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September 2001

THE ROLE OF PREDATION IN WILDLIFE POPULATION DYNAMICS

Eric M. Gese
Utah State University, eric.gese@usu.edu

Frederick F. Knowlton
Utah State University, Logan, UT

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Gese, Eric M. and Knowlton, Frederick F., "THE ROLE OF PREDATION IN WILDLIFE POPULATION
DYNAMICS" (2001). USDA National Wildlife Research Center - Staff Publications. 542.
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 THE ROLE OF PREDATIOS IN WILDLIFE POPULATION
DYNAMICS

E R I C ?vI. GESE, National Wildlife Research Center, Department of Fisheries and Wildlife,
Utah State University, Logan, UT 84322-5295.

F R E D E R I C K F. WOWLTON, National Wildlife Research Center, Department of

Fisheries and Wildlife, Utah State University, Logan, UT 84322-5295.

Abstract: The role predation plays in the dynamics of prey populations is controversial. Our
understandings of predator-prey relationships is complicated by a multitude of factors in the
environment and a general lack of knowledge of most ecological systems. Various other
factors, besides predation, may regulate or limit prey populations, and various factors
influence the degree to which predation affects prey populations. Furthermore, some factors
may create time lags, or even cause generational effects, that go unnoticed. Herein, we
review the role of predation in wildlife population dynamics, some of the factors influencing
predator-prey interactions, and attempt to indicate where the professional debate currently
is focused and where it may need to go to enhance our understanding of predator-prey
interactions.

Predation has been defined as on ungulate density. Assessments of the

individuals of one species eating living importance that wolf (Canis lupus)
individuals of another species (Taylor predation plays in regulating or limiting
1984). The role of predators in the moose (Alces alces) populations varies
population dynamics of prey species has among biologists. Interactions among
been investigated for decades, yet moose, forage, and climate have been
determining whether or not predators limit postulated to determine moose density
or regulate a prey population remains (Peterson et al. 1984, Mech et al. 1987,
controversial within the scientific Thompson and Peterson 1988). Bergerud et
profession (e.g., Erlinge et al. 1984, Kidd al. (1983), Bergemd and Snider (1988), and
and Lewis 1987, Newsome et al. 1989, Van Ballenberghe and Ballard (1994)
Sinclair 1989, Sinclair et al. 1990, Messier considered predation a major limiting factor
1991, 1994, Skogland 1991, Boutin 1992, of moose because moose density was
Pech et al. 1992). Much of the debate generally below forage canying capacity.
results from the multitude of competing Messier and CrSte (1985) and Messier
variables, including predation, that influence (1991) argued that moose and predator
demographics of prey species and the interactions were complex and that the
difficulty of conducting large-scale, long- effect of predation varied from density-
term studies with some degree of control or dependent to inversely density-dependent
replication. In a review of studies involving over the range of moose densities resulting
predation on ungulates, Connolly (1978) in population cycles, multiple stable states,
reported that 45 studies suggested predation and predator pits. Skozland (1991)
was a limiting factor, while 27 studies suggested that ths existin: data was
indicarsd predation \vas not a limitin: factor inconclusive with rsgaids to predation

Gese, E. M., and F. F. Knowlton. 2001. The role of predation in wildlife population dynamics. Pages 7-25 in The
Role of Predator Control as a Tool in Game Management. Edited by T. F. Ginnetr and S. E. Henke.
Texas Agricultural Research and Extension Center. San Angelo. Texas.
regulating moose, while Boutin (1992) 'attack, capture, or consume' commonly
argued that wolf predation as a limiting found within the predation literature. The
factor on moose populations was not relationship between the kill rate by a
supported. predator and prey density is termed the
"functional response." Lotka (1925) and
In this paper, we will attempt to Volterra (1926) provided initial
provide a foundation on predator-prey mathematical descriptions of predator-prey
theory, describe some studies illustrating interactions, which assumed that the number
the roles that predators and other variables of prey captured increased in direct
can play in the dynamics of wildlife proportion to the number of predators.
populations (mainly from the camivore- Nicholson and Bailey (1935) proposed the
ungulate literature), and suggest reasons relationship was curvilinear with the kill rate
why the debate over the influence predators decreasing as predator satiation sets an
have on prey populations continues. It is upper limit to food consumption.
not our intent to critically review all Subsequently, Holling (1959) described 3
predator-prey studies, but to use certain types of functional responses (Fig. 1). A
studies to illustrate aspects of predator-prey Type I functional response occurs when the
relationships. kill rate per predator is directly proportional
to prey density. In the Type I1 response, the
TERMINOLOGY kill rate is limited at higher prey densities by
satiation of the predator, and is thus
For our discussions of predator-prey curvilinear. The Type 111 functional
relations to be fruitful, we need to clarify response is sigmoid in shape with a lag in
some terminology. We also should kill rate at low prey density due to low
recognize that many of the initial terms and hunting efficiency or absence of a search
early theories concerning the role predators image and an upper limit set by predator
play in limiting or regulating prey satiation. Type I1 functional responses have
populations were developed by been documented between wolves and
entomologists examining relationships moose (Messier 1994; Fig. 2), wolves and
between numbers of parasites needed to caribou (Rangifer tarandzrs) (Dale et al.
regulate invertebrate pest species on 1994), as well as coyotes (Canis latrans)
agricultural crops. The terms regulating and and black-tailed jackrabbits (Lepus
limiting have often been used calforniczrs) (Stoddart et al. 2001). In
interchangeably, with regulation defined as addition to functional responses, Morris et
"any density-dependent process that tends to al. (1958) demonstrated a "numerical
stabilize population numbers over time. The response" in which predator numbers
process that causes the change(s) in increase in response to increasing prey
population size is termed limitation" abundance. This numerical response may
(Skogland 1991). We consider a limiting be from reproduction or immigation.
factor to be any mortality factor that reduces Numerical responses of coyotes to changes
the rate of population growth (Ballard et al. in black-tailed jackrabbit (Kno\vlron and
2001). We also will try to adhere to using Stoddart 1992) and snowshoe hare (L.
the term 'kill' to denote the essential uniericnnla) abundance (O'Donoghue et a[.
component of a predator's impact upon 1997) have been documented- Messier
prey, rather than the ambiguous terms (1991) found a numerical response of
wolves to changes in moose density (Fig. 3). CONSTRAINTS OF PRED.4TOR AND
The combination of the functional and PREY
numerical responses represents the "total
response." The total response may cause the Evolution has placed constraints of
predation rate to be density dependent at both predatory and prey species, with
low prey density and inversely density obvious implications for the relationships
dependent at high prey density (Holling between them. In general, comparative
1959, Messier 1994). In a compilation of body size, strength, speed, and agility
studies, Messier (1994) illustrated the total dictate a predator's ability to kill particular
response of wolves to changing moose prey, while similar constraints on prey
density. define which predators pose a threat to
them. For example, predation by swift
Other terms commonly used when foxes (V~rlpesvelo,~)on adult pronghorn
describing predator-prey relationships are antelope (A~ltilocnprcramerica~ln)is highly
"compensatory" and "additive" mortality. improbable, even though both species
Ballard et al. (2001) defined additive occupy the same prairie habitat. Similarly,
mortality as occurring when the "additional the body size and defensive capabilities of
risk of death does not cause reductions in voles (Microtzrs spp.) are no match for the
other forms of mortality, but rather increases size and agility of coyotes, hence voles must
overall mortality rate." On the other hand, rely upon other survival strategies. Such
for compensatory mortality, the "additional physical and behavioral characteristics, or
risk of death causes a reduction in other constraints, have developed over extensive
forms of mortality so that overall mortality periods and represent an "evolutionary race"
either does not change or is less than it between predator and prey. Some physical
would be if additive." Kunkel and Pletscher abilities, e.g., the speed of pronghorn
(1999) suggested that predation on cewids antelope, may even represent residual
by several predatory species (mainly wolf developments from interactions with
and cougar, Puma concolor) was additive in predators now extinct (Byers 1997).
northwestern Montana. Two terms also
worthy of definition are "obligate" and HUNTING STRATEGIES
"facultative" predator. An obligate predator
is one that specializes on one primary prey The process within which predators
species. Hence changes in the levels of the seek and kill has important implications to
primary prey will generally influence a their impact upon prey. In the case of
numerical change in the obligate predator. obligate predators, relief from predatory
In contrast, a facultative predator is a dietary pressures on the prey will certainly occur
generalist that switches among prey species with the numerical decline in predators
and is thus buffered by changes in when prey become too scarce to support the
abundance of any one prey species. A predators. Among mammalian systems, this
facultative predator in a multi-prey system is perhaps best exemplified by the patterns
can limit one prey species to low levels in abundance of lynx (Lyt1.r cntzcriiensis) and
because other prey maintain the predator snowshoe hare, or black-footed ferrets
population. (ibf~rsiela nigripes) and prairie dogs
(Ci.rio~g,s spp.). \Vhen facultative
predarors. ~viththeir abilities to switch from
one prey source to another or even from directly to the number of coyotes and
predacious to omnivorous diets, are inversely to the number of jackrabbits (the
involved, relief from predatory pressures coyote-jackrabbit ratio). If these scenarios
may not be forthcoming. When a prey are correct, a transition from a Type I to a
species comprises incidental portions of a Type I1 functional response occurs as
predator's diet, the killing rate may be a jackrabbits mature. In the relatively simple
matter of random encounter between ecological situation studied by Knowlton
predator and prey and be strictly a product and Stoddart (1992), there was an apparent
of the numerical abundances of both feed-back mechanism with the numerical
predator and prey (Hollings' Type I abundance of coyotes dictated by the
functional response). This type of situation abundance of their principle prey, adult
can be particularly hazardous to prey species jackrabbits. Thus they propose predatory
that are scarce. mechanisms that might partially explain the
cyclic nature of some predator-prey
As the relative importance of a prey interactions.
species increases in a predator's diet, or the
effort needed to acquire the prey increases, In addition to a component of strict
the functional response assumes numerical abundance, the effect that
characteristics of a Type I1 hnctional facultative predators, which switch from one
response, with satiation placing an upper prey type to another, have on prey species
limit upon the killing rate by individual probably reflects a complex integration of
predators (for this discussion, we will ignore the relative abundance, efforts to capture,
events like surplus killing, food caching, and the quantity and quality of reward
etc.). In this case, the killing rate of the prey associated with each prey species.
is directly proportional to the number of Consequently, understanding the role of
predators but inversely proportional to the predation upon a prey species in this
abundance of prey. This is perhaps best situation requires an understanding of the
exemplified by iOlowlton and Stoddart's contexi within which it occurs. We
(1992) hypothesis that predation upon acknowledge there also are components
black-tailed jackrabbit nestlings by coyotes associated with habits, learning, and
may be a Type I functional response, being traditions, among individual predators, but
merely a matter of chance encounter those issues are beyond the scope we wish
because the frequency and size of reward is to present here.
insufficient for coyotes to actively hunt for
them. On the other hand, they suggest that FACTORS INFLUENCIKG PREY
predation upon adult jackrabbits is more POPULATIONS
likely a Type I1 functional response, with
coyotes actively hunting jackrabbits as a The role predation plays in wildlife
dietary staple, but because it requires population dynamics follows many of the
significant effort to capture adult constnlcts and theories established by earlier
jackrabbits, they are only hunted when researchers working on insects. However,
coyotes are h u n , ~ . In this case, satiation of as in the entomological debates of decades
the coyotes places an upper limit upon the past, the role of predation in the population
killin2 of adult jackrabbits. Hence the d!namics of wildlife. particularly ungulates,
fraction of adult jackrabbits killed is related is far from clear (Sinclair 1991. Skogland
 1991, Messier 1991, 1994; Boutin 1992, (Sinclair 1979). Once rinderpest was
Dale et al. 1994, Van Ballenberghe and eradicated, the wildebeest population tripled
Ballard 1994). Much of the conhsion arises in size from 1963 to 1974 (Sinclair 1979).
because predation is only one of many Competition among ungulate species also
factors influencing prey populations. may influence population levels. A theory
Growth rates (increasing or decreasing) of currently proposed for the mule deer ( 0 .
prey populations may be affected by habitat hemionris) decline in some areas of the
changes, severe weather (e.g., deep snow), western United States is competition with
starvation, diseases, predation, human elk (Cervlu elaphus) and white-tailed deer.
hunting, competition with other ungulates Equally disconcerting is the likelihood of
(native and domestic), changes in sex and hybridization between mule and white-tailed
age structure, as well as interacting deer, particularly in areas where habitat
combinations of these factors (Ballard et al. modification increases the probability of
2001). For example, density-dependent interspecific hybridization (Hornbeck and
food limitation and density-independent Mahoney 2000). Competition with
adverse weather have been implicated as livestock also has been implicated as a result
factors regulating the numbers of reindeer in of cattle removing winter forage for mule
the arctic tundra (Skogland 1985, 1990). deer and elk in the western United States.
Gates et al. (1986) concluded that food Increased urbanization has resulted in loss
limitation and snow conditions regulated of suitable habitat, especially wintering
barren-ground caribou (R. t. groenlandicus) ranges, for many ungulate populations,
in northern Canada. Mech et al. (1987) and although land conversion to agculture may
McRoberts et al. (1995) reported the benefit some white-tailed deer populations.
cumulative effect of snow depth over 3 Thus, while many factors affect prey
winters influenced population parameters of population levels, for purposes of this paper,
moose on Isle Royale and white-tailed deer let us focus on the effect of predation.
(Odocoi!ezts virginianz~~) in Minnesota. In
a counterpoint, Messier (1991) postulated FACTORS INFLUENCING
that competition for food, not snow depth, PREDATION
had a regulatory effect on moose, and that
deer density and deer population growth was Skogland (1991) identified 9 factors
inversely related to wolf density; with snow that may influence predation: habitat
depth not a significant factor. Given the heterogeneity, prey refugia, nomadism
same information, various researchers (temporallspatial availability of prey), buffer
provide differing interpretations of the data. zones for prey, synchrony of the birthing
It seems unlikely the debate over the season and aggregation at birthing, prey size
hierarchy of factors influencing ungulate and age vulnerability, availability of
population dynamics will soon be resolved. alternate prey, the ratio between the
dominant prey species and alternative
Disease is another factor that may (buffer) prey, and the effects of
regulate some ungulate populations. The compensatory causes of mortality and the
introduction of rinderpest to the Serengeti effects of alternative predator species.
plains by domestic cattle caused high Several ofthese are related and interactions
mortality amons wildebeest (Connochaetes amon2 factors may cloiid our understanding
tnuril~ils) and buffalo (3.ncenis ccfler) of predator-prey systems Food is often a
limiting factor on ungulate populations Seip (1992) suggests that one caribou
(Sinclair 1979). Heterogeneity of habitat population was slowly increasing because
has been proposed to influence predation on its migratory behavior kept the caribou
prey populations. With increasing human separated from wolves and moose
modification of the landscape, prey throughout the year resulting in low wolf
populations become fra,mented, or isolated, predation. Fryxell et al. (1988) postulated
and more vulnerable to predators. Habitat that migratory ungulates on the Serengeti
degradation may increase predation of mink may escape predatory regulation by their
(Mustela vison) on water voles (Arvicola movements, while resident ungulate
terrestris) in England (Barreto et al. 1999). populations might be more vulnerable to the
Increased predation risk from habitat effects of predators. However, the seasonal
fragmentation has been implicated as cause migratory patterns observed for ungulates
for decline of game bird populations. In on the Serengeti are more likely due to
contrast, modification to urban landscapes changes in forage quality across the
may favor some prey species (i.e., white- landscape (Fryxell 1995), than predator
tailed deer), yet dissuade large carnivores avoidance. Another antipredator strategy
from these areas, forming a refuge from among ungulate species may result from
predatory pressures. In a more natural reproductive synchrony and aggregation
setting, Murie (1944) showed that Dall during and following the birthing process.
sheep (Ovis dalli) escaped wolfpredation by Reproductive synchrony and aggregation
using steep terrain and cliffs as refugia. during birthing can flood territorial
Similarly, Ferguson et al. (1988) suggested predators to the point that only a small
that one population of woodland caribou portion of the reproductive effort falls prey
reduced predation risk from wolves by to predators (Estes 1976). Although,
residingon small islands. birthing synchrony is generally related to
environmental seasonality and the plant
The temporal and spatial availability growing season (Rutberg 1987). Perhaps an
of prey also influences predator-prey equally importani effect may result from the
relationships. In northeastern Minnesota, territorial nature of most carnivores, which
white-tailed deer use buffer zones between limits their ability to respond numerically to
wolf packs, which wolves avoid for fear of aggregations of prey, especially during
intraspecific strife with neighboring packs periods of heightened vulnerability, as in the
(Mech 1977). Nelson and Mech (1981) case of yarding among white-tailed deer or
suggest that wolf predation regulates deer winter concentrations of black-tailed
numbers, but the buffer zones between wolf jackrabbits (Smith 1987).
territories allow sufficient numbers of deer
to survive and these deer can reoccupy wolf Prey vulnerability is regarded as a
pack territories when wolf numbers are low. major factor in predator-prey interactions.
Subsequently, Nelson and Mech ( 1 9 8 6 ~ ) Most predator-prey studies document how
reported that the effects of snow depth and predators target young and old animals,
vulnerability was the main factor regulating individuals in poor nutritional condition, or
deer numbers, rather than wolf predation. prey that are weakened by disease or
Migatory behavior is another mechanism physical abnormalities. In a classic study in
that reduces the effect of predation. In the Alaska. ,Murie (1944) rsported that wolves
LVells Gray caribou of British Columbia. killed Dall sheep that Ivere weak. diseased,
or very old. Mech (1966) reported that which could exacerbate the interaction
moose with heavy infestations of hydatid between a predator and another prey. In
cysts (Echinococcus grand~rlosus),calves, Montana, Hamlin et al. (1984) documented
and very old individuals were at greatest coyote predation as a major cause of mule
risk of wolf predation on Isle Royale. In deer fawn mortality. Hotvever, when
northeastern Minnesota, wolves killed a microtine rodents were abundant, mule deer
preponderance of fawns, old male deer, and fawn mortality was low. They further
individuals with abnormalities (Mech and concluded that vegetative production and
Frenzel1971, Mech and Kams 1977, Nelson winter snow cover may regulate microtine
and Mech 19866). Even the nutritional abundance, and thus fawn mortality rates.
condition of the mother and grandmother Kunkel and Pletscher (1999) reported that
may influence the vulnerability of first and where deer were present, the wolf-caused
second generation fawns to wolf predation mortality rate on moose was lower than in
(Mech et al. 1991). In Yellowstone areas where deer were absent. In contrast,
National Park, Gese and Grothe (1995) Fuller (1990) believed the effect of wolves
reported that adult elk killed by coyotes in on deer was exacerbated by the abundance
winter were in poor nutritional condition, of moose in north-central Minnesota.
based upon femur marrow fat indices. Stoddart et al. (2001) reported that coyotes
Studies also have documented the high responded numerically as black-tailed
vulnerability of new born fawns and calves jackrabbits increased during their 10-11 year
to predation by a suite of predators (e.g., cycle. When the jackrabbit population
Cook et al. 1971, Barrett 1984, Ballard et al. began to decline, coyotes switched to
1999). Vulnerability of a particular age domestic sheep and predation rates on lambs
group in the prey population can influence escalated.
population dynamics. Predators may
remove a high proportion of neonates The effects of alternative predators on
annual!y which may or may not affect a prey population can be substantial. In
population levels (Linnell et al. 1995), or if Alaska, Gasaway et al. (1992) identified
predators remove a high portion of the wolf and bear (Urstrs arctos) predation as a
reproductive cohort (e.g., prime-age does), major factor limiting moose at low densities.
the repercussions to the prey population may This multi predator system, with moose as
be substantial. the primary prey, held the moose population
well below carrying capacity. Kunkel and
Availability of alternate prey can Pletscher (1999) reported that with wolf
influence predator-prey interactions by recolonization in northwest Montana, the
either diluting or exacerbating the effects of full compliment of predators (wolves, bears,
a predator on their primary prey (Kunkel and cougars) brought about changes in the
and Pletscher 1999). Dilution could be abundance of some ungulate species. They
expected when alternate prey becomes more postulated that the mortality rate by all
vulnerable than the primary prey (Carbyn predators on the cen~idpopulations (elk,
1983, Potvin et al. 1988); in which case the deer, and moose) was additive.
'new' prey may become the primary prey,
buffering the former primary prey. In
contrast, the abundance of one prey may
cause a numerical response in the predator. Previous secrions identified many
factors that may influence prey populations "predator pit," which refers to the "narrow
-
and uredation rates. While these mizht add
confusion to predator-prey interactions,
band of densities between upper and lower
equilibrium points where ungulates can not
scientists have developed several theoretical increase because of density-dependent
models that allow testins of specific predation" (Ballard et al. 2001). The
hypotheses and predictions among multiple stable states model may exist in
competing models (e.g., Messier 1994). multi-predator and multi-prey systems.
These models, examine the role of predation
in ungulate population dynamics, are varied. Under the stable-limit cycle model,
Four models widely used in predator- prey populations may exhibit regular cycles
ungulate dynamics include: low-density of 30-40 years duration (Ballard et al. 2001).
equilibria, multiple stable states, stable-limit Ballard et al. (2001) reported that severe
cycles, and recurrent fluctuations (Boutin climate may influence the viability of young
1992, Van Ballenberghe and Ballard 1991, and the survival rates of young and adults.
Ballard and Van Ballenberghe 1997, Ballard "Predation is density independent during
et al. 2001). Similar models are presented population increases and inversely density-
by Messier (1994) and Sinclair and Arcese dependent during population declines"
(1995). Under the low-density equilibria (Ballard et al. 2001). Forage, climate, and
model, prey populations are regulated at low prey density all interact to regulate the prey
densities for long periods. The prey population. The stable-limit cycle model
population remains at a low density until typically exists in single predator and single
either a natural ohenomena or a decline in prey systems.
predator abundance (e.g., predator control)
allows the population
- - to grow.
- Limitation Sometimes a prey population
by food is not important because prey fluctuates and never reaches a state of
density never reaches carrying capacity. equilibrium. Prey densities change in
When predators recover from low numbers, response to changes in climate, forage
the prey population retums to a low density quality and quantity, and human harvest, but
(Ballard et al. 2001). This model generally the primary factor most often limiting prey
persists in systems with multiple species of density is predation (Ballard et al. 2001). At
predators and prey where predators subsist high prey density, predation is inversely
primarily on other prey. density-dependent. Prey may escape the
regulatory effect of predation and attain a
Under the multiple stable states model, higher density where ultimately food
a prey population is regulated by density- competition causes a population decline.
dependent predation at low prey density Inversely density-dependent predation may
until either a nahlral phenomena or predator accelerate or prolong the decline of the prey
removal reduces the predator population, population. Perturbations cause the prey
allowing the prey population to reach population to fluctuate without attaining a
carrying capacity and become regulated by predictable density (Ballard et al. 2001).
competition for food (Ballard et al. 2001). Both multi-predator and multi-prey systems
Food competition then regulates the prey and single predator-single prey systems may
pop~~lation at this higher equilibria e\.en exhibit recurrent fluctuations.
after ths predator popillation retums to its
forms; level. This model led to the tsm. It is unlikely that on? of th%s models
will describe any predator-prey system at all At the elevated population level, forage
times. Habitat conditions, human became suboptimal within the exclosure and
populations, climatic events, and other the general health of the deer declined.
factors are in constant flux. The acceptance Parasite loads increased, deer conceived
of one model, without periodically later, bucks retained velvet longer, males
reexamining the data on the entire system, shed antlers later, and g o s s reproductive
would be foolish in light of the competing performance decreased ( f i e et al. 1979, ECle
variables that influence both predators and and White 1985, Teer et al. 1991).
prey. The relative merits of each model Eventually, the population declined to levels
continues to be a source of debate within the comparable to outside the exclosure.
scientific profession. Only through Compensatory mortality occurred with
informative discussion, exchange of ideas, higher mortality among fawns 6-12 months
developing data sets, and testing of of age, rather than the mortality occurring
hypotheses will the debate prove fruitful. among post-natal fawns (Knowlton and
Stoddart 1992). Essentially, the addition of
EFFECT OF PREDATOR CONTROL animals above canying capacity required
ON PREY POPULATIONS management action (e.g., increased harvest),
or as in this case, compensatory mechanisms
Predator control can enhance prey (i.e., malnutrition and parasitism) returned
populations if prey is at low densities the deer population to levels as before
relative to carrying capacity. In Alaska, predator removal, but in a less healthy
predator removal programs brought about condition (Knowlton and Stoddart 1992).
irruptions of moose, which allowed for
increased human harvest of moose If predator control is considered for
(Gasaway et al. 1983, 1992, Ballard et al. enhancing ungulate populations, several
1991). In British Columbia, following factors should be considered. In a study in
reduction of wolf numbers, recruitment was Quebec, wolf reduction was conducted on 2
enhanced 2-5 times for 4 ungulate species experimental areas while wolves were not
and all populations increased (Bergerud and reduced on 2 control areas (Potvin et al.
Elliott 1998). Similarly, deer populations in 1992). They found that deer populations
south Texas increased following an increased in all 4 areas as a consequence of
intensive coyote removal program (Beasom mild winters and recommended that wolf
1974). Predator control may obtain an reduction was not a viable management tool
increase in ungulate numbers, but the in this context. Thus it is important that
addition of animals can have consequences managers determine whether or not
not often anticipated. In a study conducted predation is a limiting factor. Also, is the
on the Welder Wildlife Refuge in south ungulate population below forage carrying
Texas, coyote predation on white-tailed deer capacity? Considerations of scale, timing,
fawns was substantial. To test if coyote and method of removal need to be addressed
predation was a factor limiting population (e.g., what size of area is needed, and can
growth, a 391-hectare exclosure was erected control be cost effective). As demonstrated
on the site and the coyote density reduced throughout this paper, managers also need to
inside. Deer densities in the exclosure consider ~vhatother factors may be limiting
tripled compared to densities outside the or influencing the ungulate population. In a
exclosure. and remainsd stable for 2-3 years recent rsvieiv of the relationships behveen
predators and mule and black-tailed (0,h. in the predatory equation because they
columbian~ts)deer, Ballard et al. (2001) contribute to the base determining the
concluded that (a) the relationship of a prey numerical abundance of predators as well as
population to forage carrying capacity was provide buffers for the prey and stability for
critical to the impacts of predation, (b) prey the predators. The balance between prey
populations do not respond to predator abundance and the resources upon which
removal if prey is at or near carrying they depend is another integral part of
capacity, and (c) when prey populations are understanding the interactions because as
limited by predators and are far below their resources (i.e. food, cover, etc.)
carrying capacity, predator removal could become scarce, their vulnerability to
enhance prey survival, but increased hunter predation typically increases.
harvest may be uncertain. Equally
important is whether or not clear alternate One of the glaring lapses is an absence
values or objectives of the prey population of long-term data sets with simultaneous
are served. measures of the abundances and
demographic parameters of predators and
WHITHER FROM HERE? prey within individual ecosystems. Such
data sets provide the insights needed to
Assessing the impact of predation generate the testable hypotheses that will
upon prey populations is one of the more help define predator-prey interactions.
daunting tasks facing the wildlife Ideally these data sets should include not
profession. If it were easy, we would only routine measures of the abundances of
already know the answers. Predation predatory and prey species of primary
involves events towards the top of interest, but also those of alternate predatory
ecological trophic schemes, with events at and prey species, as well as climatic
lower trophic levels having repercussions conditions (especially deviant events),
manifested in higher levels. Consequently, primary productivity, and cause-specific
attempting to unravel relationships at the top mortality among ine age classes of each
without accounting for those below becomes prey species. Unfortunately, the difficulty
illogical. However, we would be remiss of establishing and maintaining the interest
without identifying potential means of and resource stream required for such
improving our understanding. endeavors precludes many of us from such
pursuits. It is indeed sobering to think that
It now seems largely folly to attempt a for some long-term fluctuations (e.g., the
comprehensive understanding of the role cyclic pattern in jackrabbit abundance in the
and impact of predation on prey populations intermountain\vest), it may take 10-20 years
independent of other ecolo~ical of data before we can generate the
considerations. This may be less important appropriate questions, yet alone provide the
in the case of obligate predators subsisting answers to them.
on relatively few prey types but it becomes
increasingly important as the number and We must recognize that by managing
type of suitable prey and predators increases predation, \ye may be merely sustaining
the complexity of the system (Table I ) . prey populations in habitats that are
..\vailability and abundance of altsmate nizrsinal for other reasoris. In our quest for
suitable prey constitutes a sipificani tern1 numbers. \ye ma)- bs m3kinz choices
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managers also must address the question of Linnean Society of London, London.
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Table 1. Gradients of increasing complexity in predator-prey interactions.
Simpler predator-prey interactions to More complex predator-prey
- -- -
interactions
Single prey system Multiple prey systems
Stationary prey Mobile prey
Resident prey Transient (migratory) prey
Carnivore Omnivore
Obligate predator Facultative predator
Single predator system Multiple predator system
Figure 1. Types of functional responses of predators to increasing prey density: (Type I)
predator kills a constant proportion of the prey population regardless of prey density; (Type
11) predation rate decreases as predator satiation sets an upper limit; (Type 111)predator kill
rate lags at low prey density owing to low hunting efficiency or absence of search image
(Holling 1959).
Figure 2. The functional response of wolves to changing moose density. -filling rate
(number of moose killed per wolf per 100 days) was related to moose density (numberikm2)
with a hyperbolic, Michaelis-Menton equation (data from Table 2 in Messier 1994).
 0 0.5 1 1.5 2 2.5
Moose Density (no. per sq. km)

Figure 3. The numerical response of wolves to changing moose density. Wolf density
(nurnberi1,OOO km2presented on a log,, scale) was related to moose density ( n u m b e r h 2 )
with a hyperbolic, Michaelis-Menton equation (data from Table 2 in Messier 1994).