others. In 1990, California
voters approved Proposition 117, which prohibited sport
hunting of cougars. In 1994 a citizen ballot initiative called
Measure 18 passed in Oregon, prohibiting hound hunting
for cougars, and in 1996 Washington voters approved Ini-
tiative 655, likewise prohibiting the hunting of cougars with
hounds (see Appendix 5 for more detail). Bills passed in
2004 in Washington (Beausoleil et al. 2005) and in 2007
in Oregon relaxed these prohibitions somewhat, allow-
ing hound hunting for cougars in fi ve Washington coun-
ties (increased to six counties in 2008) to address livestock
depredation concerns and allowing the use of hounds for
specifi c agency management actions in Oregon (e.g., depre-
dation incidents). Cougars were reclassifi ed in North Dakota
in 1991, from state-threatened species to furbearer. South
Dakota reclassifi ed cougars in 2003, from state-threatened
species to big game animal.
Cougars have long been extirpated from most areas east
of the Rocky Mountains except for the Florida panther,
which the state classifi ed in 1958 as endangered, with the
federal government following suit in 1967 (Lotz 2005).
Management emphasis went from documenting cougar pres-
ence to protection and recovery efforts (see Chapters 3, 12).
Cougar Management in North America 49
Presence of cougar populations in other areas east of the
Dakotas is currently unconfi rmed, but sightings and occa-
sional confi rmed reports of dead cougars have increased
in the past fi fteen years (see http://www.cougarnet.org/
network.html for examples). Confi rmed deaths and reliable
reports have been documented in several midwestern states
(e.g., Iowa, Missouri, Oklahoma). Whether these cougars
are of captive or wild origin is unconfi rmed in many cases,
but known dispersal of radio-collared cougars from South
Dakota into Oklahoma and Minnesota (Daniel Thompson,
South Dakota State University, pers. comm., 2008) supports
the notion that many of these records, at least west of the
Mississippi River, are true wild dispersers.
Additional factors also suggest potential expansion of
western cougar populations eastward: the prevalence of
males of dispersal age among the confi rmed deaths; reemer-
gence of cougar populations in eastern areas adjacent to the
Rocky Mountain West (North Dakota, South Dakota, and
possibly western Nebraska); and a recent increase in reliable
cougar reports. As noted, observers in several western states
considered cougar populations to be increasing during the
1990s, and this is supported by localized research docu-
menting increasing cougar populations from New Mexico
to Alberta (Ross and Jalkotzy 1992; Murphy 1998; Logan
and Sweanor 2001; Laundré et al. 2007; on Canada, see
below), and by genetic evidence suggesting recent demo-
graphic increase and expansion (Biek et al. 2006b). Whether
establishment of cougar populations eastward succeeds will
depend largely upon social acceptance and the proactive
involvement of local wildlife agencies. Of the states bor-
dering the known cougar distribution in the west, cougars
are designated as protected in Louisiana and Wisconsin;
game species with a zero harvest quota in Nebraska and
Oklahoma; nongame in Arkansas, Kansas, Missouri, and
Minnesota; and as unprotected in Iowa. Modifi cation of
management designations and implementation of manage-
ment planning may be necessary to accommodate and/or
prepare for cougar recolonization eastward.
Special Circumstances in Canada
Although most cougar research in North America has been
conducted in the United States, fi ndings presented in this
book on cougar population dynamics, morphology, behav-
ior, and interactions with prey are just as relevant to cougar
north of the 49th parallel. Similarly, many of the general
management issues described above also apply to Canada.
Several aspects of the status and management of cougar in
Canada are suffi ciently distinct, however, that they deserve
special attention: population distribution and abundance;
level of protection, harvest, and control; and potential for
range expansion and its implications for the northernmost
cougars.
Distribution and Population Status
Prior to European settlement, cougar were present through-
out the southern portions of Canada. Historical popula-
tion size is unknown, but by the early 1900s a reduction of
ungulates through market hunting and simultaneous perse-
cution of predators probably restricted cougar at low densi-
ties to portions of British Columbia and western Alberta. As
ungulate populations recovered by the mid-1900s cougar
numbers are also thought to have increased, but as in the
United States, bounty hunting and widespread poisoning
for predator control may have limited recovery (Jalkotzy
et al. 1992). Currently, breeding populations of cougar are
known to be present in both Alberta and British Columbia
(see Range Map, p. vii).
With the possible exception of the far north, breeding
populations of cougar are found across the entire British
Columbia mainland and have also found their way onto
the coastal islands, including Vancouver Island, where den-
sities have reached levels among the highest ever recorded
(Wilson et al. 2004). The estimated number of cougar in
British Columbia is between 4,000 and 6,000 individuals
(Austin 2005). By contrast, in Alberta, breeding populations
have traditionally been relegated only in a relatively small
region along the western edge of the province, where the
most recent published population estimate is 640 cougars
on provincial lands—that is, excluding Banff and Jasper
national parks (Jalkotzy et al. 1992). Our ongoing research
data suggest that either cougar populations have increased
north of the Bow River, where they had not previously been
directly studied, or the original estimates were too low in
this large portion of Alberta’s cougar range. In either case,
current cougar numbers in Alberta likely exceed the esti-
mate provided in the early 1990s.
Breeding populations of cougar have recently become
established outside the previously well-defi ned eastern
boundary for Canadian cougar along the foothills of
the Rocky Mountains in Alberta (see Range Map, p. vii).
In the Cypress Hills Interprovincial Park, which straddles
the Alberta-Saskatchewan border, a large number of cougar
sightings, accidental trapping of an entire family group in
snares set for coyote in December 2006, and high-quality
photographic evidence of a second family group in August
2007 ( Figure 4.2) are strong evidence that a breeding popu-
lation exists in the park. Elsewhere in Saskatchewan, large
numbers of cougar have been accidentally snared, shot, pho-
tographed, or confi rmed by wildlife personnel since 2000.
This, combined with numerous other sightings and reports
across the province, has recently prompted the government
50 Charles R. Anderson Jr., Frederick Lindzey, Kyle H. Knopff, Martin G. Jalkotzy, and Mark S. Boyce
to provide a provincewide estimate of approximately 300
cougars (Saskatchewan Department of the Environment
2007). Other than in the Cypress Hills Interprovincial Park,
it is not clear where breeding populations might be found
within Saskatchewan.
Cougar sightings also are common in Manitoba, where
a female cougar was shot and a male was trapped near the
Duck and Riding mountains in 2004, suggesting the possi-
bility of a small resident cougar population (Watkins 2005).
In Ontario, large numbers of unconfi rmed sightings in the
Great Lakes region (e.g., Ontario Puma Foundation 2007)
have led the provincial government to offi cially accept the
presence of a breeding cougar population (Ontario Minis-
try of Natural Resources 2007). Sightings are often unre-
liable indicators of cougar presence, however (Beier and
Barrett 1993), and confi rmation of breeding populations
will require further evidence. If populations are present east
of Saskatchewan, it remains unclear whether these are the
remnants of original eastern cougar populations or derive
from recent expansion out of the west or from the Dakotas
to the south (see Range Map, p. vii).
Confirmed presence of individual cougars has been
reported in many other locations across the country. In
Quebec and the Maritime provinces, for example, dedi-
cated researchers have recently obtained genetic evidence
of cougars occurrence (Gauthier et al. 2005). Cougars also
have been confi rmed as far north as the southern portion
of the Yukon Territory (Jung and Merchant 2005). Dis-
persing cougars can cover incredible distances (Thompson
and Jenks 2005), and many isolated occurrences and even
repeated sightings may represent dispersing animals (see
Chapters 5, 8, and 12). Alternatively, some of the east-
ern sightings and confi rmed occurrences could be animals
that have been released from captivity. Where confi rmed
incidents are particularly concentrated outside the known
distribution of breeding populations, they may indicate resi-
dent populations occurring at low densities.
Management and Conservation
As in most western states, cougars were managed as a boun-
tied predator in Canada until the mid-twentieth century.
They were extirpated from eastern Canada, and it was
not until 1966 in British Columbia, and 1971 in Alberta,
that the cougar achieved the status of big game animal.
In Alberta, a comprehensive management plan is now in
place (Jalkotzy et al. 1992), but it has not been updated
since it was adopted in the early 1990s. At the time of this
writing, there is still no offi cial management plan for cougar
in British Columbia, despite the fact that it is a species of
important management concern in that province (Robinson
et al. 2002).
This is quite different from the way that cougars are man-
aged in the United States, where offi cial management plans
are now the norm and are updated regularly in many states
where cougar occur (e.g., Apker 2005; Barber 2005). Lack of
a plan in British Columbia and of any updates to the Alberta
plan mean that offi cial activity does not incorporate recent
developments in our understanding of predator-prey ecology,
such as information on the ecosystem benefi ts derived from
the presence of large carnivores, including cougars (Ripple
and Beschta 2006). Stakeholder opinions, moreover, are not
offi cially acknowledged. In the United States, stakeholder
opinion is at least designated as a component of most cougar
management plans, and public action has infl uenced man-
agement through ballot initiatives in California, Oregon, and
Washington. In Alberta and British Columbia, by contrast,
public opinion and new ideas may receive consideration by
provincial agency staff in their day-to-day duties and may
have some bearing on cougar management, but they are not
refl ected in offi cial plans.
Despite the lack of an offi cial plan in British Columbia
or regular updates in Alberta, cougar management remains
an important issue. Cougar hunting is permitted and closely
monitored in both provinces. In British Columbia, with the
exception of some northern management units, cougar
hunting is permitted throughout the province. The hunt is
unlimited entry, with many units maintaining a bag limit
of two cougar per hunter per season. Some management
districts, however, have a female subquota limiting the num-
ber of females that can be taken each year. Cougar harvest
in Alberta is regulated by a strict quota system for both
males and females, with sex-specifi c harvest limits for each
of several cougar management areas in the western part of
the province (Ross et al. 1996). Both provinces permit hunt-
ing with hounds and forbid the harvest of spotted kittens
or of females traveling with spotted kittens, although this
regulatory constraint is a very recent development in Brit-
ish Columbia (Austin 2005). There is no season for cougar
in the wildlife management units of eastern and northern
Figure 4.2 A camera trap captured this cougar family in Cypress Hills
Interprovincial Park in Alberta, Canada. This is the fi rst photographic evidence
demonstrating that breeding populations of cougar have moved east to the
Alberta-Saskatchewan border. Photo by M. M. Bacon, Cypress Hills Interpro-
vincial Park Cougar Study .
Cougar Management in North America 51
Alberta, nor may cougar be hunted in any of the other
Canadian provinces or territories.
An important management issue for western Cana-
dian cougar revolves around accidental snaring of the big
cats. Snaring of wolves near carrion bait, often road-killed
ungulates, is a common recreational and economic activ-
ity in western and northern Canada. Scavenging behavior
has been demonstrated in cougars (Bauer et al. 2005), and
some individuals scavenge frequently from animal remains
(K. Knopff, unpublished data). Cougars are, therefore, sus-
ceptible to becoming by-catch in wolf and coyote snares.
Because snaring for wolves is not currently permitted where
cougars exist in the United States, cougars by-catch at wolf
bait stations is a distinctly Canadian concern. Similar con-
cerns exist, however, in states where snaring for coyotes is
permitted and cougars also are present (Wyoming Game
and Fish 2006b).
By law, all human-caused cougar mortalities must be
reported to provincial authorities. We analyzed the data for
all reported mortalities in Alberta between January 2000
and March 2006. Over that six-year period, 837 human-
caused cougar mortalities were reported. The vast major-
ity of these were cougars harvested by licensed hunters
(77 percent); the second most important source of human-
caused mortality was accidental snaring (9 percent). Less
frequent causes of mortality were removal of problem wild-
life (4.7 percent), road kills (4.7 percent), and self- defense
(2.3 percent). Trappers in Alberta cannot sell cougar hides
and must forfeit all cougars trapped or snared to the
province, so there are neither economic nor trophy incen-
tives for cougar snaring. The removal of so many cougars
through snaring is undesirable because it neither improves
the ability of managers to preserve self-sustaining cougar
populations nor assists in maximizing the benefi t to Alber-
tans through optimal allocation of the resource, two key
management objectives outlined in Alberta’s cougar man-
agement plan (Jalkotzy et al. 1992). This issue has not pre-
viously received a great deal of attention, but our analyses
show it to be an important source of human-caused cou-
gar mortality in Canada, and it may become increasingly
important in the United States if gray wolves are delisted
and wolf management practices in Montana, Idaho, and
Wyoming follow the Canadian model. Data currently
available for cougar and wolf habitat selection, prey-site
selection, and movement patterns (e.g., Alexander et al. 2006;
Kortello et al. 2007; Atwood et al. 2007) might be use-
fully applied to help reduce the probability of unwanted
by-catch of cougar at wolf bait stations, but the required
analyses have yet to be conducted.
As is true wherever cougars and people share the same
space, an important component of cougar management
in Canada involves managing cougar interactions with
livestock, pets, and people. Between 1890 and 2004, British
Columbia had thirty-nine cougar attacks on people, the
highest number of any jurisdiction in North America, with
the majority of the incidents occurring on Vancouver Island.
Alberta has only experienced a single lethal cougar attack
on a human (in 2001), but complaints involving cougars are
frequent. As noted, nearly 5 percent of all human-caused
cougar mortalities involve removal of problem animals by
wildlife offi cers, and a further 2.3 percent are the result of
self-defense. In addition, approximately as many problem
animals were relocated (thirty-six) as were killed (thirty-
nine) by provincial wildlife agencies between January 2000
and March 2006. The high number of complaints by resi-
dents has led Alberta to amend its laws to allow cougars
to be shot on sight on private land (Alberta Sustainable
Resource Development 2007). Animals so taken may not
be kept and must be turned over to provincial authorities.
Compensation for livestock loss is available in Alberta from
the Alberta Conservation Association but is unavailable in
British Columbia (Austin 2005). As in the United States,
high levels of negative interactions between cougars and
people in Canada are likely a result of larger numbers of
people living in and using cougar habitat, combined with a
general increase in cougar numbers in recent decades.
Cougar conservation and management are becom-
ing increasingly important topics in the provinces east of
Alberta, where cougars have previously been considered
extirpated but are reappearing (Watkins 2005). No offi cial
management, conservation, or recovery plans have been in
place for cougars in any of these provinces. Saskatchewan,
however, is now in the position of developing a strategy for
managing cougar populations and addressing confl ict with
people in regions where cougars are becoming established.
It and the other provinces east of Alberta will be responsible
for determining whether and to what extent cougars are able
to repopulate historic range. The Committee on the Status
of Endangered Wildlife in Canada (COSEWIC 2007) lists
the cougar in eastern Canada as data defi cient, meaning that
suffi cient information is not available to assess the status of
the eastern subspecies or assign it an extinction risk rating
(i.e., extirpated or endangered). Therefore, if small popula-
tions of eastern cougar exist, they are not currently pro-
tected under Canada’s Species at Risk Act (SARA). In 2007,
the Committee on the Status of Species at Risk in Ontario
independently designated the eastern cougar as endangered,
stating that “there have been hundreds of sightings of cou-
gars in Ontario over the years, and their presence here is
generally acknowledged” (Ontario Ministry of Natural
Resources 2007).
Consequently, the cougar now falls under the regulatory
arm of the province’s Endangered Species Act, affording cou-
gar in Ontario the highest level of protection of any cougar in
52 Charles R. Anderson Jr., Frederick Lindzey, Kyle H. Knopff, Martin G. Jalkotzy, and Mark S. Boyce
Canada. Revisions to the act, which took effect on June 30,
2008, require that no person kill, harm, or harass cougars
and that both industry and the public must refrain from dam-
aging cougar habitat. In addition, the act requires that the
Province of Ontario develop and implement a recovery plan
to bring cougar populations within the province up to a level
where they are no longer threatened with extinction (Endan-
gered Species Act, 2007, S.O., c. C-6). The Ontario cougar is
only the second subpopulation in North America, after the
Florida panther, to be classifi ed as an endangered species.
Range Expansion
Population estimates and harvest information for the states
and provinces with known breeding populations of cougars
(Beausoleil and Martorello 2005) suggest that Canada prob-
ably supports less than one quarter of North America’s total
cougar population. Relatively low human population densi-
ties, an abundance of suitable forested habitats, and large
populations of ungulate prey, however, suggest potential for
future population growth and range expansion in Canada.
Indeed, this is already occurring. Eastward expansion of
cougar may be occurring at present because it has taken
western cougar populations time to rebuild from the days
of bounties and general persecution. Cougar harvests and
population estimates reached all-time highs in most western
states and provinces around the turn of the new century
(Beausoleil and Martorello 2005). Detailed genetic analysis
(Biek et al. 2006b) lends additional support to the idea that
North American cougars recently underwent substantial
population expansion. With higher population densities,
greater numbers of dispersers can be expected, facilitating
expansion (see Chapters 4, 5, 12). Even so, range expansion
is likely to be a relatively slow process because colonizers of
both sexes must be simultaneously present and successfully
produce offspring in the new habitat, and female cougars do
not disperse long distances as often as males do (Sweanor
et al. 2000). Anthropogenic features may also slow recolo-
nization. For instance, while cougars do not generally avoid
roads (Dickson and Beier 2002), major highways can present
a barrier to dispersal if appropriate corridors or crossings are
unavailable (see Chapter 12; Beier 1995). Highway 2, which
runs north–south in Alberta and connects the major popu-
lation centers of Edmonton, Calgary, and Lethbridge, may
act as such a barrier, slowing expansion. Provided suffi cient
numbers of dispersing individuals are available, continued
eastward expansion of breeding populations into suitable
habitat where ungulate populations are high can reasonably
be expected.
Expansion north into boreal habitats that are not part of
historic cougar range is also possible. Continued industrial
development and global warming may play a role in the
potential for future cougar expansion into higher latitudes
in Canada. White-tailed deer ( Odocoileus virginianus ) have
increased in abundance in many parts of North America,
in some cases doing so well that they become a pest species
(Augustine and DeCalesta 2003). Additional forage created
by industrial deforestation combined with reduced snow
depth and cover in winter (Rikiishi et al. 2004) may cre-
ate conditions that are favorable for their expansion north-
ward. Indeed, in Alberta, white-tailed deer have increased
markedly (Latham, pers. comm., 2009) in some northern
boreal forests that were originally the domain of moose
( Alces alces ) and woodland caribou ( Rangifer tarandus cari-
bou ) and were home to very few deer (Stelfox 1993). With
increased prey densities, we speculate that cougar may suc-
cessfully colonize areas north of their traditional breeding
distribution (both present and historic), especially in ripar-
ian areas where prey is more plentiful. Between 2000 and
2004 in Alberta’s boreal region, over two hundred cougar
occurrence reports were fi led with the provincial govern-
ment. These reports range across northern Alberta and
include sightings, livestock depredation, and road-killed
cougars (e.g., districts of Athabasca, Grand Prairie, Fort
McMurray, and High Level). The reports do not confi rm
the presence of breeding populations, but, at a minimum,
they provide evidence of northern dispersal movements by
individual cougars.
Expansion back into original cougar range in eastern
Canada will serve to increase the resiliency of Canadian
cougar populations and could serve to restore some ecosys-
tem function in places where cougar have been absent for
decades or centuries. Wolf reintroduction into the north-
western United States has had important top-down effects
on ecosystems through trophic cascades (Beyer et al. 2007).
Cougars have been linked to similar kinds of trophic cascades
(Ripple and Beschta 2006), and effects on entire ecosystems
might reasonably be expected as a result of recolonization
(Chapter 10). As the Yellowstone wolf recovery program
also demonstrates, however, predator repopulation can be
controversial and can result in discontent among farmers,
ranchers, and hunters. It is unclear how cougar expansion
eastward will be received by Canadians, although there are
indications that the response may be positive in some juris-
dictions (Watkins 2005).
Expansion northward has similar potential to increase
the resiliency of Canadian cougar populations, and because
of lower human population densities in the north, it is less
likely to create controversy among residents. Ecosystem
effects of colonization are still likely, and may not always
be “positive.” Colonization of the boreal forest by cougar
and white-tailed deer, for instance, may have important
implications for other ungulates. Woodland caribou are
a species at risk in some portions of the Canadian boreal
Cougar Management in North America 53
forest (Edmonds 1991; Dzus 2001). The capacity for cougars
to impact caribou populations negatively through apparent
competition (Holt 1977) has been implied in other parts
of Canada where cougar traditionally occur (Kinley and
Apps 2001). Careful monitoring of cougar colonization
(both north and east) by the provincial wildlife agencies
would facilitate the identifi cation and effective management
of both the ecological and human confl ict issues that sur-
round cougar range expansion.
Conclusion
In Canada, the population distribution and size, level of
protection, and management of cougar are entering a period
of uncertainty and change. In the west, where cougar popu-
lations are well established, increasing human populations
and development of rural areas will likely increase inter-
actions between cougars and people. At the same time, there
is great potential in Canada for cougar range expansion to
both the north and east. Breeding populations of cougars
have likely already become established in Saskatchewan and
may also be present in Manitoba and Ontario. All this may
serve to increase the profi le of cougar in Canada and will
likely result in the need for regularly updated management
plans that account for stakeholder opinion, set protection
levels and harvest objectives, and provide response guide-
lines for human-cougar interactions. If range expansion
occurs at large scales, it will also increase the importance
of Canada as a stronghold for cougar populations and will
likely have important ecological consequences, both pre-
dictable and novel, for Canadian ecosystems.
In the United States, wildlife managers with responsibil-
ity for cougars will continue to face the same issues they
already know. Traditional stakeholders—livestock interests
and hunters—have grown to expect hunting to play a major
role in cougar management, whereas the larger public does
not share this expectation and calls for reducing a suite
of threats to cougar populations. The often diametrically
opposing views of participants in decisions make it likely
that middle ground will be hard to fi nd; each group will dis-
like some management decisions and insist that these refl ect
pressure from other stakeholders. It is easy to describe, and
perhaps even to implement, the steps agencies should take
in their management of cougars: well-developed propos-
als with clearly stated objectives and scientifi c support as
available; opportunity for stakeholder input; and trans-
parency in a decision-making process, addressing as many
stakeholder comments as possible. But even this approach
will not result in all parties being happy with management
decisions.
General growth of the human population and the trend
toward rural housing developments will increase contacts
between cougars and humans. Agencies need to hone proto-
cols for dealing with people and cougars in populated areas
and for handling incidents in which people are harmed or
threatened (see Cougar Management Guidelines Working
Group 2005, chap. 7). While most western states and Cana-
dian provinces contain vast areas of contiguous, suitable
cougar habitat, and cougars show remarkable fl exibility
in habitats that suffi ce as movement corridors, managers
should be aware that massive land use changes and human
structures can fragment habitats and compromise dispersal
corridors (Chapter 12).
In places, new issues are arising. Long before Europeans
settled North America, cougars commonly moved through
the plains states, and the frequency of recent confi rmed
reports suggests that this pattern may now be redeveloping.
If so, agencies in these states are acquiring a new responsi-
bility at a time of new complexity in public perceptions, and
they will need to be both responsive and proactive about
what to do in the novel situation of independent recoloniza-
tion efforts by a large carnivore.
Agencies and stakeholders will face many changes in
the future and need to consider their actions in the broader
context of how these will affect conservation of the spe-
cies. Currently, hunting is the single most controversial
aspect of cougar management programs. It is, after all,
premeditated killing of cougars for sport or to address
depredation, predation, or human safety concerns. Suc-
cess is easily demonstrated only for recreational hunting.
If hunting is removed from the equation, recreation is all
that will be lost. Cougars will continue to be killed to
protect livestock, to protect wildlife at risk (e.g., isolated
bighorn sheep populations), and to address human safety
concerns.
Of potential threats to the species, sport hunting is the
most visible and easily fi xed by simply banning it, but it
may be the least important in the long term. Alteration and
fragmentation of habitats for cougars and the ungulate
prey supporting them are ongoing and insidious and much
more diffi cult to control. Loss of cougars because of habitat
alteration will never be as obvious or as easily documented
as cougars killed by hunters, making it much more diffi cult
to develop support for necessary management actions. Deci-
sions to protect cougar habitat in place of human develop-
ment will be as controversial as decisions about hunting, or
more so, and much more diffi cult to implement. The next
phase of cougar management should see authority remain
with state and provincial management agencies, and manag-
ers and stakeholders should recognize that habitat manage-
ment is the crux of the long-term survival of the species.
54 Charles R. Anderson Jr., Frederick Lindzey, Kyle H. Knopff, Martin G. Jalkotzy, and Mark S. Boyce
... Additionally cause-specific mortality rates are important factors underly-ing survival, reproduction, and population structure (Quigley and Hornocker 2009). Cougars (Puma concolor) can be legally hunted in most western states of the United States (Anderson et al. 2009, Whittaker 2011) and the primary cause of adult mortality where hunting occurs is hunter harvest (Logan et al. 1986, Ross and Jalkotzy 1992, Lambert et al. 2006, Stoner et al. 2006, Robinson et al. 2014. Thus modification of hunter harvest is assumed to have the largest effect on cougar survival and as a result population growth (Clark et al. 2014). ...
... In hunted populations, the leading reported cause of mortality is generally hunter harvest (Lindzey et al. 1988, Robinson et al. 2008, Cooley et al. 2011) and female cougars tend to have higher survival than males in hunted populations where dogs are used to pursue cougars (Martorello and Beausoleil 2003, Robinson et al. 2008, Clark et al. 2014, Robinson et al. 2014, Wolfe et al. 2015, presumably because hunters can selectively harvest large males (Martorello and Beausoleil 2003, Anderson and Lindzey 2005, CMGWG 2005. Greater female survival is often the desired effect to meet the common management objective of maintaining a healthy population of cougars and maximizing opportunities for hunting (CMGWG 2005, Anderson et al. 2009). Sustained annual mortality rates of adult female cougars in excess of 20-25% can result in population decline (Lambert et al. 2006, Robinson et al. 2014. ...
... As suggested for other areas in the Great Basin, progressive land-use planning and public education will be important to the long-term conservation of cougars in this area (Wolfe et al. 2015). Further, because habitat loss and associated loss of connectivity is the most difficult threat to address for cougar populations (Sweanor et al. 2000, Logan and Sweanor 2001, CMGWG 2005, Anderson et al. 2009, Quigley and Hornocker 2009, we suggest it important to collaborate with stakeholders, particularly trappers and hunters, both of which are highly invested in wildlife and the habitat upon which it depends. Working cooperatively to determine ways to minimize incidental capture of cougars will likely be the most efficient way to implement effective practices that will be beneficial to cougars, trappers, hunters, and the public. ...
Article
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Cougars (Puma concolor) occupy mountain ranges throughout the Great Basin, Nevada, USA, where legal trapping of bobcats (Lynx rufus) is common and some non‐target captures of cougars in bobcat traps occur. Such incidental capture of cougars is an undocumented source of mortality because some cougars die from injuries several weeks after release from traps. We examined cause‐specific mortality and the effects of capture of cougars in bobcat traps on annual and overall (7‐year) survival during 2009–2015. We captured 48 cougars, of which we followed 33 until death. We estimated average annual survival rates for adult cougars and assessed the relative effects of sex, season, and long‐term effects of non‐target capture of cougars in foothold traps on estimated survival of adults using a nest survival model in Program MARK. We incorporated a time‐varying covariate to assess the long‐term effect of capture in non‐target foothold traps on survival of adult cougars. Average annual survival of non‐trapped females and males, regardless of trapping history, was significantly greater than females with a history of capture in a non‐target foothold trap; however, once partitioned across age, sex, and capture status, sample sizes were small. Our results suggest that capture in non‐target foothold traps decreases survival of adult female cougars directly by causing injuries that eventually result in mortality, and indirectly by increasing susceptibility to other forms of mortality. Mortality of adult females during the 7 years of our study caused by non‐target trapping was similar to hunting among radio‐marked female cougars, and this potential source of mortality has been unaccounted for in harvest objectives and harvest data for cougars without radio‐collars. Given anthropogenic sources of mortality accounted for 100% of recorded mortality of adult females, mortality from non‐target trapping is likely additive to other sources of mortality in our study area. We recommend regulatory agencies consider the possibility of unintentional take and potential for reduced long‐term survival of females where these large fields are sympatric with bobcats, and trapping of bobcats with foothold traps is a legal activity. Moreover, we suggest wildlife managers record information about the trapping incident, to include trap type, trap size, trap set type, location, number of days since last check, and type and severity of injuries when releasing incidentally captured cougars to inform future management decisions. Addressing other anthropogenic sources of mortality resulting from conflict with humans and road mortalities will be increasingly important as the human population expands into cougar habitat. © 2018 The Wildlife Society.
... In the early to mid-1900s, puma abundance throughout Colorado declined to a few hundred individuals, primarily due to unrestricted hunting and predator control. From 1965 to present, human-caused mortality in pumas was restricted (Anderson et al. 2010), and as a result the puma population in Colorado have grown from as low as a few hundred in 1965 to a few thousand adult and sub-adult pumas (Cahalane 1964;Anderson et al. 2010;Apker 2017). When puma numbers were low, interactions between pumas and domestic cats would consequently be limited. ...
... In the early to mid-1900s, puma abundance throughout Colorado declined to a few hundred individuals, primarily due to unrestricted hunting and predator control. From 1965 to present, human-caused mortality in pumas was restricted (Anderson et al. 2010), and as a result the puma population in Colorado have grown from as low as a few hundred in 1965 to a few thousand adult and sub-adult pumas (Cahalane 1964;Anderson et al. 2010;Apker 2017). When puma numbers were low, interactions between pumas and domestic cats would consequently be limited. ...
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Full-text available
Emerging viral outbreaks resulting from host switching is an area of continued scientific interest. Such events can result in disease epidemics or in some cases, clinically silent outcomes. These occurrences are likely relatively common and can serve as tools to better understand disease dynamics, and may result in changes in behavior, fecundity, and, ultimately survival of the host. Feline foamy virus (FFV) is a common retrovirus infecting domestic cats globally, which has also been documented in the North American puma (Puma concolor). The prevalent nature of FFV in domestic cats and its ability to infect wild felids, including puma, provides an ideal system to study cross-species transmission across trophic levels (positions in the food chain), and evolution of pathogens transmitted between individuals following direct contact. Here we present findings from an extensive molecular analysis of FFV in pumas, focused on two locations in Colorado, and in relation to FFV recovered from domestic cats in this and previous studies. Prevalence of FFV in puma was high across the two regions, ∼77 per cent (urban interface site) and ∼48 per cent (rural site). Comparison of FFV from pumas living across three states; Colorado, Florida, and California, indicates FFV is widely distributed across North America. FFV isolated from domestic cats and pumas was not distinguishable at the host level, with FFV sequences sharing >93 per cent nucleotide similarity. Phylogenetic, Bayesian, and recombination analyses of FFV across the two species supports frequent cross-species spillover from domestic cat to puma during the last century, as well as frequent puma-to-puma intraspecific transmission in Colorado, USA. Two FFV variants, distinguished by significant difference in the surface unit of the envelope protein, were commonly found in both hosts. This trait is also shared by simian foamy virus and may represent variation in cell tropism or a unique immune evasion mechanism. This study elucidates evolutionary and cross-species transmission dynamics of a highly prevalent multi-host adapted virus, a system which can further be applied to model spillover and transmission of pathogenic viruses resulting in widespread infection in the new host.
... Hatter (2019) found that annual estimates of human-caused mortality were correlated (P = 0.015) with the abundance of cougars in an interior BC cougar population. We did not use cougar kills/100 hunter days as an index because cougar harvests are dependent upon suitable snow conditions for tracking that vary considerably from year-toyear (Anderson et al. 2010). ...
... For instance, this study has found that the killing of a mountain lion is unpopular even in the case of an attack on a person, while past studies have shown killing to be the preferred management action in such circumstances (Casey et al., 2005;Manfredo, Zinn, et al., 1998;Thornton & Quinn, 2009). Likewise, compensation for lost livestock was unpopular in this study, while it is a common practice in other administrative areas (Anderson, Jr., Lindzey, Knopff, Jalkotzy, & Boyce, 2010). Policy and law within different administrative bodies often reflect these differences, as demonstrated by the lack of protection for the mountain lion in Texas, management as game in other Western states, and protection in California. ...
Article
Growth in the human population and the popularity of outdoor recreation have resulted in increasing interaction between humans and mountain lions (Puma concolor). A questionnaire was used to gauge attitudes, risk perception, and management preferences toward the species among residents near its habitat in Santa Cruz County, California. Attitudes were positive, risk perception moderate, knowledge low, and lethal control measures unpopular. More positive attitudes were found among men, respondents with more education, respondents who recreated often in natural areas, and nature organization members. Older respondents, women, those who recreated less in mountain lion habitat, and those who lived near (but not in) perceived mountain lion habitat demonstrated increased risk perception. Results could help align management actions with public preferences, and guide conservation organizations toward capitalizing on positive attitudes. Both management bodies and conservation organizations should target outreach toward addressing poor knowledge among groups with negative attitudes and higher risk perception.
... Presumably, cougar populations began to increase and possibly reoccupy historic ranges following this protection (Anderson, Lindzey, Knopff, Jalkotzy, & Boyce, 2010). Coinciding with this was the rapid expansion of human populations. ...
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Full-text available
As human populations continue to expand across the world, the need to understand and manage wildlife populations within the wildland–urban interface is becoming commonplace. This is especially true for large carnivores as these species are not always tolerated by the public and can pose a risk to human safety. Unfortunately, information on wildlife species within the wildland–urban interface is sparse, and knowledge from wildland ecosystems does not always translate well to human‐dominated systems. Across western North America, cougars (Puma concolor) are routinely utilizing wildland–urban habitats while human use of these areas for homes and recreation is increasing. From 2007 to 2015, we studied cougar resource selection, human–cougar interaction, and cougar conflict management within the wildland–urban landscape of the northern Front Range in Colorado, USA. Resource selection of cougars within this landscape was typical of cougars in more remote settings but cougar interactions with humans tended to occur in locations cougars typically selected against, especially those in proximity to human structures. Within higher housing density areas, 83% of cougar use occurred at night, suggesting cougars generally avoided human activity by partitioning time. Only 24% of monitored cougars were reported for some type of conflict behavior but 39% of cougars sampled during feeding site investigations of GPS collar data were found to consume domestic prey items. Aversive conditioning was difficult to implement and generally ineffective for altering cougar behaviors but was thought to potentially have long‐term benefits of reinforcing fear of humans in cougars within human‐dominated areas experiencing little cougar hunting pressure. Cougars are able to exploit wildland–urban landscapes effectively, and conflict is relatively uncommon compared with the proportion of cougar use. Individual characteristics and behaviors of cougars within these areas are highly varied; therefore, conflict management is unique to each situation and should target individual behaviors. The ability of individual cougars to learn to exploit these environments with minimal human–cougar interactions suggests that maintaining older age structures, especially females, and providing a matrix of habitats, including large connected open‐space areas, would be beneficial to cougars and effectively reduce the potential for conflict. This manuscript summarizes cougar–human interactions and cougar use of exurban landscapes along the Front Range of Colorado. In this manuscript, we show how cougars are using the urban–wildland interface, investigate where interactions are occurring, and discuss management options to mitigate conflict.
... Determining the influence of sex on mountain lion survival is difficult because of several factors influencing sex-specific harvest probabilities. Anderson et al. (2009) suggested that because male mountain lions generally exhibit larger daily movements, they become more susceptible to harvest, particularly by hunters aided by hounds. However, it is reasonable to think females may be more susceptible to boot hunters, predator callers, and opportunistic take because females generally make up a larger proportion of the population on the landscape (Logan and Sweanor, 2001). ...
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Mountain lions (Puma concolor) have many impacts on the ecosystems they inhabit, leading to both biological and social ramifications. Yet, due to the relatively recent natural recolonization by mountain lions of the Little Missouri Badlands Region of western North Dakota, detailed data regarding many aspects of this population have been lacking. We studied mountain lions occupying the Badlands Region to improve our understanding of the characteristics of the North Dakota mountain lion population. Our objectives included estimating annual survival rates, documenting sources of mortality in this population, and creating a population model using statistical population reconstruction (SPR) techniques. Between 2012 and 2016, the average annual survival rate estimated from known fate models was 0.456. Sex-specific annual survival was estimated at 0.589 for females and 0.259 for males. We recorded 17 cause-specific mortalities of marked mountain lions over the same 5 y period, plus the probable failures of two litters of marked dependent kittens. Results from our population model indicate annual population abundance estimates between 2005 and 2017 ranged from a low of 27 total mountain lions in 2005-2006 to a high of 165 in 2011-2012. The model indicated an increasing trend in total abundance between 2005 and 2012, then a reversal and sharp drop in abundance from 2012 to 2014, until the trend leveled with similar total abundances from 2014 to 2017. The results from our study can help inform current and future management decisions in North Dakota and may also provide insight for managers faced with potential mountain lion recolonization further eastward in the United States.
... With PopRecon, random effects can be added to either survival or hunter kill vulnerabilities, but not both, to account for annual stochasticity in one of these parameters. I chose random effects for hunter kills because successful cougar hunting is largely dependent upon suitable snow conditions for tracking, which vary considerably from year-to-year (Anderson et al. 2010). I combined males and females into 5 age classes including 1, 2, 3, and 4 year-olds, and pooled cougars > 5 years-of-age to reduce the effect of aging errors (Skalski et al. 2012b) and facilitate model fitting. ...
... Cougars are classified as a hunted game mammal across much of their current range (Anderson et al. 2010), and state and provincial managers should clearly define population and harvest objectives for cougars that do not threaten their long-term conservation. Anderson and Lindzey (2005) and Wolfe et al. (2016) found that monitoring hunter success rates and ages of female cougars harvested provided guidance on appropriate harvest rates to reduce or maintain cougar densities. ...
Article
Full-text available
Understanding bottom‐up, top‐down, and abiotic factors along with interactions that may influence additive or compensatory effects of predation on ungulate population growth has become increasingly important as carnivore assemblages, land management policies, and climate variability change across western North America. Recruitment and population trends of elk (Cervus canadensis) have been downward in the last 4 decades across the northern Rocky Mountains and Pacific Northwest, USA. In Oregon, changes in vegetation composition and land use practices occurred, cougar (Puma concolor) populations recovered from near‐extirpation, and black bear (Ursus americanus) populations increased. Our goal was to provide managers with insight into the influence of annual climatic variation, and bottom‐up and top‐down factors affecting recruitment of elk in Oregon. We conducted our research in southwestern (SW; Toketee and Steamboat) and northeastern (NE; Wenaha and Sled Springs) Oregon, which had similar predator assemblages but differed in patterns of juvenile recruitment, climate, cougar densities, and vegetative characteristics. We obtained monthly temperature and precipitation measures from Parameter‐elevation Regressions on Independent Slopes Model (PRISM) and estimates of normalized difference vegetation index (NDVI) for each study area to assess effects of climate and vegetation growth on elk vital rates. To evaluate the nutritional status of elk in each study area, we captured, aged, and radio‐collared adult female elk in SW (n = 69) in 2002–2005 and NE (n = 113) in 2001–2007. We repeatedly captured these elk in autumn (n = 232) and spring (n = 404) and measured ingesta‐free body fat (IFBF), mass, and pregnancy and lactation status. We fitted pregnant elk with vaginal implant transmitters (VITs) in spring and captured their neonates in SW (n = 46) and NE (n = 100). We placed expandable radio‐collars on these plus an additional 110 neonates in SW and 360 neonates in NE captured by hand or net‐gunning via helicopter and estimated their age at capture, birth mass from mass at capture, and sex. We monitored their fates and documented causes of mortality until 1 year of age. We estimated density of cougars by population reconstruction of captured (n = 96) and unmarked cougars killed (n = 27) and of black bears from DNA analysis of hair collected from snares. We found evidence in lactating females of nutritional limitations on all 4 study areas where IFBFautumn was below 12%, a threshold above which there are few nutritional limitations (9.8% [SE = 0.64%, n = 17] at Toketee, 7.9% [SE = 0.78%, n = 17] at Steamboat, 7.3% [SE = 0.33%, n = 46] at Sled Springs, and 8.9% [SE = 0.51%, n = 23] at Wenaha). In spring, of females known to have been lactating the previous autumn, 48% (SE = 3.3%, n = 56) had IFBFspring <2%, a level indicating severe nutritional limitations, compared to 20% (SE = 1.7%, n = 91) of those not lactating the previous autumn. These low levels of IFBFspring of lactating females likely resulted from a carry‐over effect of inadequate nutrition during summer and early autumn. We found a positive relationship between summer precipitation and IFBFautumn in NE, and that IFBFautumn of pregnant females was inversely related to birth date of their neonates the following spring (F1, 52 = 20.37, P < 0.001, R2adj = 0.27). Mean pregnancy rates of lactating females were below 0.90, a threshold indicating nutritional limitations, at Toketee (0.67, SE = 0.12, n = 15), Wenaha (0.70, SE = 0.10, n = 23), and Sled Springs (0.87, SE = 0.05, n = 47) but not Steamboat (0.93, SE = 0.07, n = 14). Of elk where we sampled femur fat during winter in NE, we saw evidence of imminent starvation in 3 of 21 juveniles (12%) with all 3 killed by cougars, and 2 of 12 adult elk (17%) that both died from non‐predation events. Birth mass was <13 kg for 6.5% and 2% of VIT neonates in SW and NE, respectively, a mass associated with reduced probability of survival in previous studies. Birth mass of VIT neonates was greater in Sled Springs ( = 18.3 kg, SD = 2.5, n = 59) than Steamboat ( = 16.3 kg, SD = 2.1, n = 21) or Toketee ( = 16.1 kg, SD = 2.8, n = 24) but not Wenaha ( = 17.1 kg, SD = 2.8, n = 36; F3, 132 = 7.63, P < 0.001). Median and mean birth date (29 May) for VIT neonates did not differ between regions (F1, 136 = 0.33, P = 0.56), but NE had greater variation around the mean, indicating a longer parturition interval. We documented 293 mortalities of juveniles across study areas and years, and predation was the proximate cause of mortality in 262 cases primarily from cougar (n = 203), black bear (n = 34), and other or unknown predation (n = 25). We also documented causes of mortality as unknown (n = 16), human‐caused (n = 8), and disease or starvation (n = 7). We recorded abandonment of 2 (1.4%) and predation mortality of 4 (2.7%) VIT neonates prior to being collared. We found 4‐fold differences between regions of subadult female and adult cougar densities (0.90–4.29/100 km2) and 2‐fold differences within study areas across years, with cougar density lower in SW than NE. Black bear densities varied from 15–20/100 km2 across our study areas. We estimated survival of neonates to 30 days, 16 weeks, and 12 months using known fates models in Program MARK. Survival of neonates born to females with VITs was associated with cougar density, IFBFspring, and female mass but not female age or neonate birth date or birth mass. Survival was higher for juveniles born to females with lower IFBF and mass in spring, opposite of what we predicted. In a post hoc analysis, we found females successful in raising their neonate to recruitment were more likely to be successful the following year compared to those not successful the previous year, which may explain this unexpected finding. As cougar density increased, survival of juveniles born to females of known nutritional condition declined. We conducted separate analyses of survival by region for all neonates captured to evaluate effects of climate, bottom‐up (but not maternal condition), and top‐down factors. In NE, juvenile survival was little affected by annual variation in climate but decreased as cougar densities increased and as birth date became later. For SW, survival was higher with less April–May precipitation and for later born neonates but less affected by cougar density than observed in NE. Across our 4 study areas, survival varied annually from 0.61 (SE = 0.08) to 1.00 during the first 30 days, 0.41 (SE = 0.11) to 0.90 (SE = 0.09) the first 16 weeks, and 0.18 (SE = 0.06) to 0.57 (SE = 0.11) through 12 months (recruitment) with survival higher in SW than NE. Survival of juvenile elk was inversely related to cougar density through 30 days (F1, 18 = 16.59, R2adj = 0.45, P < 0.001), 16 weeks (F1, 18 = 21.07, R2adj = 0.51, P < 0.001), and 12 months (F1, 11 = 18.94, R2adj = 0.60, P = 0.001). We found that as rates of cougar‐specific mortality increased, juvenile survival declined ( = −0.63, 95% CI = −0.84 to −0.42) suggesting cougar predation was partially additive mortality because the estimated regression coefficient was significantly less than 0 but greater than −1. We did not observe a similar relationship with rates of black bear‐specific mortality because the estimated regression coefficient overlapped 0, suggesting predation by black bears on juvenile elk was compensatory. Our results suggest that recruitment in NE but not SW was primarily limited by predation from cougars, which was partially additive mortality. Given that we observed nutritional limitations that influenced juvenile survival in all 4 study areas, we were unable to explicitly quantify how much of the cougar predation was additive mortality. Thus, we caution that a reduction in cougar density may not result in an equivalent increase in recruitment, and maintaining or enhancing summer and winter ranges of elk in our study areas is also vitally important for sustaining populations and distributions. In SW, where cougar densities were lower, maintaining, and enhancing existing elk habitat may be the only management option to improve recruitment. Given the differences we found between regions monitored, basing management on an incomplete understanding of causative factors affecting elk population dynamics may result in ineffective actions to address low recruitment. © 2018 The Authors. Wildlife Monographs Published by Wiley Periodicals, Inc. on behalf of The Wildlife Society. La comprensión de los factores de abajo hacia arriba, de arriba hacia abajo y abióticos, junto con las interacciones que pueden influir en los efectos aditivos o compensatorios de la depredación sobre el crecimiento de la población ungulada, se ha vuelto cada vez más importante como asociaciones de carnívoros, políticas de manejo de la tierra y cambios en la variabilidad del clima en el oeste de América del Norte. El reclutamiento y las tendencias poblacionales de los alces (Cervus canadensis) han disminuido en las últimas 4 décadas en el norte de las Montañas Rocosas del Norte y el Pacífico Noroeste, EE. UU. En Oregón, se produjeron cambios en la composición de la vegetación y en las prácticas de uso de la tierra, las poblaciones de pumas (Puma concolor) se recuperaron de la casi extirpación y aumentaron las poblaciones de osos negros (Ursus americanus). Nuestro objetivo era proporcionar a los gerentes información sobre la influencia de la variación climática anual y los factores de abajo hacia arriba y de arriba hacia abajo que afectan el reclutamiento de alces en Oregón. Llevamos a cabo nuestra investigación en el suroeste (SW, Toketee y Steamboat) y en el noreste (NE, Wenaha y Sled Springs) Oregon, que tenían ensamblajes de depredadores similares pero diferían en los patrones de reclutamiento juvenil, clima, densidades de puma y características vegetativas. Obtuvimos medidas mensuales de temperatura y precipitación a partir de Regresiones de Elevación de Parámetros en el Modelo de Pendientes Independientes (PRISM) y estimaciones del Índice de Vegetación de Diferencia Normalizada (NDVI) para cada área de estudio para evaluar los efectos del crecimiento del clima y la vegetación en las tasas vitales de alces. Para evaluar el estado nutricional de los alces en cada área de estudio, fueron capturados, envejecidos y collares con radio fueron colocados en las alces hembras adultas en SW (n = 69) en 2002 − 2005 y NE (n = 113) en 2001 − 2007. Capturamos repetidamente estos alces en otoño (n = 232) y primavera (n = 404) y medidos de la grasa corporal con exclusion de sustancias ingeridas (siglas en inglés, IFBF), masa, y estado de embarazo y lactancia. Hemos equipado a los alces embarazadas con transmisores de implantes vaginales (VIT) en primavera y capturamos a sus neonatos en SW (n = 46) y NE (n = 100). Colocamos collares con radio expandibles en estos, además de 110 neonatos adicionales en SW y 360 neonatos en NE capturados a mano o con un arma en helicóptero y estimamos su edad de captura, la masa de nacimiento de la masa de captura y el sexo. Monitoreamos sus destinos y documentamos las causas de mortalidad hasta el año de edad. Estimamos la densidad de los pumas mediante la reconstrucción de la población de los pumas capturados (n = 96) y no marcados (n = 27) y de los osos negros del análisis de ADN del cabello recogido de las trampas. Encontramos evidencia en las hembras en período de lactancia de limitaciones nutricionales en las 4 áreas de estudio donde IFBFautumn estaba por debajo del 12%, un umbral superior por el cual hay pocas limitaciones nutricionales [9.8% (SE = 0.64%, n = 17) en Toketee, 7.9% (SE = 0.78%, n = 17) en Steamboat, 7.3% (SE = 0.33%, n = 46) en Sled Springs, y 8.9% (SE = 0.51%, n = 23) en Wenaha]. En primavera, de las hembras que se sabe que estuvieron lactando el otoño anterior, 48% (SE = 3.3%, n = 56) tuvieron IFBFspring <2%, un nivel que indica limitaciones nutricionales graves, en comparación con 20% (SE = 1.7%, n = 91) de los que no lactando el otoño anterior. Estos bajos niveles de IFBFspring de hembras lactantes probablemente resultado de un efecto de arrastre de una nutrición inadecuada durante el verano y principios de otoño. Encontramos una relación positiva entre la precipitación de verano y el IFBFautumn en NE, y que el IFBFautumn de hembras embarazadas se relacionó inversamente con la fecha de nacimiento de sus neonatos en la primavera siguiente (F1,52 = 20.37, P < 0.001, Radj 2 = 0.27). Las tasas de embarazo promedio de las hembras lactantes fueron inferiores a 0.90, un umbral que indica limitaciones nutricionales, en Toketee (0.67, SE = 0.12, n = 15), Wenaha (0.70, SE = 0.10, n = 23) y Sled Springs (0.87, SE = 0.05, n = 47), pero no Steamboat (0.93, SE = 0.07, n = 14). La grasa de fémur de 3 de 21 juveniles (12%) matado por puma y 2 de 12 alces adultos (17%) que murieron por eventos de no depredación a fines del invierno en NE mostró evidencia de inminente inanición. La masa al nacer fue <13 kg para 6.5% y 2% de los neonatos con VIT en SW y NE, respectivamente, una masa asociada con una probabilidad reducida de supervivencia en estudios previos. La masa al nacer de los neonatos con VIT fue mayor en Sled Springs ( = 18.3 kg, SD = 2.5, n = 59) que Steamboat ( = 16.3 kg, SD = 2.1, n = 21) o Toketee ( = 16.1 kg, SD = 2.8, n = 24) pero no Wenaha ( = 17.1 kg, SD = 2.8, n = 36; F3,132 = 7.63, P < 0.001). La mediana y media fecha de nacimiento (29 de mayo) para los neonatos con VIT no difirió entre las regiones (F1,136 = 0.33, P = 0.56), pero NE tuvo una mayor variación alrededor de la media, lo que indica un intervalo de parto más prolongado. Documentamos 293 muertes de juveniles en áreas de estudio y años, y la depredación fue la causa próxima de mortalidad en 262 casos principalmente de puma (n = 203), oso negro (n = 34) y otra depredación desconocida (n = 25). También se documentaron las causas de mortalidad por causas desconocidas (n = 16), causadas por humanos (n = 8) y enfermedad / inanición (n = 7). Registramos el abandono de 2 (1.4%) y la mortalidad por depredación de 4 (2.7%) neonatos con VIT antes de ser colocado el collar con radio. Encontramos diferencias cuádruples las densidades de pumas hembras sub‐adultas y adultas (0.90 a 4.29 por 100 km2) entre las regiones y diferencias doble en las áreas de estudio a través de los años con una densidad de puma más baja en SW que en NE. Las densidades de osos negros variaron de 15 a 20 por 100 km2 en nuestras áreas de estudio. Estimamos la supervivencia de los neonatos a 30 días, 16 semanas y 12 meses utilizando modelos de destinos conocidos en el Programa MARK. La supervivencia de los neonatos nacidos de hembras con VIT se asoció con la densidad del puma, la IFBFspring, y la masa hembra, pero no con la edad hembra, o la fecha de nacimiento del neonato o la masa de nacimiento. Encontramos que la supervivencia fue más alta para los juveniles nacidos de hembras con un IFBF y una masa más bajos en primavera, lo contrario de lo que predijimos. En un análisis post‐hoc, encontramos que las hembras que tuvieron éxito en criar a sus recién nacidos a reclutamiento tuvieron más probabilidades de tener éxito el año siguiente, en comparación con las que no tuvieron éxito el año anterior, lo que puede explicar este hallazgo inesperado. A medida que la densidad del puma aumentó, la supervivencia de los juveniles nacidos de hembras de afección nutricional conocida disminuyó. Realizamos análisis separados de supervivencia por región para todos los neonatos capturados para evaluar los efectos del clima, de abajo hacia arriba (pero no de la condición materna) y de los factores de arriba hacia abajo. En NE, la supervivencia juvenil se vio poco afectada por la variación anual en el clima, pero disminuyó a medida que aumentaban las densidades de puma y la fecha de nacimiento se hacía más tarde. Para SW, la supervivencia fue mayor con menos precipitaciones de Abril a Mayo y para los neonatos nacidos tarde, pero menos afectada por la densidad de pumas que la observada en NE. En nuestras 4 áreas de estudio, la supervivencia varió anualmente de 0.61 (SE = 0.08) − 1.00 durante los primeros 30 días, 0.41 (SE = 0.11) − 0.90 (SE = 0.09) las primeras 16 semanas, y 0.18 (SE = 0.06) − 0.57 (SE = 0.11) a 12 meses (reclutamiento) con una supervivencia mayor en SW que en NE. La supervivencia de los alces juveniles se relacionó inversamente con la densidad del puma durante 30 días (F1,18 = 16.59, Radj2 = 0.45, P < 0.001), 16 semanas (F1,18 = 21.07, Radj2 = 0.51, P < 0.001) y 12 meses (F1,11 = 18.94, Radj2 = 0.60, P = 0.001). Encontramos que a medida que aumentaban las tasas de mortalidad por puma, la supervivencia juvenil disminuyó, ( = −0.63, CI 95% = −0.84–‐0.42), sugiriendo depredación de puma era una mortalidad parcialmente aditiva porque el coeficiente de regresión estimado era significativamente menor que 0 pero mayor que −1. No observamos una relación similar con las tasas de mortalidad específica del oso negro porque el coeficiente de regresión estimado se superpuso a 0, sugiriendo depredación por osos negros en alces juveniles fue compensatoria. Nuestros resultados sugieren que el reclutamiento en NE pero no en SW estuvo principalmente limitado por la depredación de los pumas, que fue una mortalidad parcialmente compensatoria. Dado que se observaron limitaciones nutricionales en las 4 áreas de estudio que influyeron en la supervivencia juvenil, no pudimos cuantificar explícitamente cuánta de la depredación de puma era la mortalidad aditiva. Por lo tanto, advertimos que una reducción en la densidad del puma puede no resultar en un aumento equivalente en el reclutamiento, y mantener o mejorar los rangos de alces en verano e invierno en nuestras áreas de estudio también es de vital importancia para el mantenimiento de poblaciones y distribuciones. En SW, donde las densidades de pumas eran menores, mantener y mejorar el hábitat existente puede ser la única opción de manejo y gestión para mejorar el reclutamiento. Dadas las diferencias que encontramos entre las regiones monitoreadas, basar el manejo y gestion en una comprensión incompleta de los factores causales que afectan la dinámica de la población de alces puede resultar en acciones ineficaces para abordar el reclutamiento bajo. Comprendre les facteurs ascendants, descendants et abiotiques ainsi que les relations pouvant influencer les effets additifs ou compensatoires de la prédation sur la croissance des populations d'ongulés est devenu de plus en plus important compte tenu de la transformation de l'assemblage des carnivores, des changements dans les politiques en matière d'aménagement du territoire et des variations dans le climat dans l'ouest de l'Amérique du Nord. Le recrutement et les tendances sont à la baisse chez les populations de wapitis (Cervus canadensis) depuis les 4 dernières décennies dans le Pacifique Nord‐Ouest et dans le nord des montagnes Rocheuses aux États‐Unis. Dans l’état de l'Oregon, des changements survenus dans la composition végétale et dans les pratiques d'utilisation des terres ont entraîné un rétablissement des populations de cougar (Puma concolor) qui étaient en voie de disparition et une augmentation des populations d'ours noirs (Ursus americanus). Notre but était d'aider les gestionnaires à mieux comprendre l'effet des variations climatiques annuelles et des facteurs ascendants et descendants sur le recrutement des wapitis en Oregon. Nous avons réalisé nos travaux dans le sud‐ouest (SO; Toketee et Steamboat) et dans le nord‐est (NE; Wenaha et Sled Springs) de l'Oregon. Ces régions avaient des assemblages de prédateurs similaires, mais présentaient des différences dans le recrutement des juvéniles, le climat, les densités de cougar et les caractéristiques de la végétation. Pour chaque région à l’étude, nous avons obtenu des données de température et de précipitation à l'aide de PRISM (Parameter‐elevation Regressions on Independent Slopes Model) et des estimations de l'indice de végétation par différence normalisée (IVDN) afin d’évaluer les effets du climat et de la croissance de la végétation sur les indices vitaux des wapitis. Pour évaluer l’état nutritionnel des wapitis de chaque région, nous avons capturé des femelles adultes dans le SO (n = 69) en 2002−2005 et dans le NE (n = 113) en 2001−2007, établi leur âge et installé des colliers radio‐émetteurs. Nous avons capturé ces wapitis à plusieurs reprises en automne (n = 232) et au printemps (n = 404) et mesuré leur taux de lipides sans ingesta (TLSI), leur masse corporelle, leur état de gestation et leur état de lactation. Au printemps, nous avons équipé les femelles en gestation d'un émetteur intra‐vaginal et capturé leurs nouveau‐nés dans le SO (n = 46) et dans le NE (n = 100). Nous avons installé des colliers radio‐émetteurs extensibles sur ces femelles et sur 110 nouveau‐nés dans le SO et sur 360 nouveau‐nés dans le NE capturés à la main ou avec des filets lancés à partir d'un hélicoptère. Lors de la capture des nouveau‐nés, nous avons estimé leur âge, leur masse à la naissance et leur sexe. Nous avons suivi leur développement et documenté les causes de mortalité jusqu’à l’âge de 1 an. Nous avons estimé la densité des cougars en reconstituant la population à partir de cougars capturés (n = 96) et de cougars tués non marqués (n = 27) et la population d'ours noirs en analysant l'ADN de poils recueillis dans des collets de trappage. Nous avons constaté des limites nutritionnelles chez des femelles en lactation dans les 4 régions à l’étude. Ces femelles avaient un TLSIautomne inférieur à 12%, un seuil au‐delà duquel il y a peu de limites nutritionnelles (9,8% [erreur‐type = 0,64%, n = 17] à Toketee, 7,9% [erreur‐type = 0,78%, n = 17] à Steamboat, 7,3% [erreur‐type = 0,33%, n = 46] à Sled Springs, et 8,9% [erreur‐type = 0,51%, n = 23] à Wenaha). Au printemps, 48% (erreur‐type = 3,3%, n = 56) des femelles en lactation observées l'automne précédent avaient un TLSIprintemps <2%, un niveau sous lequel les limites nutritionnelles sont importantes, comparativement à 20% (erreur‐type = 1,7%, n = 91) des femelles qui n’étaient pas en lactation l'automne précédent. Ces faibles TLSIprintemps chez les femelles en lactation étaient probablement le résultat d'un effet d'entraînement d'une alimentation inadéquate en été et au début de l'automne. Nous avons remarqué une relation positive entre les précipitations en été et le TLSIautomne dans le NE et une relation inverse entre le TLSIautomne des femelles en gestation et la date de naissance de leurs nouveau‐nés au printemps suivant (F1, 52 = 20,37, P < 0,001, R2corr = 0,27). Les taux de gestation moyens des femelles en lactation étaient inférieurs à 0,90, un seuil indiquant des limites nutritionnelles, à Toketee (0,67, erreur‐type = 0,12, n = 15), Wenaha (0,70, erreur‐type = 0,10, n = 23) et Sled Springs (0,87, erreur‐type = 0,05, n = 47), mais pas à Steamboat (0,93, erreur‐type = 0,07, n = 14). Le gras du fémur de 3 wapitis juvéniles parmi 21 (12%) tués par des cougars et 2 wapitis adultes parmi 12 (17%) morts à la fin de l'hiver dans le NE suite à des événements qui n’étaient pas reliés à la prédation a révélé une famine imminente. La masse à la naissance de 6,5% et de 2% des nouveau‐nés de femelles équipées d'un émetteur intra‐vaginal était <13 kg, dans le SO et le NE respectivement. Des études antérieures ont établi que cette valeur de masse corporelle était associée à une probabilité de survie moindre. La masse à la naissance des nouveau‐nés de femelles équipées d'un émetteur intra‐vaginal était plus élevée à Sled Springs ( = 18,3 kg, écart‐type = 2,5, n = 59) qu’à Steamboat ( = 16,3 kg, écart‐type = 2,1, n = 21) ou qu’à Toketee ( = 16,1 kg, écart‐type = 2,8, n = 24), mais pas à Wenaha ( = 17,1 kg, écart‐type = 2,8, n = 36; F3, 132 = 7,63, P < 0,001). La date de naissance moyenne et médiane (29 mai) des nouveau‐nés de femelles équipées d'un émetteur intra‐vaginal ne différait pas entre les régions (F1, 136 = 0,33, P = 0,56), mais le NE présentait de plus grandes variations autour de la moyenne, indiquant ainsi une période de mise bas plus longue. Nous avons documenté 293 mortalités de juvéniles dans les régions à l’étude pendant toute la durée de l’étude. La prédation était la cause directe de mortalité dans 262 cas. Les principaux prédateurs étaient les cougars (n = 203), suivis des ours noirs (n = 34), et des prédateurs autres ou inconnus (n = 25). Nous avons également documenté les mortalités de causes inconnues (n = 16), celles de nature humaine (n = 8), et celles reliées à la maladie ou à la famine (n = 7). Nous avons documenté 2 juvéniles abandonnés (1,4%) et 4 nouveau‐nés (2,7%) de femelles équipées d'un émetteur intra‐vaginal qui étaient morts par prédation avant que nous puissions leur installer un collier émetteur. Nous avons constaté des différences dans la densité des cougars adultes et des jeunes femelles cougars de l'ordre de 4 fois entre les régions (0,90–4,29/100 km2) et de l'ordre de 2 fois au sein même des régions à l’étude au cours des années, avec une densité de cougars plus faible dans le SO que dans le NE. La densité des ours noirs a varié de 15–20/100 km2 dans les régions à l’étude. Nous avons estimé le taux de survie des nouveau‐nés après 30 jours, 16 semaines et 12 mois à l'aide de modèles connus dans le programme MARK sur le développement des individus. Le taux de survie des nouveau‐nés de femelles équipées d'un émetteur intra‐vaginal était relié à la densité des cougars, au TLSIprintemps et à la masse de la femelle, mais pas à l’âge de la femelle, la date de naissance du nouveau‐né ou sa masse à la naissance. Le taux de survie était plus élevé chez les juvéniles nés de femelles ayant un TLSI plus bas et une masse moins élevée au printemps, contrairement à nos prédictions. Dans un analyse post hoc, nous avons constaté que les femelles qui avaient réussi à élever leur nouveau‐né jusqu'au stade du recrutement étaient plus susceptibles de réussir l'année suivante comparativement à celles qui n'avaient pas réussi l'année précédente, ce qui pourrait expliquer ce résultat inattendu. Le taux de survie des juvéniles nés de femelles dont la condition nutritionnelle était connue déclinait au fur et à mesure qu'augmentait la densité des cougars. Nous avons effectué des analyses distinctes du taux de survie par région de tous les nouveau‐nés capturés afin d’évaluer les effets du climat et des facteurs ascendants (mais pas la condition maternelle) et descendants. Dans le NE, le taux de survie des juvéniles était peu influencé par les variations annuelles du climat, mais diminuait au fur et à mesure de l'augmentation de la densité des cougars et du recul de la date de naissance. Dans le SO, le taux de survie était plus élevé lorsqu'il y avait moins de précipitations en avril‐mai et que les nouveau‐nés naissaient plus tard, mais moins influencé par la densité des cougars que dans le NE. Dans nos 4 régions à l’étude, le taux de survie a varié annuellement de 0,61 (erreur‐type = 0,08) à 1,00 au cours des 30 premiers jours, de 0,41 (erreur‐type = 0,11) à 0,90 (erreur‐type = 0,09) au cours des 16 premières semaines, et de 0,18 (erreur‐type = 0,06) à 0,57 (erreur‐type = 0,11) sur une période de 12 mois (recrutement) avec un taux de survie plus élevé dans le SO que dans le NE. Le taux de survie des juvéniles était inversement lié à la densité des cougars pendant 30 jours (F1, 18 = 16,59, R2corr = 0,45, P < 0,001), 16 semaines (F1, 18 = 21,07, R2corr = 0,51, P < 0,001), et 12 mois (F1, 11 = 18,94, R2corr = 0,60, P = 0,001). Nous avons constaté que le taux de survie des juvéniles diminuait au fur et à mesure de l'augmentation du taux de mortalité attribué spécifiquement aux cougars ( = −0,63, IC 95% = −0,84 à −0,42), ce qui semble indiquer que la prédation par le cougar représentait une mortalité partiellement additive, car le coefficient de régression était considérablement plus bas que 0, mais plus grand que −1. Nous n'avons pas observé une relation similaire dans le taux de mortalité attribué spécifiquement aux ours noirs, car le coefficient de régression estimé dépassait 0, ce qui semble indiquer que la prédation par les ours noirs représentait une mortalité compensatoire. Nos résultats semblent indiquer que la prédation par les cougars constituait la principale limite au recrutement dans le NE, mais pas dans le SO, ce qui représentait une mortalité partiellement compensatoire. Compte tenu que nous avons observé des limites nutritionnelles qui influençaient le taux de survie des juvéniles dans les 4 régions à l’étude, nous n'avons pas été en mesure de quantifier explicitement le pourcentage de prédation par les cougars qui correspondait à une mortalité additive. Nous tenons donc à souligner qu'une réduction dans la densité des cougars peut ne pas donner lieu à une augmentation équivalente dans le recrutement. Maintenir et améliorer les aires de répartition estivales et hivernales des wapitis dans nos régions à l’étude est aussi d'une importance cruciale pour soutenir les populations et les distributions. Dans le SO, là où les densités de cougars étaient les plus faibles, il est possible que le maintien et l'amélioration de l'habitat actuel des wapitis soient les seules options de gestion pour augmenter le recrutement. Compte tenu des différences que nous avons observées entre les régions étudiées, prendre des décisions de gestion sans bien comprendre les facteurs qui influencent la dynamique des populations de wapitis peut donner lieu à des actions inefficaces pour corriger un faible recrutement. Cougar predation explained much of the variation in juvenile elk survival but was a partially compensatory source of mortality because of nutritional limitations that influenced pregnancy rates, overwinter survival, and susceptibility to predation (e.g., birth date). Our results highlight the importance of simultaneously considering top‐down and bottom‐up factors when assessing juvenile recruitment and growth rates of elk populations.
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