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Winter Feeding of Elk in Western North America


Abstract and Figures

Winter feeding of elk (Cervus elaphus) is a topic that has engendered a great deal of debate among wildlife biologists, policy makers, and the general public. The first institutional feeding of elk in North America occurred inJackson Hole, Wyoming, where several thousand elk are still fed during most winters at the National Elk Refuge. Winter feeding of elk is employed on an annual basis by state agencies in Idaho, Oregon, Utah, Washington, and Wyoming. During 1995-99, an average 31,000 elk were fed in those 5 states at a cost of $1.6 million. Most feeding programs originated due to conflicts between elk and agricultural uses of historic elk winter range. Wildlife man- agers generally resorted to feeding to reduce damage by elk to crops, and to provide economic benefits of main- taining more elk than diminished winter habitat could sustain. Several negative consequences result from feeding elk. These include (1) the monetary costs of feeding, which divert dollars from other resource programs; (2) excessive herbivory that alters plant community structure and consequently affects the value of habitats near elk feedgrounds to other wildlife species; (3) changes in elk behavior that are of both spatial and philosophical sig- nificance; (4) diseases, which are more readily transmitted among densely concentrated animals, threaten the wel- fare of elk and other species, and shape resource management; and (5) public perceptions that may lead to the devaluing of habitat. These consequences argue for a shift from a production-consumption model of elk man- agement toward management that embraces conservation of all species, maintenance of ecosystem functions, and sustainability of resources. I suggest proactive alternatives to winter feeding, which may avert conflict situations that precipitate public and political pressures to feed elk.
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Winter Feeding of Elk in Western North America
Author(s): Bruce L. Smith
The Journal of Wildlife Management,
Vol. 65, No. 2 (Apr., 2001), pp. 173-190
Published by: Allen Press
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Abstract: Winter feeding of elk (Cervus
is a topic that has engendered a great deal of debate among wildlife
biologists, policy makers, and the general public. The first institutional feeding of elk in North America occurred
inJackson Hole, Wyoming, where several thousand elk are still fed during most winters at the National Elk Refuge.
Winter feeding of elk is employed on an annual basis by state agencies in Idaho, Oregon, Utah, Washington, and
Wyoming. During 1995-99, an average 31,000 elk were fed in those 5 states at a cost of $1.6 million. Most feeding
programs originated due to conflicts between elk and agricultural uses of historic elk winter range. Wildlife man-
agers generally resorted to feeding to reduce damage by elk to crops, and to provide economic benefits of main-
taining more elk than diminished winter habitat could sustain. Several negative consequences result from feeding
elk. These include (1) the monetary costs of feeding, which divert dollars from other resource programs; (2)
excessive herbivory that alters plant community structure and consequently affects the value of habitats near elk
feedgrounds to other wildlife species; (3) changes in elk behavior that are of both spatial and philosophical sig-
nificance; (4) diseases, which are more readily transmitted among densely concentrated animals, threaten the wel-
fare of elk and other species, and shape resource management; and (5) public perceptions that may lead to the
devaluing of habitat. These consequences argue for a shift from a production-consumption model of elk man-
agement toward management that embraces conservation of all species, maintenance of ecosystem functions, and
sustainability of resources. I suggest proactive alternatives to winter feeding, which may avert conflict situations
that precipitate public and political pressures to feed elk.
words: agriculture, Cervus
economics, elk, disease, feeding, feedground, growth, habitat, management,
Winter feeding of elk occurs at a number of
locations in western North America. Elk feed-
grounds are small, but because of the migratory
nature of most elk herds, landscape-scale impacts
of feeding programs are possible. The objectives
of this paper are to explore the purposes, advan-
tages, and liabilities of feeding; to review the geo-
graphic distribution of elk feeding; and to detail
the effects of feeding at the individual, popula-
tion, and community levels. Because winter feed-
ing is the exception to the system by which most
elk herds are managed, a perspective of the ori-
gin of feeding to mitigate wildlife-human con-
flicts is instructive.
Bunnell (1995) reported that nearly 1 million
elk inhabited North America in 1995-a remark-
able recovery from the 50,000 reportedly remain-
ing at the turn of the 20th century (Seton 1927).
This restoration of elk is attributed to a legacy of
conservation including: regulation of hunting;
law enforcement; the reservation of wildlands in
national forests, national parks, national wildlife
refuges, and Bureau of Land Management lands;
1 E-mail:
research elucidating the ecological requirements
of elk; and habitat improvement programs of
state and federal agencies and conservation orga-
nizations. In 1 dramatic instance, early efforts to
recover elk to presettlement numbers were also
aided by the initiation of winter feeding.
Of the 50,000 elk left in North America at the
century's end, most were found in the remote
lands in and around Yellowstone National Park
(Seton 1927). At the southernmost extent of
this refugium lay Jackson Hole, 1 of the last
intermountain valleys in the United States set-
tled by people of European descent. Here the
conflicts between settlers, their livestock, and
elk reached epic proportions between 1890 and
1910, focusing a nation's attention on the plight
of wildlife as civilization advanced westward
(Anderson 1958, Wilbrecht and Robbins 1979).
Elsewhere, elk either had retreated from the
advance of western settlement, or simply were
eliminated by hunting for sport and market.
But most importantly, competition between elk
and livestock for common resources-winter
food and habitat-all but doomed the vast herds
of elk, bison (Bison bison), pronghorn (Antilo-
capra americanus), bighorns (Ovis canadensis),
OF ELK * Smith
and other big game. Their disappearance from
many places either went unnoticed or was con-
sidered a necessary passage in the taming and
civilizing of the American West (Hornaday 1931,
Trefethen 1961).
Yet as elk vanished from most of their range,
the combination of Wyoming's grand venue and
the growing prospect of local extinction evoked a
conservation imperative that overcame the ardor
of humans for the taming of wildlands and their
wild inhabitants. The story of the people and pol-
itics that rescued the Jackson Hole elk has been
eloquently recounted (Leek 1909, Preble 1911,
Graves and Nelson 1919, Betts 1978). In short,
the recommendations of E. A. Preble and D. C.
Nowlin (Preble 1911), scientist of the Bureau of
Biological Survey and Wyoming game warden,
respectively, were largely implemented. The 1911
"Preble Report" advocated the reservation of a
permanent winter range in Jackson Hole as
"essential for the proper protection of the elk.
Such a refuge should be of considerable size,
should be situated in a valley which the elk natu-
rally seek, and should comprise pasturelands, as
well as meadows which will produce hay for feed-
ing the animals after they have exhausted the
available forage."
During 1912, the United States government
purchased about 800 ha of private lands adjacent
to the town of Jackson as a nucleus of the Nation-
al Elk Refuge (NER). With a legislative appropri-
ation of $5,000 in 1910, the state of Wyoming had
already begun the practice of winter feeding.
During 1911, the state passed a memorial entreat-
ing the U.S. Congress to financially assist with the
feeding of the Jackson Hole elk. Congress com-
plied with an appropriation of $20,000 to investi-
gate the situation, to begin trapping and removal
of elk from Jackson Hole, and to purchase hay.
Thus began the first government-subsidized feed-
ing of wildlife.
The 1872 establishment and early protection of
Yellowstone National Park by the U.S. cavalry,
which predated the beginnings of the NER by 40
years, was an equally bold conservation move. In
Yellowstone, too, winter feeding was previously
conducted to enhance survival of elk and bison,
albeit on a less extensive scale than in Jackson
Hole (Houston 1982). In addition to winter feed-
ing, several thousand elk were trapped and
translocated by truck and train from the NER
and Yellowstone to reduce those populations.
These elk restocked depleted ranges throughout
North America (Robbins et al. 1982).
A telephone survey of wildlife agencies in the
western states and provinces provided informa-
tion on winter feeding of elk (Table 1). Since the
establishment of the NER, the state of Wyoming
has established 22 additional elk feedgrounds
west of the Continental Divide during the
1950s-80s. Elsewhere in North America, winter
feeding of elk is a rarity. Only in Idaho, Oregon,
Utah, and Washington are public herds of elk
annually fed (Table 1). About 23,000 elk are fed
during winter in Wyoming and 8,000 are fed in
the other 4 states (Table 1). Thus, only 3% of the
estimated I million elk in North America are fed
by management agencies during winter.
These data represent elk feeding programs that
occur on an annual or nearly annual basis
(depending largely on weather conditions). Not
included is feeding that occurs on an emergency
basis to avert high mortality of animals during
extreme weather conditions, or to mitigate a spe-
cific elk damage problem on a nonrecurring basis.
For example, Colorado has fed elk during severe
winters, and the Colorado Division of Wildlife
has developed a priori criteria that guide deci-
sion-making regarding when feeding should be
initiated. Alberta does not feed elk on an annu-
al basis, but uses "intercept feeding" to deter elk
in specific situations from feeding on or damag-
ing crops, primarily haystacks.
Moreover, in most western states and provinces,
private citizens feed free-ranging elk, either
deliberately or unintentionally. Deliberate feed-
ing may be motivated to enhance wildlife viewing
or to improve elk survival. Most winter feeding
by citizens is purely incidental to the feeding of
livestock, and the elk are either tolerated or
unwanted on livestock feedlines. It would be dif-
ficult to quantify the numbers of elk that occa-
sionally or even regularly consume hay intended
for livestock, forage grain crops, or browse fruit
trees. Such numbers may be substantial during
severe winters. The information I present per-
tains only to winter feeding programs for elk that
are sanctioned by state or federal agencies. In
most situations, agencies conduct all program
phases, from purchase or production of the hay
to its distribution to the elk. In some cases, pub-
lic funds are used to purchase hay, but private cit-
izens do the feeding.
Throughout this paper, the elk feeding pro-
grams in Wyoming, and the NER in particular,
are highlighted. Those programs account for
J. Wildl. Manage. 65(2):2001
1. Distribution of winter
of elk in North America.
The number of elk fed and number of days of feeding
are based
on averages for
1995-99. Figures
are based on 1994-98 for Utah and Wyoming.
State Location Land No.
of No.
days of Type
of Reasons for
(herd) jurisdictiona elk fed feeding hay fed feedingb
(<17 sites)
Sun Valley-S.
Boise River
(6 sites)
Idaho Swan
Valley (3 sites)
Idaho Total
Oregon Elkhorn
(10 sites)
Oregon Wenaha
Oregon White River
Oregon Jewel Meadowsc
Oregon Total
Utah Hardware Ranch
Washington Yakima
(13 sites)
Wyoming National
Wyoming Alkalai
Wyoming Fish Creek
Wyoming Patrol Cabin
Wyoming Camp
(Fall Creek)
Wyoming Dog Creek
Wyoming Horse
Wyoming South Park
Wyoming Dell
Wyoming McNeel
Wyoming Grey's
Wyoming Forest Park
Wyoming Jewett (Piney)
Wyoming Finnegan
Wyoming Franz
Wyoming North
Wyoming Bench Corral
Wyoming Black Butte
Wyoming Green River Lakes
(Green River)
Wyoming Soda Lake
Wyoming Fall
Wyoming Scab Creek
Wyoming Muddy
Wyoming Total
USFS, Pvt 740
State, Pvt, BLM 1,400
State 523
State 350
State 275
State 490
USF&WS 9,200
USFS 663
USFS 827
State 442
State 628
USFS, Pvt 712
State 1,137
State 1,150
USFS 232
Pvt 512
State 956
USFS 810
State 612
BLM 386
BLM 426
BLM 305
State 566
State 555
USFS 468
State, BLM
140 Baled
<80 Pelleted
Baled 2, 5
= Bureau of Land
= private,
USFS = U.S. Forest
Service, USF&WS
= U.S. Fish and Wildlife Service.
b The following
reasons for
feeding elk were identified
by state wildlife
managers: (1) loss of winter
to residential devel-
opment, (2) conflicts on adjacent agricultural
lands/to alleviate wildlife
damage, (3) to increase
elk numbers for
(4) to pro-
vide public
of elk and related economic
(5) elk migrations
to winter
are short-stopped
by livestock
ations, (6) to make
and removal
easier, and (7) public
concerns about winter
of elk.
c Roosevelt elk are fed at this location.
75% of the elk fed on the continent each winter, EFFECTS OF WINTER FEEDING ON
and they are the longest running, best docu- ELK POPULATIONS AND PHYSIOLOGY
mented, and best known of the elk feeding pro-
grams. Furthermore, the Wyoming situation is Studies of survival, reproduction, and physical
singular due to widespread brucellosis infection development of winter-fed elk have been con-
of elk in the western part of the state. ducted in northwest Wyoming and Utah. These
1, 7
2, 3
2, 4
3, 7
J. Wildl. Manage. 65(2):2001
studies were designed in part to reveal individual
and population-level responses to winter food
Physical Development
Body mass of winter-fed elk from Utah and
Wyoming is similar to that reported for other
Rocky Mountain elk (C. elaphus nelsoni;
Dean et
al. 1976, Thorne and Butler 1976, Taber et al.
1982). Elk and red deer typically lose body mass
in winter, which they regain during the growing
season (Mitchell et al. 1976, Nelson and Leege
1982). When supplementally fed, elk may lose,
maintain, or gain body mass in relation to the
ration provided. Adult females maintained in
pastures with limited natural forage at the NER
lost 10.8%, 6.2%, and 4.2% of body mass from late
January to early April on rations of 0.95, 1.03, and
1.36 kg of pelleted alfalfa/100 kg of body mass,
respectively (Oldemeyer et al. 1993). Changes in
body mass of female elk held in paddocks during
winter and fed 1.7 kg of hay/100 kg body mass
varied from -9.5% when fed baled hay to -4.2%
when fed pelleted hay (Thorne and Butler 1976).
At Utah's Hardware Ranch, a ration of 3.2 kg of
meadow hay (approx. 1.4 kg/100 kg body mass)
allowed adult females to maintain body mass dur-
ing winter (Kimball and Wolfe 1984). Bailey
(1999) found no difference in body condition
between supplementally fed elk and free-ranging
elk in theJackson herd during 2 winters, based on
body fat indices and allantoin:creatinine ratios.
Winter feeding may be expected to improve
antler growth of elk by retarding loss of body mass
during winter and necessary recovery during the
period of antler growth (Kozak
et al. 1994). Antler
mass achieved by elk at the NER was similar to that
reported in unfed elk herds (Flook 1970, Wolfe
1983, McCorquodale 1989). Antler mass of 2- to
15-year-old elk that died during 1989-94 on the
? 14,000
0 12,000
E 10,000
Z 8,000
. ' '
Jackson herd
- * o
-- "Elk Refuge
- - ?C o ? Elk Refuge
1982 1984 1986 1988 1990 1992 1994 1996 1998
1. The number
of elk counted
in the Jackson
herd and on
the National
Refuge during
winters 1982-98.
NER was unrelated to the amount of feed those
elk received, and the number of days they were fed
during the winter prior to growing their last set of
antlers. Instead, early growing season tempera-
tures during March and April of the spring pre-
ceding their deaths, and ambient temperatures
when male elk were in utero (which also correlat-
ed with cohort birth mass) were correlated with
antler size (Smith 1997). The former effect
emphasizes the sensitivity of antler growth to pre-
vailing foraging conditions during each year of
life (Taber 1959, Bubenik 1982). The latter pro-
vides additional evidence that environmental
conditions during the birth year influence
growth, reproductive success, and survival of
cervids (Albon et al. 1987, Mech et al. 1991).
With elk, body mass attained by fall can influ-
ence conception rates (Sadlier 1969, Mitchell et
al. 1976). However, there is no evidence that body
mass differs between fed and unfed elk herds.
Likewise, pregnancy rates of adult elk (>2-year-
old) (87% at NER, 85% at Hardware Ranch) and
yearling elk (17% at NER, 12% at Hardware
Ranch) fed during winter (Kimball and Wolfe
1979, Smith and Robbins 1994) are similar to
those reported in other elk populations (Hous-
ton 1982, Taber et al. 1982).
Although supplemental feeding has elevated
fecundity of white-tailed deer (Odocoileus
Ozoga 1987), the same has not been shown
for elk. In western Wyoming, midwinter calf:cow
ratios are lower than in adjacent states and east of
the Continental Divide in Wyoming, where elk are
not fed during winter. Since 1982, the numbers of
elk in the
Jackson herd and the number wintering
on the NER have increased (Fig. 1). This resulted
from low winter mortality and the difficulty of
achieving desired harvests
of elk, particularly
that spend summer in national parks (Boyce 1989,
Smith and Robbins 1994, Smith and Anderson
1998). However, midwinter calf:cow ratios have
declined, and are inversely correlated with elk
numbers on the NER (Fig. 2). Moreover, summer
recruitment of calves since 1990 in Grand Teton
National Park, where half of NER elk spend sum-
mer, is inversely correlated with elk counted from
helicopter in the park's central valley (Fig. 2).
Bailey (1999) found no difference in fetal
growth between supplementally fed elk and free-
ranging elk of the Jackson herd. At the rates and
duration that supplemental feeding occurs at the
NER, winter feeding did not produce larger birth
J. Wildl.
OF ELK * Smith 177
j34I a U
32 r= 0.78
30 P = 0.013 \
600 800 1,000 1,200
Number of elk
Fig. 2. Calf:100
cow ratios
at the National
Refuge during
1983-99 regressed on number of elk counted
on the
Elk Refuge the previous
winters (A), and calf:100
cow ratios
in Grand Teton National
on number
of elk counted
1991-99 (B).
mass than reported for elk that are not fed
(Smith et al. 1997). As in red deer, cohort birth
mass varied with annual spring temperatures and
consequent growth of new grass during their
birth year (Albon et al. 1987). This is not sur-
prising because most fetal growth occurs during
the last 2 months of gestation (Nelson and Leege
1982), after winter feeding has ceased at the NER.
There was evidence that winter feeding influ-
enced sex ratios at birth. More males were born
after winters when feeding began earlier and the
digestibility of the feed was higher (Smith et al.
1996). Survival
of male fetuses, which are energeti-
cally more costly to produce than females (Clutton-
Brock et al. 1982), may be favored by nutritional
supplementation early during gestation.
Winter mortality was reported to regulate red
deer on the Isle of Rhum, Scotland (Clutton-
Brock et al. 1985), and elk in northern Yellowstone
National Park (Houston 1982, Singer et al. 1997)
in a density-dependent fashion. Winter feeding
can reduce mortality of elk during severe winters
or on overstocked ranges. Winter mortality of elk
on western Wyoming feedgrounds averages <1.5%
annually (Boyce 1989;
J. Bohne, Wyoming Game
and Fish Department, personal communication).
Annual mortality of elk at Utah's Hardware Ranch
is <1% (Lou Cornicelli, Utah Division of Wildlife
Resources, personal communication). Wildlife
managers in the other states where elk are fed
report similarly low mortality on feedgrounds.
Radiocollared calves of the Jackson elk herd
that were supplementally fed on the NER had
higher winter survival (0.886; P< 0.04) than calves
that were not fed (0.714). However, survival of
calves on the NER declined as the number of days
the elk were fed during winter increased (Smith
and Anderson 1998). Elk were fed longer during
protracted winters. Thus, feeding cannot negate
environmental stressors completely, and maintain-
ing elk on feedgrounds for protracted periods may
increase the risk of mortality from disease (Smith
and Roffe 1994, Smith and Anderson 1998).
The desire to maintain larger numbers of elk
than available
winter habitat can sustain is generally
at the root of feeding programs. In most situations,
winter habitat has been fragmented, degraded, or
usurped by land uses such as ranching, farming, sub-
or road construction. Conflicts between
elk and human uses of the land have ensued.
Among the western states and provinces, feed-
ing of elk is far more prevalent now than prior to
1950. Winter feeding of elk was generally initiat-
ed in response to political pressure and remains
popular with the public. The following reasons
are cited by wildlife managers for feeding elk in
winter: (1) Feeding can maintain a larger num-
ber of elk than remaining habitat can support,
enhancing hunting opportunities. (2) Feeding
can make elk more available for public viewing,
and commercial benefits can result. (3) Feeding
may reduce winter mortality of elk and assuage
public concerns about animal welfare. (4) Feeding
alters winter distribution of elk, helping to keep
elk off private lands where damage to crops,
orchards, and fences occurs, and off roadways
where motorist safety may be of concern.
Economic and Recreational Opportunities
Feeding elk during winter engenders recre-
ational opportunities for harvest, viewing, and
photography of elk. Feeding also enhances eco-
nomic opportunities for guiding and outfitting,
and related businesses that benefit from con-
sumptive and nonconsumptive uses of wildlife.
In Wyoming, where state law does not permit
nonresident hunters to hunt big game without a
guide in any of the 15 national forest wilderness
areas within the state, the outfitting business ben-
efits from elk feedgrounds. The more elk that are
available, the more hunting licenses are available.
As more licenses become available, opportunity
to outfit elk hunts increases. During 1980, the
r=0.69 * ? \
P = 0.003 * \
6,000 8,000 10,000
Number of elk at t-1
J. Wildl. Manage. 65(2):2001
outfitting business in Teton County, Wyoming,
generated $2.4 million in direct sales from hunt-
ing of big game animals (Taylor et al. 1981). Not
all of this was related to elk hunting, of course.
However, community businesses realize indirect
revenue from both resident and nonresident elk
hunters purchasing outdoor equipment, food,
lodging, and entertainment. Applying a multipli-
er to account for indirect revenue, the outfitting
business generated $4.2 million in economic
activity in Teton County (Taylor et al. 1981).
Expenditures related to elk hunting occur during
fall when local economies may experience a lull
between summer and winter tourist seasons.
Private contractors offer horse-drawn sleigh
rides through feedgrounds at the NER, Donnel-
ley, Idaho, and Hardware Ranch, Utah. Prices
charged for rides in 1999 ranged from $3.50/
adult at Hardware Ranch to $12.00/adult at the
NER. These private business endeavors can pro-
vide opportunities for public education. State
personnel escort the public to view and help feed
elk atJewel Meadows in Oregon.
Mitigation of Human-Wildlife Conflicts
Controlling distributions of elk provides safety,
economic, and public relations benefits. Feeding
elk may improve transportation safety by short-
stopping elk that would cross roads and highways
during spring and fall migrations to reach more
distant winter ranges than locations where feed-
grounds are established.
states and provinces have legislated wildlife
depredation laws. Winter feeding has been pro-
moted to reduce compensation payments to land-
owners. Elk that might otherwise migrate through
or winter on private lands and consume or damage
crops are attracted
or hazed to feedgrounds that are
generally located on public lands. To further limit
elk access to private lands, 65 km of 2.5-m-tall
fences have been built in western Wyoming to fun-
nel elk to the Grey's River,
Soda Lake, and Muddy
Creek feedgrounds. Eighty km of fence prevent
elk from reaching wheat and hay crops at White
River, Oregon, where 7,000 deer and 350 elk are
fed. Near Yakima,
Washington, over 160 km of elk-
proof fence prevent elk from straying into fruit
orchards. Decision-makers in those states believe it
is cheaper to fence and intentionally feed elk than
it is to compensate landowners for damaged crops.
Additional benefits accrue from discouraging elk
from using private lands, particularly
in states with
compensation laws or in states that harbor elk car-
rying brucellosis. Efforts to limit access of elk to
private lands may conciliate landowners, legisla-
tors, and local communities. Should transmission
of brucellosis from elk to cattle occur, the potential
economic hardships have been amply detailed
(Thorne and Herriges 1992, Thorne et al. 1996,
Kreeger et al. 2001). During 1992, a Wyoming
rancher sued the federal government and the state
of Wyoming in separate court actions alleging that
wild elk or bison infected his beef cattle with bru-
cellosis (Keiter and Froelicher 1993, Carlman
1994). Although both federal and state judges
did not find for the plaintiff, hard feelings and crit-
icism of wildlife managers ensued. Costs of elk
management increased as state officials intensified
efforts to haze elk from private lands, and con-
ducted depredation hunts of elk during midwinter.
Additionally, during 1997, the state of Wyoming
asked the U.S. Department of Agriculture (USDA)
to review
the state's brucellosis program. Wyoming
requested the review to ensure the state's com-
petitiveness in interstate livestock commerce. Wyo-
ming accepted the USDA recommendations, in-
cluding the testing of cattle in 6 western counties
for brucellosis. All 44,000 head of cattle tested dur-
ing 1998 were found to be disease-free (Dr.
Logan, Wyoming state veterinarian, letter dated
15 March 1999, to the Wyoming Livestock Board).
Nonetheless, efforts to eliminate brucellosis in elk
and wild bison, and to prevent brucellosis trans-
mission to cattle, have escalated in Wyoming and
the adjacent states of Idaho and Montana
(Thorne and Herriges 1992, Kreeger et al. 2001).
The financial requirements to feed elk during
winter are both capital and recurring. The NER
program has evolved from feeding loose hay from
horse-drawn sleds to a fully mechanized opera-
tion that distributes processed feed. The changes
came as a result of the loss of traditional hay sup-
plies in northwest Wyoming, and escalating labor
costs (Robbins et al. 1982). A 1974 memorandum
of understanding between the Wyoming Game and
Fish Department and the U.S. Fish and Wildlife
Service calls for a maximum of 7,500 elk to be main-
tained on the NER each winter. The capital costs
required to annually feed this number of elk in-
clude 4 >1,000-ton-capacity feed storage sheds, 3
Caterpillar crawler tractors and wagons, 1 Osh
Gosh articulated feedtruck, a feed off-loading belt-
4 forklift vehicles with 1-ton-capacity
for loading feed into trailers,
and a good mechanic.
J. Wildl. Manage. 65(2):2001
FEEDING OF ELK * Smith 179
2. Cost of feeding 7,500 elk for an average 79 days/
(the past 25-year average)
on the National
in 1999 dollars.
Item Cost/day Cost/winter
Pelleted alfalfa
(28 tons/day) 4,060 320,740
Labor 180 14,220
Fuel 32 2,528
Total 4,272 337,488
The direct, recurring costs include contracting
and purchase of about 2,400 tons of pelleted alfal-
fa, salaries of feedtruck drivers, and fuel. Indirect
costs include contracting, administrative, mainte-
nance, and biological monitoring support, and
equipment depreciation. The annual recurring
cost of just distributing feed to 7,500 elk averages
$337,488 (Table 2). The U.S. Fish and Wildlife Ser-
vice and Wyoming Game and Fish Department split
the cost of the pelleted alfalfa
fed on the NER. The
U.S. Fish and Wildlife Service pays all other costs.
I contacted wildlife managers in those states
with ongoing elk feeding programs to ascertain
feeding costs. Feeding 1 elk for 1 winter ranges
from $35 to $112 (Table 3). These costs do not
include administration, contracting, or biological
monitoring of feeding programs.
Idaho-The Idaho Fish and Game Department
feeds elk from 3 herds in the western and central
part of the state. In addition, there are 3 loca-
tions in eastern Idaho, collectively referred to as
Swan Valley, where the state has purchased hay
that private citizens or state employees fed elk in
recent years. Since 1984, when annual feeding of
elk was initiated in Idaho, the total cost of feed-
ing has been $2,400,000. Idaho spent $133,000
during 1998, exclusive of permanent employee
salaries, to feed elk. A similar amount was spent
to feed just the Sun Valley-South Boise River elk
during winter 1996-97.
Oregon-Wildlife officials began feeding elk in
Oregon in 1953 at the Wenaha Wildlife Manage-
ment Area. Both Rocky Mountain and Roosevelt
(C. elaphus
elk are now fed at 4 locations
across the state. The feeding program currently
costs the Oregon Department of Fish and Wild-
life $158,500 annually.
Utah-Feeding began at Utah's Hardware
Ranch in 1947 to prevent elk from following tradi-
tional migration routes down the Cache Valley,
where depredations on orchards and agricultural
fields were occurring. During the 1980s, Kimball
and Wolfe (1984) reported an annual cost to feed
500 elk of $75,000, or $150/elk. Estimated annual
costs to feed 490 elk at Hardware Ranch during
1995-99 were $45,000. The higher, former costs
arise from the inclusion of permanent salaries
associated with administration, contracting, and
monitoring. Thus, when all support costs and per-
manent salaries are included, true costs of feeding
elk in Utah and elsewhere significantly exceed the
direct costs of provisioning hay to an elk herd.
Washington-This state pays an average $117,500
annually to feed 3,000 out of an estimated 14,000
elk in the Yakima herd. The program began dur-
ing the early 1950s in an effort to keep elk from
damaging fruit orchards. During the severe win-
ter of 1996-97, elk broke through a 160-km-long
"elk-proof' fence in localized areas where they
were not being fed. The elk moved onto private
lands and caused extensive agricultural damage.
Wyoming-This state estimates that its annual
costs to feed about 14,000 elk, plus pay for half of
the pelleted alfalfa fed on the NER, approach
$1,250,000 annually. Costs to the U.S. Fish and
Wildlife Service average another $175,000. Addi-
tional program costs resulting from the artificial
concentration and feeding of elk are management
efforts to control and mitigate brucellosis. These
programs cost Wyoming another $250,000 annually.
Thus, the feeding programs in Wyoming are
the largest in terms of numbers of elk fed and the
state and federal budgets required to sustain the
programs. The costs of elk management, includ-
ing administering feeding operations, biological
monitoring, and disease mitigation and research,
surpass the income the Wyoming Game and Fish
Department derives from the statewide sale of elk
licenses. The revenue generated from sale of elk
licenses in 1998 was $7,770,000, but costs of elk
management were $8,820,000. Costs of elk man-
agement west of Wyoming's Continental Divide,
where the state and federal feedgrounds are
located, totaled $2,758,000 in 1998 compared to
license revenues of $1,846,000 (H. Harju,
Wyoming Game and Fish Department, personal
Habitat Changes
As early as 1911, Preble (1911) noted the nega-
tive effects that elk were having on their habitat in
Jackson Hole. He observed intense competition
for food during winter, noting the elk "were driven
to browse on the willows and other shrubs already
nearly destroyed during previous winters. They
soon eat the smaller twigs and then are forced by
hunger to attack the bark and larger branches....
J. Wildl. Manage. 65(2):2001
Table 3. Cost/winter
of feeding
elk in several western states. Costs include
feed, labor
to distribute
feed, and fuel costs. Costs
and Washington
are averages for 1995-99. Costs in Utah and Wyoming
were averaged
for 1994-98.
State Location
(herd) No. of elk Cost/elk Total cost
Sun Valley-S.
Boise River
Swan Valley
White River
Jewel Meadows
Hardware Ranch
National Elk
Refuge (Jackson)
Fish Creek
Patrol Cabin
(Fall Creek)
Dog Creek
Horse Creek
(Fall Creek)
South Park
Dell (Hoback)
Forest Park
Piney (Piney)
Bench Corral
Black Butte
(Green River)
Green River Lakes
(Green River)
Soda Lake
Scab Creek
Muddy (Pinedale)
a Winter
1998-99 only.
b Winter
1996-97 only.
c Elk
are fed only
5 times/week,
and hay is harvested on site by sharecropper.
d Elk are fed only
once/week, and hay is harvested
on site by sharecropper.
Haystacks about ranches are, of course, eagerly
sought. When they find the stacks securely fenced,
large numbers die immediately around them."
Craighead (1952) and Murie (1944, 1951) like-
wise detailed the decline of palatable deciduous
woody vegetation. In 1944, Murie cautioned:
There has been too much reliance on feed-
ing of hay as a solution, rather than herd
reduction to range carrying capacity. Hay
feeding concentrates the animals and is the
surest way to destroy the browse of a range
where it is practiced... Willows on the refuge
are almost gone. Serviceberry is barely able to
keep alive. Aspen groves are on the way out,
and have been so heavily browsed that ex-
cept for falling leaves, they no longer fur-
nish much feed. Even many conifers here
have been trimmed up so far as elk can reach.
J. Wildl. Manage. 65(2):2001
FEEDING OF ELK * Smith 181
Heavy hedging of palatable woody plants occurs
adjacent to feedgrounds in Idaho, Oregon, Utah,
Washington, and Wyoming. On a landscape
scale, as one moves away
from areas where elk are
concentrated during winter, the vigor and health
of woody plants improve (Kay 1985, Romme et al.
1995). Supplementally fed white-tailed deer also
exhibited reduced browse utilization with dis-
tance from feeding sites (Doenier et al. 1997).
Because of their size and food requirements, elk
can be particularly damaging when they are con-
centrated near feedgrounds (Thorne and Butler
1976). They can prevent successful regeneration
of aspen by annually browsing suckers and strip-
ping bark from tree trunks (Krebill 1972, Hart
and Hart 1989). Declines in aspen regeneration
and growth, as elk populations have increased
during the 20th century, have been reported in
most Rocky Mountain national parks of the U.S.
and Canada (White et al. 1998).
Following the clearcutting of aspen stands on the
NER to stimulate vegetative reproduction, aspen
regeneration was measured inside and outside
small exclosures (Dieni et al. 2000). Post-treatment
density of aspen suckers was greater within exclo-
sures during 9 years of study. After 9 years, <5% of
stems outside exclosures were >2m tall, compared
to 68% of stems within exclosures. Dieni et al.
(2000) concluded that repeated annual browsing by
elk was suppressing aspen recruitment and growth.
Elk are primarily grazers and do quite well on a
diet of herbaceous vegetation. Excessive grazing
near feedgrounds was not mentioned by wildlife
managers I interviewed, probably because of the
limitations of deep snow on availability of herba-
ceous vegetation. In addition, the migratory
of elk provide grazed plants a period of recovery
during the growing season, and seasonal grazing
may stimulate above-ground production (Augus-
tine and McNaughton 1998, Frank 1998).
Heavy browsing by high concentrations of
ungulates that limits growth and health of woody
vegetation, however, may produce community-
level consequences. Deterioration of woody plant
communities may occur slowly and be impercep-
tibe to the casual observer. Only careful moni-
toring may reveal cascading consequences of
habitat deterioration to other biota. Western
aspen communities, for example, support a high-
ly diverse fauna, including 56 species of mammals
and 135 species of birds (Flack 1976, DeByle
1985, Stelfox 1995). For most species of birds,
abundance was correlated with canopy hetero-
geneity and successional stage of stands in Alberta.
Richness and abundance of bird species was
greatest in 120+ year-old aspen stands. Mammal
species richness in Alberta was also greater in old
(120+ years) stands of aspen, compared to young
or mature stands (Stelfox 1995). Carothers et al.
(1974) reported that where ungulates reduce the
vertical complexity of woody vegetation, bird
species diversity is likely to decline. Berger et al.
(2001) found inverse relationships between
moose densities and avian diversity and abun-
dance in willow riparian communities of the
Greater Yellowstone Ecosystem. There is growing
evidence that birds which use woody habitats as
breeding, feeding, roosting and brood-rearing
habitat are generally less abundant in habitats
used by high densities of ungulates (Casey and
Hein 1983, deCalesta 1994).
Changes in Elk Behavior
Among the motives for feeding elk is the desire
to modify their distribution. A common motiva-
tion expressed by wildlife managers for justifying
winter feeding was to prevent elk from migrating
to lower elevation, more snow-free winter ranges
(Table 1). Interception of fall migrations pre-
vented elk from occupying historic winter ranges
that were in private ownership. In most cases,
short-stopping migrations averted elk reaching
and causing damage on agricultural lands, where
hay, grains, or orchards were produced, or where
cattle were pastured and fed in winter. In some
cases, such as the NER and Sun Valley, Idaho, the
damage that elk were likely to cause would also
have included ornamental plantings in residen-
tial areas.
Some feedgrounds were established on winter
ranges, such as those for the Jackson elk herd in
Wyoming and Wenaha in Oregon. Many others
were established on transitional range between
summer and winter ranges (Elkhorn and White
River in Oregon, Swan Valley in Idaho, and most
of the feedgrounds of the Piney, Green River,
Pinedale herds in Wyoming). Still others were
located on elk summer range (Hardware Ranch,
Utah, and Forest Park feedground in Wyoming).
Elk readily habituate to feeding operations. Yet
if feeding is inconsistent, animals will move else-
where to satisfy their appetites, as occurred at
Yakima, Washington, when feed could not be dis-
tributed to elk during a winter storm. Thus, bait-
ing of elk is initiated at some Wyoming feed-
grounds as early as November, to keep elk from
moving onto private lands, well before feeding
winter maintenance rations is necessary.
J. Wildl. Manage. 65(2):2001
Where elk are prevented from migrating to
areas of more accessible forage, the hay they are
fed may constitute the bulk of their diets because
standing forage is buried beneath snow. Only
woody plants may remain readily available to elk
and subsequently suffer severe hedging. Where
feedgrounds are contiguous with winter range,
such as theJackson elk herd, Yakima,
and Wenaha and White River in Oregon, feeding
serves to supplement elk diets with high-quality
forage to control distributions and reduce winter
mortality (Robbins et al. 1982).
Wildlife managers in Idaho, Oregon, and
Wyoming have noted some movement of elk,
from 1 winter to the next, among adjacent feed-
grounds and feeding sites. Nevertheless, elk fed
during winter generally display high fidelity to
winter feedgrounds (Tanner 1965, Smith and
Robbins 1994). Feeding likely reinforces fidelity
to wintering areas (Smith 1994).
Feedground attendance and arrival at feed-
grounds varies considerably with winter severity
(Boyce 1989, Smith and Robbins 1994). Conse-
quently, initiation of feeding varies with weather,
particularly snow accumulations. At the NER,
initiation of feeding was correlated (R2
= 0.96) with
December snow depths and the number of elk on
the NER (Smith et al. 1997). Weather conditions
also influence the composition of elk attending
feedgrounds in theJackson elk herd. When feed-
ing began later in winter, a higher proportion of
calves remained off feedgrounds. Radiocollared
calves were more likely to winter off feedgrounds
than were older radioed elk (Smith 1994). Calves
wintering off feedgrounds had poorer winter sur-
vival than those on feedgrounds (Smith and Ander-
son 1998); but the negative effect on overall cohort
survival was mitigated by fewer calves wintering off
feedgrounds during severe winters (Smith 1994).
Other concerns about the effects of feeding on
elk behavior are more subjective or emotional in
nature. Clearly, feeding elk during winter is pop-
ular with much of the public. A 1994 survey of
Idaho citizens revealed that 93% of hunters and
79% of nonhunters supported the Idaho Depart-
ment of Fish and Game's spending money to feed
big game animals during winter (Duda and
Young 1994).
People viscerally relate to feeding wildlife. It is a
step early
humans took many times in the process of
domesticating cats, dogs, horses, cattle, goats, etc.
habituation of elk to human presence and
to following feedwagons, rather than rustling for
wheatgrasses, troubles some observers. As 1 wild-
life manager in Oregon observed, "Once you con-
trol the food of the critter,
you control the critter."
Diseases affect the species composition of many
ecosystems, and are likely to play an important role
in management of wildland ecosystems in the
future (Real 1996). Infectious and parasitic dis-
eases can be important regulating mechanisms of
animal populations at high densities (Anderson
and May 1979, May 1983). Previous workers have
suggested that overstocked ranges could lead to in-
creased disease in ungulate populations (Cowan
1950, Murie 1951), or they have described wildlife
epizootics associated with high-density populations
(Matschke et al. 1984). Increased animal density
results in greater demand on the finite resources
of the available habitat and closer proximity of the
potential hosts of disease. Consequences include
poorer host nutrition and increased socio-behav-
ioral stressors, perhaps leading to reduced
immunocompetence (Sinclair 1977, Kistner et al.
1982), and increased opportunity for disease
transmission through animal-to-animal contact
and availability
of pathogens in the environment.
Viral, bacterial, and parasitic diseases of impor-
tance among free-ranging populations of feed-
ground elk include coronavirus and rotavirus in
neonates (Smith and Anderson 1996), septicemic
pasteurellosis and brucellosis (Thorne 1982a, b;
Franson and Smith 1988; Smith and Roffe 1994),
and psoroptic mange (Murie 1951, Samuel et al.
1991). Additionally, bovine viral diarrhea and
bovine respiratory syncitial virus were identified
in NER elk during winters 1997, 1998, and 2000
from serum antibody titers (T. Roffe, U.S. Geo-
logical Survey, unpublished data).
Psoroptic mange, or scabies, predisposes 20-30
adult male elk to die each winter on the NER
(Samuel et al. 1991). Scabies in elk is of little con-
cern to humans or the livestock industry due to
the host specificity of Psoroptes
cervinus. However,
clinical scabies does affect the aesthetics of
afflicted animals, and reduces the survival of tro-
phy size bull elk and the quality of capes of har-
vested animals. Elk may possibly serve as a reser-
voir for infection of sympatric bighorn sheep
populations (Lange 1982). Experimental injec-
tions of scabby bull elk with Ivermectin have pro-
duced short-term reductions of mite infestations
(Muschenheim 1988). Too little is known about
the host-parasite ecological relationships to pro-
vide practical management alternatives for eradi-
cation of scabies in elk.
J. Wildl. Manage. 65(2):2001
OF ELK * Smith 183
Septicemic pasteurellosis is an acute disease of
wild and domestic ruminants caused by Pasteurella
multocida. The hemorrhagic septicemic form of pas-
teurellosis is an acutely
fatal disease and rare on this
continent. It has been reported from dairy cattle
(Carter 1982) and several species of free-ranging
wildlife. Pasteurellosis has occurred at Wyoming's
Camp Creek feedground (S. Smith, Wyoming
Game and Fish Department, personal communica-
tion). Periodic outbreaks of the disease have been
documented on the NER (Thome 1982b,
and Smith 1988) with the largest number of ani-
mals dying during winter 1992-93 (T. Roffe and
B. Smith, unpublished data). From DNA finger-
printing, Wilson et al. (1995) suggested that P
multocida recovered from NER was a pathogen,
rather than an opportunist bacteria. Although
pasteurellosis is rarely reported elsewhere in elk
(Franson and Smith 1988), feedground elk are
more intensively monitored than most elk herds.
The lack of reports lends little to a conclusion
regarding importance of host density in the epi-
demiology of septicemic pasteurellosis. The
known epidemiology suggests that a wide range
of factors is important. Rapid progression and
usually fatal outcome of pasteurellosis makes its
epidemiology quite different from brucellosis
and scabies (Smith and Roffe 1994).
Bovine brucellosis is an infectious disease of cat-
tle occurring in at least 120 countries around the
world. The hallmark clinical sign of the disease
is abortion. Elk may also experience synovitis
arthritis, which causes lameness in some infected
animals (Thorne 1982
a). Transmission is by direct
contact with Brucella abortus contaminated repro-
ductive products associated with abortion or birth.
A national brucellosis eradication program has
nearly eliminated brucellosis in cattle in the Unit-
ed States. Since the 1980s, eradication efforts have
focused on the potential for wild bison and elk to
transmit brucellosis to cattle herds (Cheville et al.
1998). Elk may have contracted brucellosis from
infected cattle shipped from Europe (Tunicliff
and Marsh 1935), or secondarily from American
bison (Bison bison) that were initially infected by
the cattle (Thorne 1982a). In North America,
significant levels of brucellosis in wild elk occur
only in Greater Yellowstone Ecosystem (Thorne
and Herriges 1992). Bison in Yellowstone Nation-
al Park and bison that were introduced to Jack-
son Hole and winter on the NER are also infect-
ed with brucellosis (Williams et al. 1993).
Seroprevalence among adult female elk in the
western Wyoming feedground complex has
averaged 37% since 1970 (Thorne and Herriges
1992). During herd reductions of the 1960s, 1.7%
of 6,027 elk on Yellowstone National Park's north-
ern range were brucellosis test reactors (Smith
and Robbins 1994). Elsewhere, 2 of 178 Wyoming
elk not associated with feedgrounds tested posi-
tive in 1990 (Thorne and Herriges 1992). Brucel-
losis is absent or at nonsignificant levels of preva-
lence in other states (Smith and Roffe 1994), with
the exception of eastern Idaho, where elk on 2
feedgrounds tested positive in 1998 and 1999.
Experiments conducted at the NER indicated
that abortion could potentially reduce the annu-
al calf crop by 7% (Oldemeyer et al. 1993). How-
ever, the primary concern is the potential for
transmission of brucellosis to domestic cattle
raised within the distribution of elk herds in the
Yellowstone Ecosystem. USDA regulations that
restrict sale and shipment of brucellosis-infected
cattle and domesticated bison create concerns
about financial hardship among agricultural
interests in Wyoming, Montana, and Idaho that
harbor infected wild bison and elk. Litigation
against state and federal wildlife management
agencies, and threatened livestock market sanc-
tions against those states, are among the recent
repercussions of wildlife brucellosis.
Although elk and bison have experimentally
transmitted brucellosis to cattle in confined condi-
tions, transmission of brucellosis from elk to cattle
under field conditions has not been documented
(Thorne and Herriges 1992). High animal den-
sities that occur on feedgrounds are necessary for
transmission and maintenance of a high preva-
lence of brucellosis in elk (Thorne et al. 1996).
Olaus Murie, who first discovered brucellosis in
Wyoming elk in 1930, could not have envisioned
the consequences that feeding elk would pro-
duce. The costs, controversies, lawsuits, and ill
feelings that brucellosis has caused are well doc-
umented (Keiter and Froelicher 1993, Carlman
1994, Brimmer 1999). For a disease that is rela-
tively benign in elk, brucellosis has whipped up a
firestorm that has spread to the highest levels of
state and federal government. The predictable
spin-offs of interagency committees, environmen-
tal assessments, environmental impact state-
ments, feedground management plans, and
innumerable research efforts create work and
consume dollars that are often redirected from
wildlife and habitat management programs.
Most recently, the state of Wyoming sued the fed-
eral government to assert authority over wildlife
on the NER in order to force vaccination of elk.
J. Wildl. Manage. 65(2):2001
The 1999 federal court ruling on the case found
for the federal government, reasserting the statu-
tory authority of the Secretary of the Interior
over wildlife on national wildlife refuge lands
(Brimmer 1999). That ruling is under appeal.
Brucellosis has elevated the feeding issue to a
new level of public awareness. More citizens ques-
tion the justification for feeding when the prac-
tice is responsible for the spread and mainte-
nance of disease in elk.
Biologically, brucellosis is a red flag. It warns us
that out of a million elk in North America, only
those associated with the winter feeding programs
in western Wyoming and adjacent eastern Idaho
maintain this disease at any significant prevalence.
It warns us that the conditions experienced by elk
concentrated on feedgrounds are ripe for the
transmission of other, more pathogenic diseases.
Brucellosis is difficult to maintain in a free-ranging
population of elk, due to the restricted route of
transmission of the bacterium in reproductive
products. Other diseases that could be spread
through mutual grooming, shared food, and
aerosol would spread more rapidly through an
immunologically naive host population. These
include diseases such as bovine tuberculosis,
which devastated elk game farms in western Cana-
da and the United States in the early 1990s (Roffe
and Smith 1992), and chronic wasting disease.
The latter is poorly understood, has varietal forms
that affect a variety of mammals (including cattle
and humans), is popping up in game farms in the
western United States and Canada, and is in wild
populations of elk and mule deer of southeastern
Wyoming and northern Colorado (Williams and
Young 1993). Should either disease become
established within the Greater Yellowstone Ecosys-
tem, the number of infected herds could rapidly
expand. Twenty-five herds totaling 120,000 elk
and 2 herds totaling 3,000 bison winter in the 7
million ha Greater Yellowstone Ecosystem (Toman
et al. 1997). Because distributions of adjacent
herds overlap, generally during summer and fall,
bovine tuberculosis or chronic wasting disease
could spread to many herds (Fig. 3).
In cooperation with federal land management
agencies and the Rocky Mountain Elk Foundation,
the Wyoming Game and Fish Department has been
the brucellosis-feedground issue by con-
ducting elk winter habitat improvement projects.
The Wyoming Game and Fish Department also
implemented a brucellosis vaccination program in
1985 at the Grey's
feedground. Although it is
debatable whether the Strain 19 vaccine developed
for cattle is effective in elk (Peterson 1991), the
program has expanded to 21 of the state's 22 elk
feedgrounds (Thorne and Herriges 1992).
Keiter and Froelicher (1993) reviewed the law-
suit brought against the federal government by
Parker Land and Cattle Company and the suit's
fallout. They suggested that the only fully effec-
tive means of eradicating brucellosis from the
Greater Yellowstone Ecosystem's elk and bison
populations would be depopulation, "an extreme
policy choice, with serious political, ecological,
and economic repercussions." They went on to
say that, "In Wyoming, at least, any effective
response to wildlife brucellosis will almost cer-
tainly require reduction-if not elimination-of
the elk feedgrounds, which will undoubtedly im-
pact elk population numbers and hunting oppor-
tunities." Given the polarization and politicization
of the brucellosis issue, they advocated a regional
brucellosis control policy based on the principle
of risk reduction, not disease eradication.
In 1994, the Greater Yellowstone Interagency
Brucellosis Committee (GYIBC)
was formed. It is
comprised of land management, wildlife manage-
ment, and agricultural agencies from the federal
Fig.\3 Distri butio Yellowstone
e m
fedrud ad ENational
Park efue
I i <
\ I y
7 J
Fconsists of public lands. F c m
- C
fuge ^\ [ Boundary
J'Pvt j I ^ ' ') F
= Feedgrounds
50 km
Fig. 3. Distribution
of winter
feeding of elk at 22 Wyoming
and the National Elk
Refuge in the Greater Yel-
Ecosystem. Shown also is the distribution
of private
lands (Pvt)
within the 7 million-ha
ecosystem, which largely
consists of public
J. Wildl. Manage. 65(2):2001
government and the states of Idaho, Montana,
and Wyoming. The GYIBC meets 3 times each
year to coordinate brucellosis research and pub-
lic information efforts. A position statement
developed and adopted by the GYIBC in 1994
recognizes the link between concentration of
ungulates at feedgrounds and disease problems.
The statement concludes, "...the GYIBC strongly
recommends that winter feeding of elk should be
discouraged, and no additional public or private
feedgrounds be established in the Greater Yel-
lowstone Area. Establishment of emergency or
permanent feedgrounds for other wild ungu-
lates, which may act as an attractive nuisance and
concentrate elk or bison, is likewise discouraged."
Winter feeding of elk can be viewed as a means
of conflict resolution, generally spawned by
intense public pressure. It is not based on scien-
tific principle and sustainable resource manage-
ment policy. Administrators may see winter feed-
ing as the least painful remedy for producing
immediate results to appease differing groups:
agricultural interests that desire rapid resolution
to crop damage, and pro-wildlife constituencies
that oppose reductions in elk populations despite
wildlife-human conflicts or dwindling habitat.
As such, winter feeding fits comfortably into the
context in which wildlife management developed
as an agricultural paradigm that employed sim-
plified concepts of ecosystems in an effort to pro-
duce abundant numbers of certain species for
harvest (Lancia et al. 1996).
The potential for spread and maintenance of
epizootic disease in artificially crowded elk popu-
lations, as evidenced by brucellosis in Greater Yel-
lowstone Ecosystem feedgrounds and bovine
tuberculosis and chronic wasting disease in
North American game farms, argues for a shift
from a production-consumption model of elk
management toward an ecological paradigm
advocated for the wildlife profession (Lancia et
al. 1996). Leopold's (1966) philosophy of con-
servation matured from the production of pre-
ferred species to an appreciation of the land as a
complex organism of interdependent and neces-
sary components. This shift moved the wildlife
profession beyond single-species management to
embrace conservation for all species, mainte-
nance of ecosystem functions, and sustainability
of resources (Holt and Talbot 1978).
Except for limited supplemental feeding of elk
in northern Utah during the 1940s and 1950s, the
only location in which termination of feeding
programs conducted by a state or provincial
agency has been attempted is Idaho's Swan Valley.
Although the Idaho Fish and Game Department
advocated phasing out that elk feeding program
previously, public pressures perpetuated the feed-
ing until 1998. With the discovery of brucellosis in
elk tested at Rainy and Conant Creek feed-
grounds, the governor of Idaho appointed a task
force to make recommendations on management
of elk. This task force, composed of state officials
and private citizens, recommended reduction of
elk numbers and elimination of the feeding pro-
grams. The threat of brucellosis transmission from
elk to area cattle herds was the driving force (Dave
Koehler, Idaho Fish and Game Department, per-
sonal communication). A herd management
plan has been developed, with public input, to
liberalize elk seasons and bring elk numbers in
balance with available winter range and to reduce
crop depredations. Improvements of elk winter
habitat on public lands are intended to perpetuate
huntable elk herds in Swan
Valley. Winter feeding
continues for the next several years in Swan Val-
ley, while elk are trapped at feedgrounds and
translocated to alternative winter ranges.
It may not seem so in the face of a tide of pub-
lic and political pressures that rise in support of
initiating feeding, but it is easier to not begin
feeding elk than it is to terminate a feedground
once it is established. Some feeding programs
arose from an emergency or temporary situation,
rather than with the intent of feeding elk on a
long-term basis. A severe winter, a summer
drought, or an unusual movement of elk to pri-
vate farm or ranchlands may have prompted a
state agency to reduce anticipated winter mortal-
ity or bait elk away from private lands. In other
cases, private feeding of elk became public feed-
ing of elk when pleas from landowner or legisla-
tor prompted the state agency to assume the
responsibility to feed hay-conditioned animals.
situations are susceptible to varying
interpretations. Once elk are fed during a tough
winter, public expectation may render most win-
ters emergencies. Guidelines to define what con-
stitutes an emergency, such as those developed in
Colorado that specify when feeding is necessary
to prevent high losses of female elk or deer, must
exist before emergencies are declared.
Alternatives to winter feeding of elk are largely
recognition that wildlife management is people
management. People, via activities and use of the
landscape, affect wildlife. We influence animal
J. Wildl. Manage. 65(2):2001
distributions and the capacity of land to sustain
populations when we usurp and fragment open
spaces. Subdivisions, fences, agricultural
roads, and commercial developments alter the
capacity of elk winter ranges to sustain populations
or create conflicts that humans find unacceptable.
The notion of maintaining population levels in the
face of the erosion of habitat is simply unrealistic,
at least from an ecological perspective. Manipu-
lating elk populations in ways
that border on semi-
domestication, such as feeding and fencing them,
may permit numbers to be artificially maintained
on an eroded habitat base, but at what costs?
A number of actions can be taken before feed-
ing programs are established to avoid artificial
maintenance of elk herds: (1) Control elk num-
bers to keep them within the capacity of habitats
to sustain them on a long-term basis. This is a
basic tenet of sound wildlife stewardship
(Leopold 1933). (2) Practice land-use planning
that truly integrates the needs of elk populations.
This includes avoiding encroachment of subdivi-
sions and other development onto winter range,
and adopting regulations that discourage people
and dogs from harassing elk on winter ranges.
(3) Avoid fragmentation of elk ranges that short-
stops migration routes between summer and win-
ter ranges. This requires conserving migration
corridors as well as winter habitats. (4) Use tech-
niques that maintain or improve the productivity
of elk winter ranges. This includes forage pro-
duction enhancement through prescribed burn-
ing, seeding of degraded rangelands, proper
stocking of livestock if such grazing occurs on elk
winter ranges, and control of noxious weeds that
may compete with forage species valued by elk.
Many elk winter ranges on public lands border
lands or are a composite of private
and pub-
lic ownership. This clearly complicates elk con-
servation and increases the risk of conflict between
public resource stewardship and private property
rights. Fee title purchase of critical winter ranges,
and land exchanges that trade public lands or their
mineral rights for private lands, have served to con-
solidate and preserve winter ranges for the public
good. Short of land acquisition, conservation
easements may serve to maintain private lands in
private ownership and accomplish public resource
objectives. In other cases, cooperative ventures
between public agencies and private landowners
can promote the value of private lands as elk win-
ter range by compensating landowners for
accommodating wildlife. These ventures may in-
clude compensating landowners for production
and reservation of standing elk feed, or by
landowners charging hunters access fees.
In a recent essay,
Dave Stalling (1998) wrote, "As
throngs of people settle where elk once wintered,
fostering a piecemeal whittling of habitat, wildlife
management options dwindle. Alternatives to
winter feeding decline, social pressures mount,
biopolitics expands, and elk grow less wild."
Indeed, the surest path to keeping the wild in
wildlife is to maintain wildlands. This, unfortu-
nately, is easier said than done because Americans
are in the midst of a pilgrimage from the metrop-
olis to the mountains. One justification that has
been put forward for feeding elk during winter is
that summer ranges are stocked below carrying
capacity. Lovass (1970) points out that feeding
elk is an effort to have more elk than the range
will support. He adds that feeding can at best
only compound the existing imbalance between
elk and range. Boyce (1989) contends that winter
feeding of the Jackson elk herd is justified. He
reasons that, "only winter range is out of balance,
and this is due to human encroachment." True
enough. But following this dictum, there remain
fewer and fewer places in western North America
where winter feeding of elk is then not justified.
Elk habitat is wildlife habitat.
As noted above, too
many elk can degrade habitats used by many spe-
cies of wildlife. Likewise, loss of elk winter range
to occupation and use by humans displaces not
only elk, but wildlife in general. Although the elk
may be fed on an adjacent site, other species may
experience a net loss in the land's capacity to sus-
tain them. The more that wildlife officials feed
the animals, the more the public may accept feed-
ing as mitigation for development. This erosion
of habitat, the currency of wildlife, confronts and
frustrates town and county planners throughout
the West. Good intentions to maintain open space
and the integrity of winter ranges are overwhelmed
by the rising value of real estate advertised "with
wildlife right in your backyard." Sustainability of
elk, warblers, and willows depends on public and
private stewardship of wildlife real estate. Leopold
(1966) articulated this stewardship responsibility
several decades ago: "We abuse land because we
regard it as a commodity to us. When we see land
as a community to which we belong, we may
begin to use it with love and respect."
Creative opportunities for habitat conservation
still abound. They meld the goodwill of landown-
ers and private citizens and the use of public and
J. Wildl. Manage. 65(2):2001
FEEDING OF ELK * Smith 187
private funds. Prime examples are recent efforts
to secure winter range for Yellowstone's northern
elk herd in Montana (McMillion 1999). Howev-
er, as time passes, opportunities to protect elk
winter ranges for the long-term sustainability of
herds likewise pass. Ultimately, 2 questions
should be answered when considering whether
winter feeding, or an alternative solution, should
be pursued to maintain elk in a conflict situation.
What are the inherent economic, ecological, and
political costs? Is this solution sustainable?
Sinclair (1991) contended that wildlife man-
agement and research are not separate entities.
He cautioned that management lacking the
application of the scientific method leads to mis-
management through acceptance of untested
hypotheses as dogma. Through the inductive sci-
entific method of seeking knowledge (Romes-
burg 1981), managers may conclude and advo-
cate that the long-term feeding programs in
western Wyoming have maintained large num-
bers of elk on diminished habitat and that those
programs have reduced expected wildlife-human
conflicts, in contrast to not feeding. It is not sur-
prising then that elk feeding programs are gen-
erally well supported by sportsmen and agricul-
tural interests for these immediate benefits.
Similarly, managers may inductively reason that
the liabilities apparently associated with crowding
elk on feedgrounds, including budgetary costs,
habitat changes, behavioral changes in elk, and
disease, would not occur in the absence of feed-
ing. These positive and negative associations are
correlative. Although based on years of observa-
tions, they provide administrators, managers, and
the public scant and potentially unreliable infor-
mation about the comparative utility of alterna-
tive management approaches. Without rigorous-
ly testing alternative methods 'for managing
human-wildlife conflicts, we cannot reliably con-
clude that winter feeding is the best, or worst,
solution for elk or the ecosystems they inhabit.
Conducting management experiments to test
and learn from alternative approaches to manage-
ment problems in which there is a high degree of
uncertainty about outcomes, or a high degree of
risk associated with incorrect decisions, allows
administrators and managers to evaluate alterna-
tives and to improve decision-making. Conducted
as rigorous experiments using the hypothetico-
deductive process (Romesburg 1981, Murphy and
Noon 1991), researchers can increase knowledge
of wildlife systems and managers can evaluate
management alternatives and provide the public
better rationales for decisions on complex issues.
Given the weight of socioeconomics and politics,
could this adaptive management approach to deci-
sion-making be applied to elk-human conflict res-
olution? Perhaps. Where conflicts arise in 2 or
more similar situations, alternative management
approaches could be tested experimentally. One
alternative may endorse winter feeding, another
may use public-private partnerships to plan and
promote habitat improvements and conservation
to maintain populations. Careful formulation
and testing of hypotheses of individual, popula-
tion, and community-level effects, as well as socio-
economic consequences, would benefit future
decision-making with respect to whether elk
should or should not be fed during winter. How-
ever, given the liabilities of feeding and the diffi-
culty of terminating ongoing feeding programs, I
could justify initiating a new feeding program only
if such a policy decision was inevitable and exper-
imentation was a consensus function of its use.
Another possible application of adaptive re-
source management to winter feeding of elk exists
in the western Wyoming feedground complex.
These ongoing programs afford an opportunity
to compare management alternatives to remedy
the brucellosis disease issue (Peterson 1991). Elk
in all feedgrounds are infected with brucellosis,
and vaccination against brucellosis occurs at 21 of
22 state feedgrounds. Hypotheses about preva-
lence of brucellosis and other population process
variables could be formulated and tested to com-
pare elk using feedgrounds and elk for which 1
or more feedgrounds are phased out. For the for-
mer, feeding and vaccination of elk would con-
tinue. For the latter experimental groups, man-
agement may include any of the 4 strategies listed
previously, as well as reestablishing elk migrations
to historic winter ranges on-public lands (Allred
1950). Such experiments with free-ranging popula-
tions of animals are complicated and run inherent
risks of failure resulting from lack of administra-
tive commitment to the experiment. Nonethe-
less, applied as adaptive management, managers
are not committed to a single model and can con-
sider the merits of 2 or more models simultane-
ously. Importantly, adaptive management can
also lead to evaluation of values and implicit
assumptions that often underlie existing manage-
ment policies (Lancia et al. 1996).
Proactive testing of management alternatives
may avert crisis management. Ultimately, agencies
J. Wildl. Manage. 65(2):2001
that have elected winter feeding to resolve
elk-human conflicts may be forced to examine that
choice subsequent to changes in public values,
legal challenges, or disease in feedground elk. The
loss of Michigan's tuberculosis-free status for cattle,
following the state's tuberculosis epizootic in white-
tailed deer, compelled the state government to ban
private feeding of deer and launch substantial
reductions in the infected deer population. Such
events can preempt careful evaluation of manage-
ment alternatives, potentially at the expense of
agency credibility and precious public resources.
I am indebted to the wildlife managers and biol-
ogists that provided information and frank discus-
sions on wildlife management and winter feeding
of elk in their states and provinces. I also thank the
many past and present staff of the National Elk
Refuge who have contributed to the long-term
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J. Wildl. Manage. 65(2):2001
... Humans provide forage to ungulates both intentionally, by delivering it to feeding sites, and unintentionally, by leaving food leftovers at compost piles and rubbish dumps (Smith 2001;Sorensen et al. 2014) or not protecting agricultural crops (Oro et al. 2013;Milner et al. 2014). Winter supplementary feeding, which involves providing forage to feeding sites during winter, is a widely practiced management approach for ungulates in temperate ecosystems of Europe and North America (Milner et al. 2014). ...
... Nevertheless, winter supplementary feeding of ungulates is also implemented for various other management goals. For instance, diversionary feeding aims at modification of ungulate movement and spatial behaviour to decrease their impact on forest and agricultural crops (van Beest et al. 2010a;Smith et al. 2001;Náhlik et al. 2005;Katona et al. 2014;Krasińska and Krasiński 2013;Jaroszewicz et al. 2017), limit ungulate-vehicle collisions (Wood and Wolfe 1988;Andreassen et al. 2005) or prevent disease transmission between livestock and wildlife (Brook 2008;Sorensen et al. 2014). ...
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Supplementary feeding is a commonly used wildlife management practice, but while it may bring benefits and fulfil management goals, it can also result in unintended negative consequences. In the temperate zone, winter supplementary feeding may reduce movement of some ungulate species, leading to increased utilization of feeding sites by individuals and, consequently, animal aggregation. However, the intensity of feeding site utilization can vary depending on various extrinsic factors, e.g. habitat type, population density or weather conditions. Here, we predicted that the index of feeding sites utilization, expressed as the distance of European bison (Bison bonasus) to feeding sites, would be positively associated with the severity of winter conditions; thus, bison will be closer to feeding sites on colder days and in the presence of snow cover. We analysed winter (December to March) tracking data of 43 VHF- and GPS-collared European bison (24 males and 19 females) collected from 2005 to 2012 in Białowieża Primeval Forest (NE Poland), where bison are supplementally fed with different intensity throughout winter. Female bison were closer to feeding sites than males throughout winter, and regardless of sex, bison were the closest to feeding sites in mid-winter (January to February) and on colder days independently of the time of the season. Additionally, the distances of bison to feeding sites were significantly related to snow cover and depth; i.e. bison were closer to the feeding sites on days with present snow cover and deeper snow. Hence, the winter area occupied by bison differed with changing weather severity — being 4 and 28 times larger in the warmest periods compared to the coldest days with snow cover (for females and males, respectively). This may have direct and indirect ecological consequences for the ecosystem due to potential impact on nutrient cycling, seed dispersal, interspecific competition, vegetation growth, forest succession and carrion distribution. Given these ecological impacts of bison and weather-dependent utilization of supplementary fodder, we recognize the possible need in the future to revise and adapt winter supplementary feeding to annual and seasonal variation in winter severity to meet management goals while optimizing the costs.
... Manipulating the availability of forage through the provision of additional food is a widely used management practice whose objective is to maintain or increase hunters' harvest yields and mitigate human-wildlife con icts (Smith 2001;Milner et al. 2014). Although supplementary feeding can positively affect populations, this practice may also have a negative effect on ungulates' health (Sorensen et al. 2014;Felton et al. 2017). ...
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The supplementary feeding of wild ungulates is a common and pervasive practice throughout Europe, but the understanding on its unintended ecological effects is still limited. This management action has different degrees of intensity from artificial grasslands to supplementation in feeders, since the type and criteria of each supplementary feeding programme depend on managers’ objectives (e.g. maintaining harvesting yields, increasing animal body condition, mitigate agriculture or forest damage). The decision on how, when and where feed may determine the magnitude of the effects. Here, based on a long-term and well‐replicated dataset and using mixed structural equation model (mixed-SEM), we investigated the effects of supplementary feeding on red deer and wild boar abundance and aggregation and how directly and indirectly it affects the prevalence of tuberculosis-like lesions (TBL). We, therefore, hypothesized that supplying food avoiding the use of feeders would have less effect as regards increasing transmission rates, despite the fact that its efficiency as regards modulating population dynamics may be similar to the provision at feeders. Supplementary feeding was characterised in field surveys carried out on 60 hunting estates (south-central Spain) over four seasons (2002/2003, 2006/2007, 2012/2013 and 2018/2019). The abundance, aggregation and prevalence of TBL in red deer and wild boar were also determined. Collectively, our results showed that most of the variability in population aggregation could be directly explained by the population abundance and, to a lesser extent, by the degree of supplementation. Regarding red deer, there was a positive interaction between population abundance and aggregation in relation to the prevalence of TBL, suggesting that the positive effect of abundance on the prevalence of TBL is intensified by population aggregation. The model also showed a significant and positive correlation between the prevalence of TBL in both species. Finally, we reported a negative relationship between the degree of supplementary feeding and the prevalence of TBL in wild boar. Our findings shed light on the drivers of TBL prevalence in wild ungulate populations. Our results are relevant for the design of disease-control actions aimed at reducing the prevalence of tuberculosis and other shared diseases favoured by intensive wildlife management.
... The erection of fences to facilitate the movement of LR animals to handling facilities, or, using a means of transport to move animals off the rewilding area, can be useful, but serious thought should be given to ensure that they are effective and not a danger to other wildlife in the area (Hanophy 2014). In large areas it is unlikely that relatively short lengths of fencing will facilitate the driving of animals to handling facilities, unless they can be lured to a restricted area, for example, by providing food during periods of shortage (Smith 2001). Rewilding programmes must assess the adequacy of the habitat for the rewilded species, minimizing the need for artificial shelters against cold or hot conditions, at least after the adaptation phase. ...
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Rewilding is a restoration strategy that aims to return anthropogenic ecosystems to a “self-organized” state, by reinstating trophic complexity through disturbance (e.g. predation, herbivory), dispersal and connectivity. In depopulated areas of Europe, lite versions of rewilding, that maintain but minimize the management of rewilding species (e.g. predators, large herbivores) is gaining support. Livestock rewilding (LR) is a form of rewilding-lite, that uses livestock landraces as keystone species in the restoration of herbivory (the functional integrity of ecosystems) offering ecosystem services, such as ecotourism and the sale of livestock population surpluses, that can mitigate the economic and social effects of rural depopulation. Many challenges remain to implementing LR, including (i) more empirical evidence is required of the feasibility of LR across a variety of habitats and conditions, and (ii) understanding the hurdles that legislation poses for LR, the latter being the aim of this study. To accomplish this, we reviewed the EU legislation on environmental protection, animal health and welfare, identification and traceability, and ownership and civil responsibility, to assess how this might apply to LR. Although there is no specific EU legislation prohibiting LR, the review indicates that it is not clear what legislation applies to LR, as LR’s status lies between that of livestock and wild species. As such the existing legislation can be a serious impediment to the development of LR programmes. We highlight the needs for a legal definition, and status of LR species and their ownership. We propose ways to adapt this legislation to support the application of LR programmes in abandoned areas of EU, for example, by using legal exceptions intended for livestock under extensive animal farming systems.
... There is a range of justifications for providing supplementary feed to wildlife. These include the desire to reduce crop and forest damage through diversion ("diversionary feeding", Milner et al., 2014), enhance an animal's reproductive success and winter survival (Schwartz and Hundertmark, 1993), increase hunting opportunities and game viewing (Smith, 2001), or manipulate migration patterns (Gundersen et al., 2004). In Sweden, surveys have shown that the primary reason given by hunters and forest owners for supplementary feeding is to increase the survival of targeted species. ...
Populations of large herbivores, including members of the deer family Cervidae, are expanding across and within many regions of the northern hemisphere. Because their browsing on trees can result in economic losses to forestry and strongly affect ecosystems, it is becoming increasingly important to understand how best to mitigate resultant damage. Previous research has highlighted the importance of regulating deer density and the availability of alternative forage to reduce browsing damage levels in conifer production stands. However, often only one or two proxies of forage availability have been used instead of applying a broad foodscape approach and more knowledge is needed to understand which types of alternative forage best mitigate damage. We conducted field inventories of damage that occurred during the previous fall/winter in 112 production stands in southern Sweden, while also measuring forage availability and cervid faecal pellets in the surrounding landscape (16 ha). Local landowners provided data on supplementary feeding. We found that variation in cervid (Alces alces, Capreolus capreolus, Cervus elaphus and Dama dama) browsing damage to top shoots or stems of young Scots pine trees (Pinus sylvestris, hereon pine), was better explained by the availability of alternative natural forage (using several indices and species of trees and shrubs) than by supplementary feeding. The proportion of damaged pine trees was higher in stands with a lower density of pine stems; in landscapes with a lower density of key broadleaf tree species (genera Sorbus, Salix, Populus and Quercus); and in landscapes with more open land (agricultural fields and paddocks). Damage was also higher in stands where relatively large amounts of moose faeces was found, while not related to the amount of faeces from other cervid species. The amount of supplementary feed (silage or other types such as root vegetables) did not explain variation in pine damage, but the result was possibly affected by relatively few study areas supplying sufficient data on supplementary feeding. The results from our inventory illustrate the efficacy of using naturally growing forage to mitigate browsing damage to young pine trees in managed landscapes. Creation of such forage is also recommended over supplementary feeding because of co-benefits to forest biodiversity and ecosystem services.
... Similarly, agricultural conversion has limited the availability of high-quality winter resources for elk, which serve as reservoir hosts of Brucella abortus ( Figure 2C). Large elk populations are now supported by lower-quality supplemental feeding, which reduces migration and promotes high-density aggregations, thereby increasing the spread of Brucella among these animals and potentially spillover to livestock [87][88][89][90] . Climate change may further exacerbate loss of phenological diversity and interrelated shifts in animal movement; however, this has not been explicitly linked to zoonotic spillover 91 . ...
Human-mediated changes to natural ecosystems have consequences for both ecosystem and human health. Historically, efforts to preserve or restore 'biodiversity' can seem to be in opposition to human interests. However, the integration of biodiversity conservation and public health has gained significant traction in recent years, and new efforts to identify solutions that benefit both environmental and human health are ongoing. At the forefront of these efforts is an attempt to clarify ways in which biodiversity conservation can help reduce the risk of zoonotic spillover of pathogens from wild animals, sparking epidemics and pandemics in humans and livestock. However, our understanding of the mechanisms by which biodiversity change influences the spillover process is incomplete, limiting the application of integrated strategies aimed at achieving positive outcomes for both conservation and disease management. Here, we review the literature, considering a broad scope of biodiversity dimensions, to identify cases where zoonotic pathogen spillover is mechanistically linked to changes in biodiversity. By reframing the discussion around biodiversity and disease using mechanistic evidence - while encompassing multiple aspects of biodiversity including functional diversity, landscape diversity, phenological diversity, and interaction diversity - we work toward general principles that can guide future research and more effectively integrate the related goals of biodiversity conservation and spillover prevention. We conclude by summarizing how these principles could be used to integrate the goal of spillover prevention into ongoing biodiversity conservation initiatives.
... Similarly, agricultural conversion has limited the availability of high-quality winter resources for elk, which serve as reservoir hosts of Brucella abortus ( Figure 2C). Large elk populations are now supported by lower-quality supplemental feeding, which reduces migration and promotes high-density aggregations, thereby increasing the spread of Brucella among these animals and potentially spillover to livestock [87][88][89][90] . Climate change may further exacerbate loss of phenological diversity and interrelated shifts in animal movement; however, this has not been explicitly linked to zoonotic spillover 91 . ...
Human-mediated changes to natural ecosystems have consequences for both ecosystem and human health. Historically, efforts to preserve or restore ‘biodiversity’ can seem to be in opposition to human interests. However, the integration of biodiversity conservation and public health has gained significant traction in recent years, and new efforts to identify solutions that benefit both environmental and human health are ongoing. At the forefront of these efforts is an attempt to clarify ways in which biodiversity conservation can help reduce the risk of zoonotic spillover of pathogens from wild animals, sparking epidemics and pandemics in humans and livestock. However, our understanding of the mechanisms by which biodiversity change influences the spillover process is incomplete, limiting the application of integrated strategies aimed at achieving positive outcomes for both conservation and disease management. Here, we review the literature, considering a broad scope of biodiversity dimensions, to identify cases where zoonotic pathogen spillover is mechanistically linked to changes in biodiversity. By reframing the discussion around biodiversity and disease using mechanistic evidence—while encompassing multiple aspects of biodiversity including functional diversity, landscape diversity, phenological diversity, and interaction diversity—we work toward general principles that can guide future research and more effectively integrate the related goals of biodiversity conservation and spillover prevention. We conclude by summarizing how these principles could be used to integrate the goal of spillover prevention into ongoing biodiversity conservation initiatives.
... Despite this, there are already various ways in which we are currently aiding these animals. They include, among others, rescuing trapped animals and sheltering those in danger and suffering due to weather conditions (Bovenkerk et al. 2003); saving sick, injured and orphan animals (Delahay et al. 2009); rescuing animals in natural disasters (Anderson & Anderson 2006); vaccinating animals against severe diseases (Blancou et al. 1988); and providing water and food to thirsty and starving animals (Smith 2001). All these actions and programs make an important difference for the animals involved. ...
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Foraging on crops by wild ungulates may create human–wildlife conflicts through reducing crop production. Ungulates interact with and within complex socio‐ecological systems, making the reduction of crop damage a challenging task. Aside from ungulate densities, crop damage is influenced by different drivers affecting ungulate foraging behaviour: food availability and food quality in the landscape (i.e. the foodscape) as well as fear from hunting and scaring actions (i.e. the landscape of fear) may together affect the degree of damage via both direct and indirect effects. A better understanding of the individual effects of these potential drivers behind crop damage is needed, as is an appreciation of whether the effects are dependent on ungulate density. We investigated this by applying path analysis to test indirect and direct links between ungulate density, foodscape, landscape of fear and human management goals on crop damage of oats and grass, respectively. Our results suggest that crop type is the major driver behind crop damage, with more damage to oats than to leys, implying that human decisions (i.e. changing crop type) influence the level of crop damage. We found that management goals and actions influenced the foodscape and the landscape of fear, by affecting the amount of forage produced in the agricultural landscape and the amount of scaring actions. Additionally, we found that supplementary feeding influenced the local ungulate densities in the area. Our results highlight the importance of including human actions on multiple levels when assessing drivers behind damage by ungulates in managed landscapes. We suggest that more studies using path analysis on multiple scales are needed in order to tackle complex issues, such as crop damage and other human–wildlife conflicts.
Animals, like humans, suffer and die from natural causes. This is particularly true of animals living in the wild, given their high exposure to, and low capacity to cope with, harmful natural processes. Most wild animals likely have short lives, full of suffering, usually ending in terrible deaths. This book argues that on the assumption that we have reasons to assist others in need, we should intervene in nature to prevent or reduce the harms wild animals suffer, provided that it is feasible and that the expected result is positive overall. It is of the utmost importance that academics from different disciplines as well as animal advocates begin to confront this issue. The more people are concerned with wild animal suffering, the more probable it is that safe and effective solutions to the plight of wild animals will be implemented in the future.
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Abstract 1. Wildlife aggregation patterns can influence disease transmission. However, limited research evaluates the influence of anthropogenic and natural factors on aggregation. Many managers would like to reduce wildlife contact rates, driven by aggregation, to limit disease transmission. We develop a novel analytical framework to quantify how management activities such as supplemental feeding and hunting versus weather drive contact rates while accounting for correlated contacts. We apply the framework to the National Elk Refuge (NER), Wyoming, USA, where the probable arrival of chronic wasting disease (CWD) has magnified concerns. 2. We used a daily proximity index to measure contact rates among 68 global positioning system collared elk from 2016 to 2019. We modelled contact rates as a function of abiotic weather-related effects, anthropogenic effects and aggregation from the prior day. The winter of 2017–2018 had greater natural forage availability and little snow, which led to a rare non-feeding year on the NER and provided a unique opportunity to evaluate the effect of feeding on contact rates relative to other conditions. 3. Supplemental feeding was the strongest predictor of aggregation, and contact rates were 2.6 times larger while feeding occurred compared to the baseline rate (0.34 and 0.13, respectively). Snow-covered area was the second strongest predictor of contact rates highlighting the importance of abiotic factors to elk aggregation, but this effect had half the strength of feeding. These results are the first to show, even in animals that congregate naturally, how greatly supplemental feeding amplifies aggregation. Contact rates were also 23% lower during times when elk hunting was active (0.10) compared to the baseline. 4. Synthesis and applications. Supplemental feeding increased contacts between elk well above the natural effects of weather, even after accounting for correlated movement expected in wintering ungulates. Similarly, differences in hunting season timing with adjacent areas led to an increase in contacts, suggesting an additional management option for reducing aggregation. The analytical framework presented supports the evaluation of temporally varying management actions that influence aggregation broadly and can be easily implemented whether the interest in changing aggregation is related to reduction of disease transmission, human–wildlife conflict or inter-species competition.
Supplemental feeding of wintering white-tailed deer (Odocoileus virginianus) has increased in recent years, yet little data address its implications. During January-March 1993 and 1994, consumption of supplemental feed, provided ad libitum, was monitored daily at 2 recently established supplemental feeding (RSF) sites in northcentral Minnesota. We also monitored browse availability and use monthly at the 2 RSF sites and at 2 control (no supplemental feeding) sites by a sequential clipping method. Cumulative feed consumption and deer-visits were consistent between sites within years, but not between years; greater feed consumption and deer-visits occurred during the more severe Winter 1994. Browse pressure differed (P < 0.0001) by month during Winter 1993, and by month, twig density (sparse, moderate, dense), and site during 1994. A site x distance interaction (P < 0.0001) in 1993 was not significant in 1994 when browse pressure was higher. During 1994, mean browse pressure at control and RSF sites was different (P< 0.0001). A key finding in our study was that supplemental feeding effects differed annually. Furthermore, in both years, browse pressure means of both RSF sites differed, in the same manner, from mean browse pressure at the 2 control sites. The observed consistency of response among replicates, within treatments and years, suggests that supplemental feeding effects may apply generally to other local deer populations.
White-tailed deer (Odocoileus virginianus) populations have been maintained at high densities in Pennsylvania for several decades with unknown effects on songbirds and their habitats. I evaluated effects of white-tailed deer density on songbird species richness, abundance, and habitat. I simulated 4 deer densities (3.7, 7.9, 14.9, and 24.9 deer/km2) within individually fenced enclosures on 465-ha forest areas in northwestern Pennsylvania. Within all enclosures, 10% of the area was clear-cut and 30% was thinned. Enclosures were subjected to 10 years of deer browsing, 1980-90, at the 4 simulated densities. I conducted bird counts in 1991. Varying deer density had no effect (P > 0.1) on ground- or upper canopy-nesting songbirds or their habitat, but species richness of intermediate canopy-nesting songbirds declined 27% (P = 0.01) and abundance declined 37% (P = 0.002) between lowest and highest deer densities. I did not observe the eastern wood pewee (Contopus virens), indigo bunting (Passerina cyanea), least flycatcher (Empidonax minimus), yellow-billed cuckoo (Coccyzus americanus), or cerulean warbler (Dendroica cerulea) at densities >7.9 deer/km2, and the eastern phoebe (Sayornis phoebe), and American robin (Turdus migratorius) were not observed at 24.9 deer/km2. Threshold deer density for effect on habitat and songbirds within managed (100-yr rotation) forests was between 7.9 and 14.9 deer/km2.
A decade after Romesburg admonished wildlife biologists to establish and test hypotheses to gain more "reliable knowledge," we have added an incentive to bring rigor to our science. Wildlife biologists are finding themselves defending their science against often savage criticism. At least 2 factors are central to producing solid, defendable science: (1) the rigorous application of scientific methods and (2) the development of clear operational definitions for terminology. The hypothetico-deductive (H-D) process, in the form of statistical tests of hypotheses based on experimental data, is hailed as the superior means of acquiring strong inference and reliable knowledge. Results from experimental studies, however, are seldom available, and most management decisions are made on the basis of incomplete information. We argue that even in the absence of experimental information, the H-D process can and should be used. All management plans and conservation strategies have properties that can be stated as falsifiable hypotheses and can be subjected to testing with empirical information and with predictions from ecological theory and population simulation models. The development of explicit operational definitions for key concepts used in wildlife science-particularly terms that recur in legislation, standards, and guidelines-is a necessary accompaniment. Conservation management and planning schemes based on the H-D process and framed with unequivocal terminology will allow us to produce wildlife science that is credible, defendable, and reliable.
In a Minnesota ecosystem, mass of female white-tailed deer (Odocoileus virginianus) fawns and adults, and survival of adult females in the face of wolf (Canis lupus) predation, were directly related to maternal nutrition during gestation. Mass of single male fawns produced by 2-year-old females, and survival of yearlings to 2 years of age were related directly to the nutrition of their grandmothers.