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Winter Feeding of Elk in Western North America

May 2001
Journal of Wildlife Management 65:173-190

Authors:

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.

. Cost of feeding 7,500 elk for an average 79 days/ winter (the past 25-year average) on the National Elk Refuge, Wyoming, in 1999 dollars.

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Winter Feeding of Elk in Western North America

Author(s): Bruce L. Smith

Source:

The Journal of Wildlife Management,

Vol. 65, No. 2 (Apr., 2001), pp. 173-190

Published by: Allen Press

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Paper:

WINTER FEEDING

OF ELK

WESTERN

NORTH AMERICA

BRUCE

SMITH,1

National

Elk

Refuge, P.O.

Box 510, Jackson,

83001, USA

Abstract: 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.

JOURNAL

OF WILDLIFE MANAGEMENT

65(2):173-190

Key

words: agriculture, Cervus

elaphus,

economics, elk, disease, feeding, feedground, growth, habitat, management,

population.

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: bruce_smith@fws.gov

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),

173

174 WINTER

FEEDING

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).

DISTRIBUTION

OF FEEDING

PROGRAMS

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

WINTER FEEDING OF ELK * Smith 175

Table

1. Distribution of winter

feeding

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

Idaho

Donnlley

Stanley

(<17 sites)

Sun Valley-S.

Fork

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

Elk

Refuge

Wyoming Alkalai

(Jackson)

Wyoming Fish Creek

(Jackson)

Wyoming Patrol Cabin

(Jackson)

Wyoming Camp

Creek

(Fall Creek)

Wyoming Dog Creek

(Fall

Creek)

Wyoming Horse

Creek

(Fall

Creek)

Wyoming South Park

(Fall

Creek)

Wyoming Dell

(Hoback)

Wyoming McNeel

(Hoback)

Wyoming Grey's

River

(Afton)

Wyoming Forest Park

(Afton)

Wyoming Jewett (Piney)

Wyoming Finnegan

(Piney)

Wyoming Franz

(Piney)

Wyoming North

Piney

(Piney)

Wyoming Bench Corral

(Piney)

Wyoming Black Butte

(Green

River)

Wyoming Green River Lakes

(Green River)

Wyoming Soda Lake

(Green

River)

Wyoming Fall

Creek

(Pinedale)

Wyoming Scab Creek

(Pinedale)

Wyoming Muddy

(Pinedale)

Wyoming Total

Pvt

USFS, Pvt

USFS

100

240

925

USFS, Pvt 740

2,005

State, Pvt, BLM 1,400

State 523

State 350

State 275

2,548

State 490

3,000

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

State, BLM

BLM

USFS

785

776

566

677

23,391

140 Baled

<80 Pelleted

Pelleted

Baled 2, 5

145

80-100

101

113

120

138

135

129

133

153

160

154

136

148

104

126

164

145

Baled

Pelleted

Baled

a BLM

= Bureau of Land

Management,

Pvt

= 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

range

to residential devel-

opment, (2) conflicts on adjacent agricultural

lands/to alleviate wildlife

damage, (3) to increase

elk numbers for

hunting,

(4) to pro-

vide public

viewing

of elk and related economic

benefits,

(5) elk migrations

to winter

range

are short-stopped

by livestock

oper-

ations, (6) to make

trapping

and removal

easier, and (7) public

concerns about winter

mortality

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

2,4,6

2, 4

1,2,3,7

3, 7

3,7

J. Wildl. Manage. 65(2):2001

176 WINTER

FEEDING OF ELK * Smith

studies were designed in part to reveal individual

and population-level responses to winter food

supplementation.

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

18,000

16,000

? 14,000

0 12,000

E 10,000

Z 8,000

6.000

4.000n

. ' '

Jackson herd

- * o

-- "Elk Refuge

- - ?C o ? Elk Refuge

1982 1984 1986 1988 1990 1992 1994 1996 1998

Year

Fig.

1. The number

of elk counted

in the Jackson

herd and on

the National

Elk

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).

Reproduction

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

virgini-

anus-,

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

those

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.

Manage.

65(2):2001

WINTER FEEDING

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

Elk

Refuge during

winters

1983-99 regressed on number of elk counted

on the

National

Elk Refuge the previous

winters (A), and calf:100

cow ratios

in Grand Teton National

Park

regressed

on number

of elk counted

during

summers

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.

Survival

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).

RATIONALES

FOR FEEDING

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-

divisions,

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.

BENEFITS OF FEEDING ELK DURING

WINTER

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

178 WINTER

FEEDING OF ELK * Smith

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.

Many

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

drift

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.

James

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).

COSTS OF FEEDING ELK

Economic

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-

veyor,

4 forklift vehicles with 1-ton-capacity

buckets

for loading feed into trailers,

and a good mechanic.

J. Wildl. Manage. 65(2):2001

WINTER

FEEDING OF ELK * Smith 179

Table

2. Cost of feeding 7,500 elk for an average 79 days/

winter

(the past 25-year average)

on the National

Elk

Refuge,

Wyoming,

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

roosevelti)

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

communication).

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

180 WINTER FEEDING OF ELK * Smith

Table 3. Cost/winter

of feeding

elk in several western states. Costs include

feed, labor

to distribute

feed, and fuel costs. Costs

for

Idaho,

Oregon,

Utah,

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

Donnlley

Stanley

Sun Valley-S.

Fork

Boise River

Swan Valley

Total

(mean)

Elkhorn

Wenaha

White River

Jewel Meadows

Total

(mean)

Hardware Ranch

Yakima

National Elk

Refuge (Jackson)

Alkalai

(Jackson)

Fish Creek

(Jackson)

Patrol Cabin

(Jackson)

Camp

Creek

(Fall Creek)

Dog Creek

(Fall

Creek)

Horse Creek

(Fall Creek)

South Park

(Fall

Creek)

Dell (Hoback)

McNeel

(Hoback)

Grey's

River

(Afton)

Forest Park

(Afton)

Jewett

(Piney)

Finnegan

(Piney)

Franz

(Piney)

North

Piney (Piney)

Bench Corral

(Piney)

Black Butte

(Green River)

Green River Lakes

(Green River)

Soda Lake

(Green

River)

Fall

Creek

(Pinedale)

Scab Creek

(Pinedale)

Muddy (Pinedale)

Total

(mean)

100

240

1,193

740

2,273

1,400

523

350

275

2,548

490

3,000

9,200

663

827

442

628

712

1,137

1,150

232

512

956

810

612

386

426

305

566

555

468

785

776

566

677

23,391

112

(81)

(43)

(56)

7,500

10,389a

134,000b

67,733a

219,622

130,000

11,500C

4,000d

13,000

158,500

45,000

117,500

325,128

27,316

39,696

21,658

28,888

33,464

44,343

50,600

19,024

31,232

55,448

50,220

40,392

25,862

30,672

17,995

20,376

42,180

22,932

28,260

46,560

48,676

43,328

1,094,250

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.

Idaho

Oregon

Utah

Washington

Wyoming

J. Wildl. Manage. 65(2):2001

WINTER

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

nature

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,

and

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

182 WINTER

FEEDING OF ELK * Smith

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,

Washington,

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.

However,

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."

Disease

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

WINTER FEEDING

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,

Franson

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

and

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

184 WINTER

FEEDING OF ELK * Smith

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

addressing

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

River

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

YELLOWSTONE

ECOSYSTEM

Fig.\3 Distri butio Yellowstone

e m

fedrud ad ENational

Park efue

I i <

\ I y

MNational

Park

7 J

Fconsists of public lands. F c m

- C

Elk

fuge ^\ [ Boundary

J'Pvt j I ^ ' ') F

= Feedgrounds

50 km

Fig. 3. Distribution

of winter

feeding of elk at 22 Wyoming

feedgrounds

and the National Elk

Refuge in the Greater Yel-

lowstone

Ecosystem. Shown also is the distribution

of private

lands (Pvt)

within the 7 million-ha

ecosystem, which largely

consists of public

lands.

J. Wildl. Manage. 65(2):2001

WINTER FEEDING OF ELK * Smith 185

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."

ALTERNATIVES TO FEEDING

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.

"Emergency"

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

186 WINTER

FEEDING OF ELK * Smith

distributions and the capacity of land to sustain

populations when we usurp and fragment open

spaces. Subdivisions, fences, agricultural

plantings,

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

private

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.

PERCEPTION AND STEWARDSHIP

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

WINTER

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?

THE ROLE OF SCIENCE

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

188 WINTER FEEDING OF ELK * Smith

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.

ACKNOWLEDGMENTS

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

studies of elk population ecology cited in this

paper. D. E. Huff, T.J. Roffe, andJ. C. Rhyan pro-

vided helpful reviews of the manuscript.

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Linking winter severity to space use of European bison around feeding sites in Białowieża Primeval Forest (NE Poland)

Article

Full-text available

Jun 2023
EUR J WILDLIFE RES

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.

Disentangling the effects of supplementary feeding on population abundance and the aggregation of wild ungulates as drivers of the prevalence of tuberculosis in south-central Spain

Preprint

Full-text available

Mar 2023

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.

Legislative hurdles to using traditional domestic livestock in rewilding programmes in Europe

Article

Full-text available

Dec 2022

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.

Forage availability, supplementary feed and ungulate density: Associations with ungulate damage in pine production forests

Article

Jun 2022
FOREST ECOL MANAG

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.

Human-mediated impacts on biodiversity and the consequences for zoonotic disease spillover

Article

Oct 2021

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.

Human-mediated impacts on biodiversity and the consequences for zoonotic disease spillover

Preprint

Sep 2021

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.

Changing attitudes towards animals in the wild and speciesism

Article

Jul 2016

Oscar Horta

Direct and indirect effects of food, fear and management on crop damage by ungulates

Article

Full-text available

Aug 2023

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.

References

Chapter

Jan 2023

Catia Faria

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.

Human activities and weather drive contact rates of wintering elk

Article

Full-text available

Jan 2021
J APPL ECOL

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.

ARM! For the future: Adaptive resource management in the wildlife profession

Article

Full-text available

Jan 1996

Effects of winter supplemental feeding on browse consumption by white-tailed deer

Article

Jun 1997

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.

Effects of Heavy Browsing on a Bird Community in Deciduous Forest

Article

Jul 1983

The Ecology of Reproduction in Wild and Domestic Mammals

Article

Aug 1970

Maximum Fecundity in Supplementally-Fed Northern Michigan White-Tailed Deer

Article

Nov 1987

John J. Ozoga

The African Buffalo: A Study of Resource Limitation of Population

Article

Jul 1978

Effect of White-Tailed Deer on Songbirds within Managed Forests in Pennsylvania

Article

Oct 1994

David S. deCalesta

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.

Coping with Uncertainty in Wildlife Biology

Article

Oct 1991

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.

Effects of Maternal and Grandmaternal Nutrition on Deer Mass and Vulnerability to Wolf Predation

Article

Feb 1991

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.

Lives of games animals.

Book

Jan 1953

E. T. Seton

Winter Feeding of Elk in Western North America

Abstract and Figures

Recommendations