HomeMy WebLinkAboutAPA2285Chapter 2 3
HABITAT CHANGES AND
MANAGEMENT
M i c hae l L . W olfe
Associate Professor
Department of Wildlife Sciences
Utah State University
Logan, Utah
The purpose of this chapter is two-fold.
First, I will sketch the nature of land-use
ch anges in North America that have oc-
curred since 1935 and some of their implica-
tions for big game resources. The second half
of the chapter will present a brief overview
of the current state of the arts of habitat
management practices for big game ani-
mals.
L AND-USE C H ANGES:
1935-P R ESENT
Sweeping alterations of the North Ameri-
can landscape during the first four centuries
of European man's habitation of the
continent were described in Chapter 17. The
Twentieth Century also witnessed striking
changes in major human uses of the land,
349
Big Game of North America
virtually all of which affected big game
habitat. Some merely were continuations or
reversals of trends in plant-successional
trends resulting from earlier disturbances of
the pristine habitat and involved logging,
livestock grazing, fire suppression and agri-
cultural practices.
However, the Twentieth Century involved
a new dimension, namely the multiplicity of
environmental disturbances in the wake of
the rapid growth of human population and
its increasing mechanization. This genre of
land-use changes-including urban sprawl,
development of transportation systems,
mineral exploration and energy develop-
ment-differed in that the impacts of these
changes on the landscape generally are in-
delible. In many cases, the consequence of
Twentieth Century land use has been the
permanent or long-term loss of big game
habitat.
Thus, for purposes of discussion, I will dis-
tinguish between successional changes and
technological and sociological influences.
These two categories obviously are not
mutually exclusive, since improved tech-
nology enables us to increase both the extent
and intensity of a particular disturbance
and, hence, its impact on big game habitat.
Successional changes
1. Forestry practices. Modern forest science
in North America emerged in the present
century. It predicated a shift in consump-
tive emphasis from mere exploitation to
management. Timber harvest still re-
mains the most visible aspect of forest
management. However, numerous prac-
tices now associated with intensive
forestry, such as fire control, prescribed
burning, reforestation, timber stand
improvement and road construction, have
considerable effect on wildlife resources.
With respect to big game habitat,
perhaps the most ubiquitous of these
trends has been the progressive exclu-
sion of wildfire as a result of sophisti-
cated fire detection and suppression tech-
niques. The elimination of fire as a major
ecological disturbance has not been
limited to forested areas, but also has oc-
curred on vast acreages of western brush-
lands and grasslands as well. The statis-
tics in Table 40 illustrate this trend
clearly. The figures for the latter half of
the 1960s show an approximate eight-fold
decrease in the total acres burned and a
five-fold decline in average fire size in
contrast to figures from a period three
decades earlier.
Impacts of fire suppression on big game
habitat have varied with respect to both
the plant communities and animal popu-
lations involved. In many forest eco-
systems, the effect has been maturation
of plant communities toward climax con-
ditions, often with attendant declines in
forage productivity and nutrient quality.
This succession has resulted in reduced
quantity and quality of habitat for some
big game populations, including those of
deer and moose, which thrive best on
ranges dominated by early and/or mid-
successional vegetation. Longhurst et
al. (1976) considered the decreased
amount of acreage subject to wildfires
and prescribed burns in California as a
major factor contributing to recent deer
declines in that state.
Table 40. Average annual wildfire statistics for the United States".
Area burned Average size
of burn
Number of fires Hectares Acres
Period (thousands) (millions) (millions) Hectares
1936-1940b 211 12.6 31.2 59.9
1965-1969 120 1.5 3.8 12.7
aFrom annual wildfire statistics compiled by the U':lited States Forest Service (1969) for federal, state and private lands.
•Figures for this period do not include those of Alaska.
350
Acres
148.0
31.5
''
In certain plant communi ties, however,
the influence of fire exclusion on big
game habitat has been somewhat posi-
tive . This was the case for extensive·
acreages of shrub-grass communities,
such as sagebrush , pinyon-juniper and
the desert grasslands of the Southwest. In
these communities , the effect of con-
tinued fire suppression , in concert with
excessive livestock grazing, has favored
the shrub component at the expense of
competing grass es, thereby creating
improved habitat for mule deer.
Generalization about the net effects of
timber removal during the Twentieth
Century on big game habitats and popu-
lations is difficult. Farming, settlement
and timber exploitation of forest lands in
eastern North America partially replaced
the natural role of wildfire in terms of re-
juvenating pristine forest ecosystems. In
the Northeast, many vast, once-culti-
vated areas were abandoned by settlers
in s earch of more productive land. In
t ime , some areas reverted to s econd-
Habitat Changes and Management
growth forests of benefit to white-tailed
deer and, to some extent, moose popu-
lations. However, maturation of these
second-growth woodlands during the past
several decades has resulted in the de-
terioration of big game habitat. In the
eastern United States, timber growth
now exceeds timber cut.
' By the 1930s, most of the original
southern hardwood forest had been re-
moved as a result of agricultural develop-
ment and logging. Subsequent decades
have seen the emergence of this region as
the most important timber-producing
area of the United States. Contemporary
silviculture in much of the South is based
largely on even-aged management of
several rapidly growing pine species.
Prescribed burning is employed exten-
sively to reduce fuel accumulations and
control the encroachment of competing
understory hardwoods . Periodic burning
induces low sprout growth that is within
reach of browsing deer (Lay 1956, 1957,
Lewi s and Harshbager 1976). In some
Cut-ove r area s and s tump farm s, foll ow i ng settlem ent of eastern North America , temporarily set ba ck
fores t successio n in som e area s to the ben efit of big game. Photo by C. H . Park; courtesy of the National Ar-
chiv es .
351
Big Game ofNorthAmerica
areas, however, the excessive application
of this technique has led to degradation of
deer habitat by the virtual exclusion of
mast-and browse-producing hardwoods
in pine monocultures with "clean" under-
stories.
Although logging activities in forests
of the West date back to the late 1800s,
large scale timber harvest on public lands
commenced in the 1930s and did not
increase substantially until after World
War II. In comparison to successional
changes caused by early wildfires and
livestock grazing, logging per se probably
was not a major factor in the mule deer
population "booms" experienced by sev-
eral western states in the 1950s and early
1960s. However, in dense coastal forests
of the Northwest, i~creased levels of
palatable forage on recently logged areas
temporarily improved black-tailed deer
habitat.
Modern logging practices and public
attitudes toward timber harvesting have
changed. Some earlier logging operations
were linked to the railroads. A common
practice was to clearcut progressively all
timber adjacent to rail lines (Hooven
1973). The advent of heavy-duty logging
trucks and crawler-type tractors made
practicable the harvest of timber on
smaller units. In the interim, land
managers were forced to accept the fact
that very large clearcuts are of limited
value to most wildlife species, since such
practices merely substitute one monocul-
ture for another. Increased public aware-
ness of the unsightliness oflarge clearcvt
tracts also influenced timber interests to
harvest smaller units, with resultant
benefits for some big game species and
populations.
Other practices associated with inten-
sive silviculture for production of conifers
on a monocultural basis frequently are
inimical to big game populations. The
increasing application of reforestation
techniques to shorten the cutting cycle
may limit regrowth of desirable forage
species following logging, thus.reducing
352
the usefulness of the "disturbed" area for
big game (Hines 1973). In some forests on
the west coast, timber stand improve-
ment practices include removal of com-
peting hardwoods, especially oaks, to
favor conifers. This practice can be harm-
ful to deer, for which acorns are a nu-
tritious food source. Another interesting
consequence of intensive forestry was
noted in the Appalachians by Beeman et
al. (1977), namely the removal of large
decadent trees that provide den sites for
black bears.
Road construction, a by-product of both
timber harvest and fire control, has both
positive and negative implications for big
game. Forest road systems provide access
to forest areas by conventional as well as
off-road vehicles and, thus, increase the
vulnerability of big game populations to
human disturbance, particularly during
hunting seasons. Logging roads, how-
ever, are not always detrimental to big
game. Secondary and lower-level roads
within logging areas may facilitate move-
ments of big game to foraging areas
created by logging. Seasonal closures of
these roads following logging have
minimized human disturbances and
enhanced use of logged areas by big
game, particularly elk.
2. Livestock grazing. The dust bowl era of
the 1930s focused attention on abuses of
public lands in the West. The year 1934
marked passage of the Taylor Grazing
Act, which was intended to regulate graz-
ing on these lands. This law established
the predecessor of the Bureau of Land
Management, curtailed grazing on se-
verely overgrazed areas and brought
some control to grazing on other areas.
The forerunner of the Soil Conservation
Service was established in 1935 to control
soil erosion on public lands and provide
private landowners with financial and
technical assistance for the proper man-
agement of their lands.
Wagner (1977) analyzed records of the
United States Department of Agricul-
ture's Statistical Reporting Service to as-
en z
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certain historical changes in the number
of domestic livestock grazed on public
lands of the 11 westernmost states
(Figure 67). From 1935-1975, sheep
numbers declined significantly. However,
this decline was offset by a gradual but
continuous increase in cattle numbers.
Thus, Wagner concluded that the
potential forage demand of domestic
livestock on western rangelands was at
an all-time high in 1975. However,
increased use of supplemental feeding
and implementation of modern range-
management practices, such as fencing,
water development and measures to dis-
tribute grazing pressures, partially com-
pensated for the effects of increased num-
bers of domestic livestock on public lands.
Certain facets of this scenario have
produced changes in big game habitat.
The excessive grazing pressures of the
late Nineteenth and early Twentieth
30
/'NO. SHEEP
I \
Habitat Changes and Management
centuries occasioned the invasion or in-
crease of woody plants on grassland areas
of the West, thereby creating favorable
forage and cover conditions for deer.
Throughout much of the West during the
first half of this century, deer populations
increased to unprecedented levels.
In some areas of the Mountain and In-
termountain West, where grazing has
been eliminated or drastically curtailed
in recent years, there appears to be a re-
version of brushy foothill ranges to the
original bunchgrass vegetation (Smith
1949, Wagner 1969). While such changes
probably have improved the quality of
habitat for elk, they have worked to the
detriment of mule deer.
Other live.stock-oriented activities
such as range-improvement programs,
predator control and fencing also have af-
fected big game habitat. Vegetation type
conversions and water developments will
300
I \ ...... ~--\ I .........
1 \ I -,
20
10 I
I
I
I
I
I
1875
/ , --J
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I ' I I 'J
1900 1925 1950 1975
en z
0
200 ....J
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100 ~ ::>
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Figure 67. Chronological trends of sheep and cattle numbers (excluding lamb and calf crops), and esti-
mated total livestock forage need (Animal Unit Months) in the 11 westernmost states (after Wagner
1977).
353
Big Game of North America
be treated later in this chapter, and the
impact of predator control efforts is
covered in other chapters.
Fencing is an integral aspect of live-
stock production. Certain economic fac-
tors have resulted in a substantial
increase in fencing of both public and
private rangelands in the West. In the
past, herders tended sheep on both winter
and summer ranges, without need for
fences. In areas where sheep remain, the
cost of hiring capable herders has led to
fencing of some sheep ranges.
The general transition from sheep to
cattle ranching throughout much of the
West also has necessitated extensive
fencing. This trend promises to continue
as federal land management agencies im-
plement rest-rotation systems for grazing
lands. These systems involve systematic
rotation of grazing on different subunits
("pastures") of a given grazing allotment
at monthly, seasonal or yearly intervals .
Within a given rotation cycle, each pas-
ture is "rested" during at least one in-
terval to allow its vegetation to recover .
The use of such systems requires fencing
of the component pastures within each
allotment.
While most big game animals can clear
livestock fences easily, occasional mor-
talities occur when animals become
entangled in the wire. Pronghorn are af-
fected most severely by fences. They do
not leap fences readily, but can pass
through or under barbed wire fences used
to confine cattle. However, woven-wire
fences represent virtually impenetrable
barriers to pronghorn movements. Where
such fences confine pronghorn during
severe winters, many may die of starva-
tion.
In recent years, feral horses and burros
have become a significant ecological fac-
tor in western North America . The
forerunners of the modern horse evolved
The effects of livestock grazing on big game ranges are striking when contrasted to "rested " or ungrazed
land. The soils of heavily grazed areas can be compacted-and, thereby, contribute to rapid erosion and top-
soil loss . Rest-rotation systems have shown that multiple uses of rangeland can be compatible. Photo
courtesy of the U .S. Bureau of Land Management.
354
in North America and spread to Asia via
the Bering Strait land bridge, after which
other modern equid forms such as asses •
and zebras arose (Clabby 1976, Stirton
1959). Horses suffered the same mys-
terious demise as many other large pre-
historic land mammals, becoming extinct
on this continent some 12,000 years B.P.
Spanish exploration and settlement oc-
casioned the reintroduction of horses and
burros to the American West in the
Sixteenth Century. Subsequently, many
of the animals escaped from captivity and
proliferated in the wild. By the mid-
Eighteenth Century, virtually all Indian
tribes of the West had horses, and their
warriors were skilled horsemen. Esti-
mates of the number of free-roaming
horses in North America at that time
range from 2-5 million (McKnight 1959).
During the Nineteenth Century, most
horse populations occurred west of the
Rocky Mountains. Wild herds were aug-
mented by animals released or lost by
ranchers and the U.S. Army during the
late 1800s and into the present century.
By 1935, an estimated 150,000 feral
horses existed on public lands in the 11
western states (Zarn et al. 1977). Sub-
sequently, these numbers were reduced
severely by commercial exploitation and
removal to reduce competition with do-
mestic livestock.
Public concern about the demise of
feral horses prompted passage of protec-
tive federal legislation-most recently
the Wild Horse and Burro Act (Public
Law 92-195) in 1971. Under the protec-
tion afforded by this law and in the
absence of effective natural predator
populations, feral horse and burro popu-
lations have increased dramatically,
some by as much as 20 percent per year.
In 1975, there were more than 50,000
horses and 5,500 burros on public lands
in the West.
At present, ecologists know very little
of the impact of these animals on the
desert and mountain ecosystems they oc-
cupy (primarily shrub-dominated habi-
Habitat Changes and Management
tats). Potential competition for food
exists among feral equids and several big
game ungulate populations of mule deer,
elk, pronghorn and bighorn sheep. Of
particular concern is the possibility that
burros may exclude desert bighorns,
whose future is already precarious, from
vital water sources. The question of com-
petition and conflict among feral equids
and native big game animals has not yet
received adequate study. One thing is
clear, however: lacking some form of con-
trol, the continued increase of horses and
burros will intensify whatever competi-
tion does exist. Very probably, this will
result in the degradation of habitat of all
species involved.
Technological and sociological influences
An in-depth treatment of the many in-
fluences of our growing technological society
on big game resources is beyond the scope of
this chapter's brief overview. However, a
few major influences must be considered.
1. Urban sprawl. The continuing flux of an
increasing human population into urban
centers and the attendant urban sprawl
of large and small cities alike in recent
years have resulted in significant and
permanent losses of big game habitat.
This trend is particularly acute in the
mountainous areas of the West, where
suburban subdivisions often are located
in foothill areas that formerly provided
crucial wintering ranges. Excessive snow
depths prevent the animals from winter-
ing at higher elevations. Uncontrolled
dogs comprise another problem asso-
ciated with the encroachment of suburbia
on big game habitat. Where these ani-
mals are allowed to roam freely, they
may inflict losses on local deer popula-
tions.
2. Transportation systems and vehicles.
A serious consequence of technological
progress during the Twentieth Century
has been the proliferation of vehicular
355
Big Game of North America
transportation systems. Highways in
particular have had substantial direct
and indirect impacts on many big game
populations. While highways and
highway construction result in some
losses of big game habitat, more im-
portant are their effects on migration
routes, the separation and isolation of
otherwise contiguous habitats, and more
ready access to remote natural areas by
recreationists. The construction of the
interstate and other multilaned highway
systems in recent decades has aggravated
these conflicts.
Not to be overlooked is the loss of big
game in collisions with vehicles. Puglisi
et al. (1974) reported that deer-vehicle
collisions in Pennsylvania increased by
218 percent from 1960-1967. They noted
that this increase was due in part to the
construction of an interstate highway
across the state. Longhurst et al. (1976)
noted that at least 20,000 deer are
killed annually on California highways .
This represents approximately 60 per-
cent of the average number of deer
harvested annually on a statewide basis
for the period 1970-1975. In recent years,
some efforts have been made to reduce
deer-vehicle collisions by the c:;onstruc-
tion of high fences along segments of
rights-of-way that intersect known sum-
mer or winter range areas or traditional
migration routes. These fences force the
animals to use special underpass struc-
tures to cross the highways.
356
The phenomenal increase in the popu-
larity and number of off-road recrea-
tional vehicles (ORVs) during the past
15 years represents yet another mani-
festation of advanced technology, af-
fluence and increased leisure time. Prior
to the mid-1950s, production of fot~.r
wheel-drive vehicles, motorcycles, snow-
mobiles, and most recently, all-terrain
vehicles was insignificant. A report of the
United States Department of Interior
(1972b) estimated the total number of all
types of these vehicles in the United
States at more than 5 million. These
figures represented a trend, and at least
with regard to snowmobiles, the trend
undoubtedly applies to Canada as well.
At present, the impact of ORVs on big
game animals is not well-documented.
However, at least two major impacts are
obvious. One is the increased disturbance
and possible displacement of animals
from areas subject to heavy ORV traffic
(Dorrance et al. 1975). These effects are
most critical during seasons when young
are born and during winter. In the latter
case, forced movements of animals de-
plete energy reserves at a time when they
already are under environmental stress.
Such disturbance also can displace ani-
mals from areas of vital shelter and food
resources. In terms of habitat degrada-
tion, the long-term effects of ORVs
represent an even greater liability. Soil
Gullies and rills initiated by OR V traffic pre-
cludes restoration of the area to big game habitat
for many years, once the traffic is banned. Photo
courtesy of the United States Geological Survey.
.....
. ' Habitat Changes and Management
Sheep graze along a hillside on public land in California where ORV hill-climbing contests took place the
weekend before. Livestock grazing, in some places, can accelerate erosion and reduce vegetative growth.
When land, almost anywhere, is exposed to persistent livestock grazing plus recreational use, its viability as
big game habitat is seriously diminished or lost. Photo courtesy of the U.S. Bureau of Land Management .
or snow compaction, erosion, destruction
of vegetation, and change of species com-
position all are potential impacts of ORV
traffic on big game habitat. Such habitat
damages vary in terms of duration, but
all are of serious consequence.
3. Water developments. Large-scale water
impoundments, developed largely since
the 1930s, have inundated millions of
hectares of big game habitat. A compila-
tion by Martin and Hanson (1966) re-
vealed more than 1,500 large reservoirs
in the United States with a total im-
poundment acreage of almost 6 million
hectares (15 million acres). As with
highway construction, the detrimental
effects of these impoundments are not
limited to the areas actually flooded.
Large reservoirs often disrupt big game
migration patterns and may result in the
isolation of otherwise suitable habitat.
Indeed, where critical winter ranges of
migratory populations are inundated, the
total effective loss of habitat may involve
a much larger area.
4. Mineral exploration and energy develop-
ment. Looking to the future, mineral ex-
ploration and energy development hold
considerable potential for the destruction
of big game habitat. According to Platts
(1974), surface mining currently ac-
counts for approximately 80 percent of
the ore and solid fuels produced. By 1971,
in the United States alone, some 1.6
million hectares (4 million acres) ofland
had been disturbed by surface mining
and related activities. Wildlife habitat
accounted for approximately one-half of
the disturbed land. While in the past
mining was centered in the Appalachian
and midwestern states, the West will
bear the brunt of future mineral extrac-
tion activities. For example, 42 percent
of the United States' known phosphate
357
Big Game of North A me rica
reserves are located in the West. Also, a
study by the National Academy of
Sciences (Box 1973) indicated that 0.6
million hectares (1.5 million acres) of the
51 million hectares (126 million acres) of
coal underlying the western United
States could be surface mined using cur-
rent methods. This same study projected
a total surface disturbance of about 780
square kilometers (300 square miles) by
the year 2000. While the magnitude of
this disturbance may seem small in rela-
tion to the total area impacted by forest,
range and agricultural activities as well
as urban sprawl, it represents only a frac-
tion of the total disturbance for all
mineral and energy reserves in North
America. Oil shale and tar sands also un-
derlie vast areas in the western United
States and Canada.
Surface mining is one of the more
potentially devastating environmental
disturbances on big game habitat, since it
involves not only the removal of vegeta-
tion but disruption of the soil profile and
local topography as well. In many cases,
however, properly executed reclamation
of disturbed sites can create habitat that
actually is more attractive to big game
than that which existed prior to mining
activities. These effects are particularly
important where mineral reserves un-
derlie extensive tracts of presently
marginal habitat.
Site disturbance represents only one of
many factors involved in the impact of
mineral and energy development on big
game habitats and populations. Other
consequential factors include increased
harassment and adverse effects asso-
ciated with support facilities such as
roads, construction camps and pipelines.
For example, studies (Klein and Hem-
ming 1976) confirmed initial fears that
the construction of large diameter
pipelines for transfer of crude oil and
natural gas over arctic habitats can
impede movements of caribou and to a
lesser degree, other ungulates. Where
pipelines intersect traditional migration
358
corridors specific modifications are
necessary to minimize their impact.
HABITAT IMPROVEMENT
PRACTICES
As human populations and land uses
continue to impinge upon big game habitat,
we can no longer rely merely on the largess
of the land for big game production. Future
big game populations will require deliberate
and effective habitat management strate-
gies.
The following discussion is predicated on
the assumption of continued emphasis on
consumptive use of big game resources.
Should hunting no longer be a viable big
game population management option or
opportunity in coming decades, habitat
management concerns likely will change.
Wildlife habitat still will be important, but
not in terms of optimizing big game produc-
tion as is now the case.
A cautionary note at the outset of this dis-
cussion must be interjected. The limited
scope of this chapter precludes enumeration
of the specific effects of various habitat
manipulation practices in all the numerous
climatic and vegetative regimes of North
America. At the same time, the response to a
particular treatment is largely site specific,
thus rendering generalizations difficult at
best. The reader should bear in mind that
beneficial effects obtained by a given prac-
tice in one area may not be duplicated on
another site where plant composition and
growth conditions differ markedly.
The essence of habitat management for
any big game population is to provide op-
timum interspersion of those vegetation
types required by the animals for food and
cover. This usually entails manipulation of
existing vegetation either to maintain or
alter its successional stage. In terms of ef-
fects on vegetation, different manipulative
methods may be employed to achieve similar
results. For example, given a closed pinyon-
juniper stand with a sterile understory, a
manager may use either fire or mechanical
' .. Habitat Changes and Management
When juniper--a low quality forage-is browsed to this extent by deer on winter range, the habitat is not
adequate to maintain a healthy and productive herd. A Wildlife Management Institute photo; taken by
Seth Gordon.
means to rehabilitate the stand for use by
deer or elk. The choice will be determined by
economic and aesthetic constraints as well
as biological considerations.
Where the weight of such constraints does
not dictate an alternative course of action,
"natural" methods of vegetation manipula-
tion such as prescribed burning, controlled
grazing and silvicultural practices should
receive higher priority than artificial ma-
nipulations, including mechanical and
chemical methods. Unexpected side effects,
which many plant communities cannot
readily absorb, more frequently are incurred
with artificial techniques than with natural
methods or phenomena.
The wildlife literature contains the re-
sults of numerous studies that purportedly
document beneficial effects for big game
populations of various habitat manipulation
practices. Most studies report marked in-
creases in animal utilization of treated
areas, but few show conclusive population
responses such as increased birth rates
and/or survival and population growth. Such
responses might simply reflect redistribu-
tions of static populations with no increases
in numbers. A notable exception is the work
of Biswell et al. (1952) in which the investi-
gators documented substantial increases in
both the density and reproductive rate of a
black-tailed deer population in response to
opening up dense stands of chamise brush in
northern California. While differential at-
tractiveness of treated areas to big game
cannot be rejected summarily, the utility of
future habitat improvement measures must
be evaluated in terms of definitive popula-
tion responses, not just circumstantial evi-
dence.
Most habitat improvement practices are
aimed at increasing forage supplies. These
359
Big Game of North America
practices often are based on the sometimes
erroneous assumption that food resources in
a given area are a limiting factor to the big
game population(s) of that area, in terms
of either nutritional quality or available
quantity. Such an assumption can lead to
manipulations of vegetation that produce
foods that are neither needed nor used. In
the process, the actual limiting factor may
be ignored to the further detriment of the
population(s).
It is imperative that managers not over-
look the fact that residual, untreated areas
of vegetation usually provide animals with
essential cover as refuge from human ac-
tivity and natural predators as well as pro-
tection from adverse weather conditions. To
a certain extent, the nutritional status
mediates the dependency of an animal on
protective cover for thermoregulation. How-
ever, microclimatic attributes of some cover
types are virtually indispensable to the sur-
vival of big game animals during periods of
climatic extremes. This is the case with
winter "yarding" areas in northern portions
of white-tailed deer range. Numerous inves-
tigators, including Verme (1965b) and
Ozoga (1968), showed that the dense,
usually coniferous cover of preferred yard-
ing areas has less snow accumulation,
warmer ambient temperatures, and lower
wind velocities than do surrounding up-
lands. In addition, Moen (1968b) demon-
strated that a dense canopy of swamp con-
ifers markedly reduced radiation heat
losses from deer, particularly on clear and
cold nights when, without overhead cover,
emissions would have been excessive.
Cover may be equally important in pro-
viding animals with protection from heat
stress. Linsdale and Tomich (1953) noted
that California deer sought out chamise
brush and closed woods for protection from
summer heat. Similar behavior was re-
ported for peccaries by Bissonette (1976).
Edgerton and McConnell (1976) attributed
higher summer elk use of dense, unlogged
conifer stands to the more stable thermal en-
vironment found there than in adjacent
partial-cut and clearcut stands. Also, al-
360
though moose are not affected adversely by
extreme cold, they are not well-adapted to
high temperatures. In the southern limits of
moose range, the shade of forest stands
provides moose with a vital refuge from
extreme summer temperatures (Kelsall and
Telfer 1974).
The point of this discussion is that the size
and spatial distribution of openings will de-
termine their utility to big game animals
regardless of the method of treatment em-
ployed to create openings in forested stands.
Depending on the species, the animals
generally will venture only a limited dis-
tance into open areas to feed. Openings
whose dimensions exceed this distance will
be utilized only at their periphery.
Recommended sizes for forest openings
prescribed by several authors for various big
game species are summarized in Table 41.
Although the figures relate primarily to log-
ging practices, they also should serve as
guidelines for other methods of vegetation
conversion. Of the statistics given, those for
maximum width (or diameter) of a treat-
ment unit are most critical. Treatment units
exceeding the recommended maximum area
may be acceptable, provided widths are not
appreciably greater than prescribed maxi-
mums. Aside from the maximum areas and
widths specified, individual treatment units
should be well-dispersed within a larger
management block to provide a balanced
mosaic of food and cover tracts.
Controlled grazing
Since at least the turn of the century,
sportsmen, scientists and conservationists
have debated whether domestic livestock
grazing is detrimental to big game habitat.
The subject of grazing is inherently too com-
plex to permit pat generalizations or conclu-
sions. For any given situation, the impact of
livestock grazing on wildlife habitat is de-
termined by feeding behavior of the species
i:Qvolved, stocking rates, the plant com-
munity and the season in which the grazing
occurs.
Habitat Changes and Management
Table 41. Recommended maximum sizes of openings in forest or woodland cover for various big game populations.
Species and source
White-tailed deer
(McCaffery and Creed 1969)
Mule deer and elk
(Reynolds 1966a, Patton 1974)
Mule deer and elk
(Reynolds 1966b)
Mule deer
(Terrel1973)
Deer and elk
(Leopold and Barrett 1972)
Deer and elk
(Hooven 1973)
Moose
(Telfer 1974)
Moose
(Peek eta!. 1976)
Location andior"
vegetation type
Northern Wisconsin, mixed
hardwood and conifers
Arizona, Ponderosa pine
Arizona, spruce-fir
Utah, pinyon-juniper
California
Oregon, Douglas fir
Canada, boreal forest
Minnesota, spruce fir
Excessive grazing sometimes causes ir-
reparable habitat damage and often is det-
rimental-at least in the short term-to
some big game populations. Extreme graz-
ing pressures of the late Nineteenth and
early Twentieth centuries resulted in loss
of habitat for bighorn sheep, elk and
pronghorn populations in the West. While
successional changes caused by grazing ulti-
mately proved beneficial to some deer and
elk populations, these same changes appear
to have eliminated bighorns and pronghorns
permanently from much of the animals'
former ranges.
Given this somewhat pessimistic intro-
duction, the positive aspect of grazing
should be emphasized, namely its potential
as a tool for manipulation of wildlife
habitats. The practice of grazing, regulated
with respect to timing and intensity, to
maintain a specific plant community or
produce desired successional changes rep-
resents a relatively new and viable manage-
ment strategy.
Prescribed grazing involves deliberate ap-
plication of forage consumption by one
species of domestic herbivore on a plant com-
munity to modify competition among the
plants of that community, thereby enhanc-
ing production of forage species preferred
by wild herbivores. Successful use of this
Allowable maxima
Area Width
Hectares Acres Meters Feet
2 5 100 330
18 46 490 1,600
8 20 320 1,060
10-30 25-75 320-640 1,060-2,120
8 20 200 660
12-24 30-60
130 320 500 1,640
80 198
method requires that: (1) stocking rates are
such that the domestic grazers forage on
their preferred food, and (2) timing and du-
ration of grazing be applied at the appro-
priate stage of plant growth to effect desired
changes in the plant community. Failure to
observe these constraints spells the dif-
ference between desired optimum utilization
and unwanted direct competition.
In most cases, populations of two or more
herbivore species can utilize primary pro-
duction of a given plant community more ef-
fectively than can a population of a single
herbivore species. Conversely, total animal
biomass that a unit of habitat can support on
a sustained basis usually is greater with
multiple-species use than with single-
species use. An example is the relationship
of big game and livestock populations in the
Intermountain West. In recent studies,
Smith and Doell (1968) and Jensen et al.
(1972) investigated the compatibility of
spring grazing by cattle and sheep on deer-
elk winter ranges, where the primary
browse species was bitterbrush. These inves-
tigators found that spring livestock grazing
caused little competition with big game for
forage provided that grazing was restricted
to the early growing season before rapid
growth of shrubs. In fact, removal of her-
baceous vegetation around the bitterbrush
361
Big Game of North America
plants by grazing livestock increased mois-
ture available to bitterbrush. This signifi-
cantly increased browse production for
winter use by deer and elk.
Similar results in vegetation manipula-
tion can be obtained by the use of an appro-
priate combination of big game animals. In
fact, in natural grazing communities, the
feeding niches of wild herbivores show
considerable diversification that minimizes
direct competition for food and allows effec-
tive utilization of available forage in a given
habitat. An interesting example of such
interactions among native ungulates was
found at Elk Island National Park in
Canada, described by Holsworth (1960) and
more recently by Wagner (1969). In this
ecosystem, browsing by elk and moose was
largely responsible for maintenance of
grassy openings utilized by bison. In the
absence of browsing pressure by elk and
moose, openings would have been invaded
by shrubs and trees, ultimately resulting in
the exclusion of bison.
Use of the grazing animal to manipulate
habitat for big game represents an effective
and ecologically sound management tool. In
terms of cost effectiveness, this approach
usually is less expensive than use of me-
chanical methods that would produce com-
parable results because the tool itself
represents a marketable product.
Forest management
Virtually all silvicultural practices have
been shown to affect forest-dwelling big
game animals in one way or another.
These practices include timber harvest and
slash disposal, site preparation and re-
generation efforts, rotation lengths and
timber stand improvement measures. Of
these, timber-cutting programs have by fa'r
the greatest impact. Indeed, Shaw (1970)
suggested that 90 percent of habitat
manipulations required by forest wildlife
can be achieved by properly planned cutting
programs. Given this premise, the following
discussion is framed primarily in the. context
362
of enhancing big game habitat through
timber-harvest procedures.
Most forest-dwelling big game animals,
in either deciduous or coniferous forest
habitats, thrive best where a diversity of age
and composition classes of plants occur in
.relatively small stands interspersed with
small openings (Telfer 1974). There is even
some evidence that woodland caribou, in-
habitants of extensive stands of boreal
forest, also may benefit from a diversity of
cover types (Bergerud 1971c). Fire was the
major primeval agent that maintained this
diversity. With progressive exclusion of
wildfire from managed forest ecosystems,
timber harvest constitutes the most practi-
cable means of creating or restoring the
necessary variety of cover types.
A major ecological consequence of forest
maturation and closure of the forest canopy
is a decrease in diversity and production of
shade-intolerant shrubs and herbaceous
plants in the understory. This generally
results in a reduction of palatable forage for
big game. Conversely, the primary benefit of
opening up a dense forest stand is to allow
light to reach the forest floor, thereby stimu-
lating the production of understory forage
plants.
The wildlife literature contains numerous
references documenting increased diversity,
productivity and nutrient content of forage
plants following logging, as well as in-
creased big game utilization of cutover
tracts. In this respect, moderate-sized
clearcuts or patch cuts generally are more
beneficial than selective-cutting or thinning
operations (Murphy and Ehrenreich 1965).
For example, the great increase in Scandi-
navian moose populations during the
present century has been attributed largely
to the shift from selective-cutting systems to
clear cutting (Lykke and Cowan 1968).
There probably exists for each combination
of forest cover type and site potential some
threshold below which residual canopy cover
or basal area must be reduced to stimulate
ap. appreciable increase in forage produc-
tion. With respect to the transition zone
between coniferous and deciduous forests in
Canada's Maritime provinces, Telfer (1973)
stated that the residual basal area of a
logged stand must be reduced below 17.2 •
square meters per hectare (75 square feet
per acre) before increased browse production
results.
Increased light penetration represents '
only one cause for increased forage produc-
tion often observed following logging. Other
factors include: (1) increased availability of
soil moisture, and (2) the release of nu-
trients previously tied up in tree biomass. Of
course, some nutrients are removed perma-
nently from the site when it is logged, but
Horwitz (1974) estimated that two-thirds of
the nutrients in trees are left on the logging
site in the form of roots, branches and other
unharvested material. Hence, the method of
"slash," or logging debris, disposal becomes
an important consideration for the release of
nutrients for future forage production and
the utility of a cut for big game. In areas of
high precipitation, such as eastern North
America, organic decomposition of slash will
result in relatively rapid return of the nu-
trients to the soil. However, in the drier cli-
mate of the mountainous West, slash may
remain largely intact for many years. Under
these circumstances, slash disposal by
prescribed burning will insure more rapid
nutrient release.
These facts do not imply that all logging
operations are inherently beneficial to big
game animals. Pengelly (1972) identified
some detrimental aspects of large clear-
cuts, including increased snow accumula-
tions and wind velocities, barriers created
by logging debris, and losses of vegetative
diversity. Pengelly also pointed out that suc-
cess in rehabilitating big game ranges by
logging often varies along a moisture
gradient. Moderate-sized clearcuts may
improve habitat for deer and elk in dense,
coastal forests where forage supplies are
limiting. However, a cut of the same size in
the sparser and moisture-limited forest
stands of the eastern Rocky Mountains
likely will be less beneficial, since browse
regeneration often is poor on drier sites. The
moisture variable also will determine, to
Habitat Changes and Management
some degree, the relative longevity of those
benefits to big game that might result from
logging. In areas with lower rates of annual
precipitation, the seral stages that follow
the disturbance of timber harvest generally
persist longer.
Whether silvicultural practices are bene-
ficial or detrimental to big game is de-
termined by many factors, the most im-
portant of which are the size and pattern of
the treatment units and the site potential
for both plants and animals. Cutting
schemes and the ensuing practices of slash
disposal, site preparation, reforestation and
timber stand improvement should be
planned and executed with purposeful, not
incidental, benefits in mind.
Prescribed burning
Some biologists have long recognized the
role of fire to maintain or rejuvenate habitat
quality for certain big game populations.
Indeed, as we have seen, species like deer
and moose benefited fortuitously from early
wildfires and from some fires prescribed for
timber management. Only recently, how-
ever, has the planned use of fire gained some
measure of acceptance as a valuable tool to
enhance and improve big game habitat.
The objective of prescribed burning is
periodic application of controlled fire to
produce the ecological benefits of a natural
state, while minimizing the negative effects
of wildfire.
Fire may be used to alter plant species
composition and increase production of se-
lected species. The response of the vegeta-
tion to a given burn is determined by an
assortment of factors too numerous to
consider in detail here. Some of the more im-
portant variables include existing plant
community composition, season, weather,
intensity of the burn (heat) and fire fre-
quency.
Impressive and sometimes spectacular
increases of herbaceous and browse plant
species have been observed following fire.
Such responses can occur for a number of
363
Big Game of North America
reasons, including increased availability of
nutrients released in the ash and decreased
competition among new-growth plants for
available light and soil moisture. Fre-
quently, increased levels of protein and
other nutrients in plants on burned-over
areas accompany the quantitative increases
in forage production. The duration of ele-
vated nutrient levels depends on local site
and climatic conditions as well as the nature
of vegetative cover prior to burning, but it
seldom exceeds three to five years.
Where browse has grown out of reach of
big game animals, fire damage to the aerial
portions of the plants often induces prolific
sprouting from root stocks. As a result, stem
densities frequently increase dramatically
over preburn levels, thereby producing an
abundant browse supply that remains
available to big game animals for several
years.
A prime example of the use of fire to
improve big game habitat can be drawn
from recent studies in northern Idaho and
Montana by Leege and Hickey (1971) and
Gordon (1976), respectively. In these areas,
extensive wildfires of the early 1900s
created seral brush fields that were im-
portant winter ranges for elk, moose and
deer. The principal browse species in these
areas include redstem ceanothus, willow,
red osier dogwood, serviceberry, mountain
An experimental area in Florida before the last of
a series of prescribed burns.
364
maple, chokecherry and aspen. In the ab-
sence of recurring fire, browse production in
many of these areas decreased because of
invading conifers, and the remaining pal -
atable shrub species grew too tall to be
utilized effectively. During the past decade,
prescribed spring and fall burning has been
used effectively to curb conifer growth and
rejuvenate production of preferred accessi-
ble browse species.
When contemplating the use of prescribed
burning for big game habitat, the manager
should heed Komarek's (1966) advice that
wildlife needs may not be met by application
of burning techniques developed for other
purposes. The forester and range manager
seek clean burns and maximum coverage,
whereas burning appropriate for big game
usually is less intensive and thorough. Tim-
ing, frequency and size of burns for wildlife
purposes do not necessarily coincide with
other land-use interests, but vary according
to the species, habitat and region. The
challenge to the land manager is to optimize
beneficial effects of prescribed burning for
big game -in conjunction with other rec-
ognized land-use objectives .
Mechanical and chemical methods
Since World War II, numerous mechanical
and some chemical methods for vegetation
Same area as in previous photo, soon after the last
prescribed burn.
,.,
conversion have been developed . These in-
clude bulldozing, cabling, chaining, railing,
root plowing, and aerial or ground-based ap-
plication of herbicides. Such treatments
Habitat Changes and Management
have been used extensively on western
rangelands 1n projects variously termed as
"brush control," or "range rehabilitation."
The shrub types involved are varied and in-
Same area as in previous two photos , eight years after last prescribed burn. This sequence demonstrates the
powerful i nfluence fire has in regulating the composition and physical structure of vegetation . In the hands
of an experienced wildlife manager, fire can b~ used to develop or maintain the diversity of flora on nearly
any landscape. When , where , and at what frequency and intensity fire is employed can provide suitable
forage and shelter for one or more species of wildlife including big game animals. Photos by Ro y Komarek;
courtesy of the Tall Timbers R esearch Station .
365
Big Game of North America
elude mesquite, pinyon-juniper, sagebrush
and chaparral. As the result of overgrazing
and fire suppression, many of these brush
communities developed into "closed stands"
with little or no herbaceous understory.
The treatment regimen is fairly standard;
namely, removal of the brushy cover fol-
lowed by seeding to a mixture of grasses and
forbs (and sometimes browse). Early conver-
sion programs were conducted with increas-
ing forage production for livestock as the
primary objective. Wildlife and watershed
considerations were of secondary im-
portance, and what enhancement of big
game habitat did occur was largely acci-
dental. Uniform conversion oflarge tracts to
homogeneous grasslands often nullified the
potential benefits of such "improvement"
practices to big game populations of deer,
pronghorn, elk and bighorn sheep. For
example, between 1950 and 1964, 1,200
projects converted some 1.2 million hectares
(3 million acres) of pinyon-juniper woodland
in the United States. This translates to an
average treatment unit of 1,000 hectares
(2,500 acres). Admittedly, not all of the
treatment units were this size; the point is
that many far exceed the recommended
maximum sizes shown in Table 41.
The "Big Mac" is the ultimate weapon among
mechanical methods of vegetation type conver-
sions. Photo courtesy of the U.S. Forest Service.
366
In some areas, juniper eradication by chaining
can enable regeneration of nutritious understory
vegetation. Photo by Don Domenick; courtesy of
the Colorado Division of Wildlife .
Vale (1974) estimated that approximately
10-12 percent of the total area (40 million
hectares: 99 million acres) of sagebrush
vegetation in the western United States has
been subjected to some form of control.
Despite the relatively slight impact of these
treatments on the total extent of this vegeta-
tion type, projects involving winter ranges
represent a potential threat to big game
habitat. Specifically, sagebrush is a staple,
nutritious winter food for many mule deer
populations . Its large-scale removal can ap-
preciably decrease a habitat's winter carry-
ing capacity.
Vale (1974) stated: "If designed, however,
to achieve a heterogenous vegetation of
small grassy regions, local areas of dense
brush, and expanses of open shrubs with
abundant herbaceous growth, sagebrush
control should help both wildlife and do-
mestic livestock. Control projects already
completed have been planned to produce
not this type of vegetation, but pure
·homogenous grasslands. Rangeland envi-
ronments with little brush are beneficial
only for livestock, not wildlife."
As suggested, such programs need not be
detrimental to big game. In fact, where
properly designed, substantial enhancement •
of big game habitat can be achieved. The
specifics of a properly planned and executed
project will differ according to the vegeta-
tion type and primary animal population(s)
involved, but some generalizations can be
made.
Individual treatment units should be
small and well-interspersed throughout a
larger complex of residual cover. This means
abandonment of the massive area approach.
Terrel (1973) considered that the proper con-
cept for pinyon-juniper management on deer
winter ranges was to "punch" strategically
spaced holes in the forest stands rather than
leaving islands of woody vegetation in units
cleared by chaining. Treatment should not
be done on sites where terrain, soil type or
average annual precipitation is inadequate
to insure the desired conversion (Plummer
et al. 1968). Likewise, treatment should be
avoided on ridgetops that provide important
cover tracts. Where reseeding of disturbed
brushfields is part of the treatment, palat-
able browse species should comprise a
substantial component of the seed mixtures
used. Since establishment of browse-produc-
ing shrubs may take several years, livestock
grazing on treated areas should be deferred
initially. Some reduction in densities of big
game populations through liberal harvests
also may be necessary to insure establish-
ment of seeded species.
Lyon and Mueggler (1968) described the
results of efforts to increase browse produc-
tion by herbicide treatment of several shrub
species in northern Idaho. They found some
lag in the mortality of competing, undesir-
able shrub species. Desired browse plants
showed relatively quick recovery from
crown dieback and poor persistence of
sprouting, but the most desirable browse
species, redstem ceanothus, was killed by all
treatments. The investigators concluded
that herbicide spray projects for browse
improvement should be based on considera-
tion of the plant composition of the shrub
community and careful weighing of the posi-
Habitat Changes and Management
tive and nE;gative effects of spraying at dif-
ferent times of the year.
Other habitat improvement practices
1. Water developments. Water represents
the third vital element of the habitat tri-
logy for any wildlife population. Water
requirements differ seasonally, among
species and populations, and even among
sex and age classes within a given popu-
lation. For example, lactating does have
greater water demands than do bucks. In
arid areas where the distribution of sur-
face water sources is a limiting factor, the
carrying capacity of habitat for some big
game animals may be improved by ad-
ding water areas. This may entail modifi-
cation of existing springs or, more
frequently, construction of "guzzlers."
Basically, these devices consist of large
and impervious rain-collecting aprons
that drain water into permanent storage
tanks for later use. Guzzlers originally
were developed for desert game birds, but
may be of particular benefit to desert big-
horns, deer and pronghorns. Big game
animals will also use simple dugouts in-
stalled for livestock as a part of range-
improvement programs. To a large de-
gree, optimum distribution of such water
sources depends on the cruising radius of
the target animal(s).
2. Browse rejuvenation. Many hardwood
browse species sprout vigorously fol-
lowing moderate mechanical injury. This
phenomenon may be employed advanta-
geously to stimulate browse production,
especially where browse plants are
excessively tall, dense or decayed. In the
East, hand cutting has been used in some
hardwood stands to improve browse
production for white-tailed deer. Similar
results may be obtained through properly
conducted thinning or "cleaning" opera-
tions during timber stand improvement
(Della-Bianca and Johnson 1965). Me-
chanical treatment is most beneficial
when conducted immediately prior to or
367
Big Game of North America
during winter, because the downed ma-
terial yields a browse supply that
otherwise may be unavailable.
Numerous other methods of browse re-
juvenation have been employed, par-
ticularly in shrub-dominated commu-
nities in the western United States.
Ferguson and Basile (1966) found that
"topping" of old-age bitterbrush plants
resulted in a nine-fold increase in twig
growth the following year. The mag-
nitude of response declined substantially
in subsequent growing seasons but, even
after four years, production of the treated
plants was twice as great as that of
untreated plants. In California chapar-
ral, brushfields have been treated by
crushing, mowing, rolling and chopping
to encourage new growth in the form of
crown sprouts or seedlings (Dasmann et
al. 1967).
3. Supplemental feeding. Artificial feeding
of wild ungulates to carry them through
stress periods long has been championed
by sportsmen as a panacea, but generally
decried by biologists as impractical.
Specifics of winter feeding are covered in
the chapter on nutrition. The following
brief discussion focuses primarily on the
rationale involved.
368
Artificial feeding is rarely, if ever, jus-
tified as an alternative (1) to improve
natural food supplies through habitat
manipulation, or (2) to implement ade-
quate harvest regulations to maintain a
big game population within carrying ca-
pacity limits. Certain special circum-
stances, however, may dictate its use. In
some cases, the economic returns to be
gained by supporting overwinter popula-
tion levels of big game in excess of
natural carrying capacity may justif:x the
expense of supplemental feeding. Such
situations do exist on commercial hunt-
ing reserves or on intensively managed
areas. One example of the latter situation
is provided by the forest areas of middle
Europe where winter feeding of big game
has been practiced for centuries. Other
considerations may dictate judicious and
usually local use of winter feeding, such
as: (1) to keep deer and elk out of com-
mercial orchards, (2) to supplement tem-
porary habitat losses on winter range due
to highway construction, suburban
development, etc., and (3) to help bring
an ungulate population through an
unusually severe winter as an emergency
measure.
The success of an artificial feeding
program depends both on the foods used
and the species involved. In the past, al-
falfa hay was used widely but, in re-
cent years, pelleted foods for deer and
even pronghorn have become increas-
ingly popular. Being broad-spectrum
feeders, elk generally fare reasonably
well in winter-feeding operations. With
deer, however, unless feeding-espe-
cially of hay-is commenced early in the
winter before the animals experience nu-
tritional stress, their rumen micro-
organisms will not be able to cope
with abrupt change in diet. Under these
conditions, mortality may be as great or
greater than it would be without supple-
mental feeding. This problem may vir-
tually negate the use of short-term, spon-
taneous, artificial-feeding programs as
an emergency measure for deer in severe
winters. For white-tailed deer, a browse-
cutting program as described earlier may
represent a more effective emergency
measure.
Regardless of the food materials used
or the big game species involved, over-
utilization of and damage to vegetation
from concentration of animals on feeding
grounds represents a deleterious side ef-
fect of artificial feeding. There is also an
ethical consideration that should be men-
tioned, namely the obligation to wild ani-
mals to allow them the opportunity to
remain wild (Leopold 1933). Habitat
management provides that opportunity.
This document is copyrighted material.
Permission for online posting was granted to Alaska Resources Library and Information Services
(ARLIS) by the copyright holder.
Permission to post was received via e-mail by Celia Rozen, Collection Development Coordinator,
on September 15, 2013, from Steve Williams, President, Wildlife Management Institute.
This chapter is identified as APA no. 2285 in the Susitna Hydroelectric Project Document Index
(1988), compiled by the Alaska Power Authority.
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QL
ECOLOGY AtiD MAtiAGEMEtiT
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