Content uploaded by Vernon C. Bleich
Author content
All content in this area was uploaded by Vernon C. Bleich on Jan 27, 2016
Content may be subject to copyright.
Conservation and management of ungulates in
North America
PAUL R. KRAUSMAN*yAND VERNON C. BLEICHz
yBoone and Crockett Program in Wildlife Conservation, Wildlife Biology, University of Montana,
Missoula, MT 59812, USA; zDepartment of Biological Sciences, Idaho State University, Pocatello,
ID 83029, USA
The North American Model of Wildlife Conservation had its genesis, in part, with the conservation
of native ungulates. Thus, it is important to understand what these species are and how they have
been and are, currently, managed. Ungulates are hooved quadrupeds that include those that are
odd-toed (e.g. tapirs [Tapirus bairdii]) and even-toed (e.g. deer). Contemporary management of the
ungulates evolved through three stages: exploitation, concern for survival and conservation through
wise use. The success of conservation of ungulates has been attributed to the merger of biology, an
understanding of habitat and the role of human dimensions. This triad of wildlife management has
been effective. Most ungulate populations in North America have been restored; viable populations
exist. Challenges to the conservation of large mammals remain, however, including increased
human population growth, climate change and sources of funding.
Keywords: Conservation; Management; North America; Ungulates
North American ungulates
Ungulates are hooved quadrupeds belonging to two Orders: Perissodactyla, which are
odd-toed ungulates (e.g. horses [Equus spp.], rhinoceroses [Rhinoceros spp.], and tapirs
[Tapirus bairdii]), and Artiodactyla, which are even-toed ungulates (e.g. pigs, deer and
pronghorn (Antilocapra americana). Only one native ungulate of the Order Perissodactyla,
the tapir, occurs in North America (i.e. Mexico). Other native ungulates in North America
are in the Order Artiodactyla, which includes 10 recent families, 86 genera and 221 spe-
cies throughout the world. Although Artiodactyls are not native to the West Indies, New
Guinea, Australia, New Zealand, Antarctica or most oceanic islands [1], some ungulates
have been translocated to these areas. Groups of the living Artiodactyla are provided by
Nowak [1] and include pigs, peccaries (Pecari spp.), hippopotamuses (Hippopotamus
spp.), llamas (Lama spp.) and camels (Camelus spp.), mouse deer (Tragulus spp.), giraffes
(Giraffa spp.), musk deer (Moschus spp.), deer, pronghorn, antelopes, bison, cattle, goats
and sheep.
Artiodactyls native to North America include collared peccaries (Pecari tajacu),
white-lipped peccaries (P. pecari), elk (Cervus canadensis), mule or black-tailed deer
*Corresponding author. Email: paul.krausman@umontana.edu
International Journal of Environmental Studies, 2013
Vol. 70, No. 3, 372–382, http://dx.doi.org/10.1080/00207233.2013.804748
Ó2013 Taylor & Francis
Downloaded by [98.143.143.254] at 05:00 27 June 2013
(O. hemionus), white-tailed deer (O. virginianus), moose (Alces alces), caribou (Rangifer
tarandus), brown brocket deer (Mazama gouazoupira), red brocket deer (M. americana),
pronghorn, American bison (Bos bison), mountain goat (Oreamnos americanus), muskox
(Ovibos moschatus), mountain sheep (Ovis canadensis) and Dall’s sheep (O. dalli). Exotic
species (i.e. those that are non-native) that have been introduced to North America include
feral ass (Equus asinus) and feral horse (E. caballus), which are odd-toed ungulates.
Additional exotic ungulates are members of the Order Artiodactyla, including feral pigs
(Sus scrofa), axis deer (Axis axis), sika deer (C. nippon), sambar deer (C. unicolor), nilgai
(Boselaphus tragocamelus), gemsbok (Oryx gazelle), blackbuck (Antilope cervicapra),
Himalayan thar (Hemitragus jemlahicus), ibex (Capra ibex), European mouflon sheep
(O. aries), Barbary sheep (Ammotragus lervia) and others. Many of these exotics from
Asia and Africa have established free ranging populations in the southwestern USA and
elsewhere.
As a result, there are numerous problems for native species including competition for
natural and financial resources, the transfer of diseases, displacement of native species,
alteration of habitats, altered genetic makeup of native species via interbreeding, and crop
damage. Additionally, exotic species often promote commercialization, which can jeopar-
dize conservation of native species [2].
Pronghorn are the only ungulate endemic to North America and numbers once were
similar to, or even exceeded, those of the American bison (e.g. millions). All native Artio-
dactyla, however, have received considerable attention from biologists and wildlife manag-
ers in North America for numerous reasons. Those ungulates were hunted and used for
food and clothing by the early Europeans that settled North America, and the early Euro-
peans also used ungulates to pay debts to the Crown for their journey to the New World
by sending hides to Europe as payment. Ungulates were also conspicuous among the spe-
cies that North Americans saw ‘disappear’in their lifetimes due to unregulated market
hunting and a lack of management. They were also the group of animals that led early
conservationists to initiate actions to prevent the further demise of all wildlife [3], which
was the beginning of wildlife management in the USA and led to the beginning of the
North American Model of Wildlife Conservation.
Pre-european status of ungulates in North America
Although modern wildlife management began in North America, the roots of the profes-
sion can be traced back to the Old World with chronicles indicating that big game (i.e.
large mammals, which includes ungulates) were particularly important [4] to human beings
for food, clothing and other necessities of life. Numerous practices and policies of modern
management can be traced back to founding European cultures but none of them included
the ‘… organized efforts to protect, wisely use, or deliberately control wildlife populations
and habitats for the societal benefits now associated with wildlife conservation and man-
agement’[4:293]. There were efforts to protect and wisely use wildlife but the use of wild-
life was for society’s leaders instead of for the general population.
Our current management system evolved through three general stages that began with
the arrival of European immigrants in North America. As Europeans entered the New
World, they were met with abundant resources that they exploited with no concern for
conservation. Over time, unregulated harvesting led to concern, outrage and the demands
by sportsmen and other visionaries to stop overexploitation of wildlife and protect the
Conservation and management of North American ungulates 373
Downloaded by [98.143.143.254] at 05:00 27 June 2013
remaining stocks. The final stage was the rise of conservation and associated efforts to pro-
tect and conserve big game and the habitats they depended on. This effort led to conserva-
tion and management for all wildlife [4].
North America was vastly different prior to the arrival of Europeans and their influence
on big game was dramatic. Prior to the arrival of settlers, the exact numbers of big game
were not known but chronicles and estimates by early naturalists placed numbers up to 10
times higher than today. For example, in pre-settlement times, there were estimated to be
13 million mule and black tailed deer, 500,000–2,000,000 mountain sheep and up to mil-
lions of bison, elk and pronghorn [5,6] in North America. Recent estimates place numbers
at <5,000,000 mule deer, >14,000,000 white-tailed deer, 71,000 mountain sheep, 500,000
bison (including bison-livestock hybrids) and 1,000,000 elk and pronghorn [5,6]. Current
numbers are the result of aggressive conservation and management over the past century.
Changes with the arrival of Europeans
There are three general stages of conservation that occur for most imperiled species:
exploitation, concern for their survival and finally conservation through wise use. Several
examples of a species moving through the three stages of conservation exist, and perhaps
the best-known example is the American bison. Bison occurred throughout most of the
USA and the slaughter began east of the Mississippi River and led to the extermination of
the bison in that area by 1830. From 1720 to 1830, the large herds on the plains were
essentially untouched until a variety of factors led to increased human use of the Great
Plains: the gold rush of 1849, the homestead laws of 1862 and the incursion of railroads
in the 1860s attracted more people to the west. The additional human population created
large commercial opportunities to provide fresh and dried meat, tallow, tongues, robes,
chips (manure), sinews and an overwhelming market for bison hides [7].
The bison was attractive for human use: it was large (<1,000 kg), had excellent meat,
was easily killed with rifles, provided an excellent robe and hide and the hair was useful
for insulation. They were nearly extirpated over a short period. Their decimation proceeded
rather slowly at first but the real destruction began around 1830 and was complete in
50 years.
The near extirpation of bison was much more rapid in the western USA when com-
pared to that in eastern North America. Bison hunting expeditions began in 1820 using
Native Americans. In 1840, a single expedition included 1,210 transport carts, 620 Native
Americans and hunters, plus women and children. When the railroads came they adver-
tised the game, split the herds and began the rapid elimination of large populations. From
1872 to 1874, white hunters killed 3,158,730 bison. Native Americans killed about
390,000 and settlers an estimated 150,000 bison. Five thousand hunters and skinners
shipped about 200,000 hides in 1882, but went out of business the following year when
they shipped only 300 hides. By 1889, there were only remnants left of the North Amer-
ican population and only 200 bison in Yellowstone National Park, USA [8]. When the
Cree and white hunters moved up the Cannon Ball River in North Dakota to hunt the
last large herd of bison they ended a chapter of American history started only a few
years before [8]. It was at this time that private individuals, and later visionaries, like
Theodore Roosevelt, George Bird Grinnell, and Gifford Pinchot, and budding conserva-
tion organizations such as the Boone and Crockett Club, intervened and prevented the
total elimination of bison [9]. That effort began with the conservation of the species,
374 P.R. Krausman and V.C. Bleich
Downloaded by [98.143.143.254] at 05:00 27 June 2013
which continues with plans to expand their range throughout Canada, the USA and
Mexico [10,11].
Pronghorn are another very good example of an ungulate that declined sharply and rap-
idly as a result of habitat alteration and human exploitation, but has recovered substantially
as a result of intensive efforts to conserve and manage them [12]. White-tailed deer and
mule deer similarly have recovered from low-levels resulting from overexploitation or hab-
itat modification [13,14]. Management and conservation have led to recovery of most spe-
cies of big game in North America, the majority of which went through similar reductions
in numbers; recovery was only possible with scientific management that started with the
elimination of outright slaughter.
Beginnings of wildlife conservation in North America
Wildlife conservation had its beginnings in Canada and the USA in the late 1800s [9], but
Mexico’s conservation efforts began later. There is a diversity of ungulates in Mexico
including mule and white-tailed deer, brown and red brocket deer, collared and white-
lipped peccaries, tapirs, mountain sheep and pronghorn. Mexico had a longer history of
land exploitation than either Canada or the USA. Prior to the arrival of Spaniards in 1521,
the Mayan Indians practised intensive agriculture, cleared forests, and harvested plants and
animals, all of which destroyed wildlife habitats and locally depleted wildlife populations
[15]. Environmental impacts increased with the arrival of Spaniards as they mined precious
metals, harvested timber, introduced livestock and ranching, enhanced intensive agriculture
and continued with the largely unregulated exploitation of wildlife and fish [16]. With the
introduction of livestock, rangelands were overused and wildlife habitat was transformed,
which continues today [15]. In addition, there has been a continuous alteration of land use
laws in Mexico, resulting in limited incentives for conservation and a national attitude that
did not consider wildlife an economically sustainable resource until recently. Conservation
efforts are rapidly gaining ground in Mexico, and the Wildlife Conservation and Produc-
tion Diversification in the Rural Sector, initiated in 1997, is the most ambitious wildlife
management initiative in Mexico to date [15]. Mexico’s new conservation ethic is benefi-
cial to all wildlife in North America.
Objectives of conservation
The term conservation was popularized by Theodore Roosevelt, who proposed that
renewable natural resources, such as wildlife, forests, ranges and waterpower, if harvested
sustainably and not faster than they can reproduce, could last indefinitely [17:17].
Conservation is, perhaps, best defined as what Roosevelt referred to as ‘wise use’of those
natural resources [9:20], a definition that has persisted in the field of wildlife management
for many decades. Gabrielson [18] identified three primary obstacles to conservation: short-
sightedness of the human race; the tendencies to seek panaceas rather than real remedies;
and lack of knowledge and understanding. Much progress has been made in overcoming
these obstacles in North America but many challenges remain. Indeed, six North American
ungulates (i.e. Sonoran pronghorn (A. americanus sonoriensis), peninsular pronghorn
(A. a. penninsularis), Nelson’s bighorn sheep (O. canadensis nelsoni) in the Peninsular
Ranges of California, USA, Sierra Nevada bighorn sheep (O. c. sierrae), woodland caribou
Conservation and management of North American ungulates 375
Downloaded by [98.143.143.254] at 05:00 27 June 2013
(R. tarandus caribou) and Columbian white-tailed deer (O. virginianus leucurus) have been
afforded protection by the Endangered Species Act (ESA) in the USA, and woodland cari-
bou is afforded protection under Canada’s Federal Species at Risk Act (SARA). Further,
land use changes including urbanization, agricultural conversion, oil or gas development,
and alternative energy projects continue to destroy, modify or fragment habitats upon which
ungulates depend for their existence [19]; habitat conservation must be a primary objective
if large mammals are to persist as sustainable populations. A constantly expanding human
population, both in North America and worldwide, will continue to create challenges to
the conservation of ungulates and all wildlife, and particularly large mammals, that are
dependent upon vast expanses of suitable habitat and the ability to travel long distances
unencumbered between seasonal ranges.
Biology
Understanding life history characteristics of ungulates is critical to their successful manage-
ment. Even the earliest steps towards maintaining viable populations via sustainable use
programmes required a solid education in the biological sciences. At the birth of the scien-
tific discipline of wildlife management, Aldo Leopold [17] identified ecology, ornithology,
mammalogy, botany, entomology, herpetology, parasitology, bacteriology, meteorology,
agronomy, geology, land economics, general zoology, physiology and biometry as the core
science-related disciplines wildlife managers should have some knowledge of. Understand-
ing the natural history of big game and their evolutionary relationships with the landscape
and other species is the foundation upon which all management of large mammals
depends. Over the years, additional knowledge of the habitat for ungulates, information
about their life history characteristics (e.g. birth rates, survival, numbers, density, relation-
ships with other animals and humans), and advanced ways of measuring these characteris-
tics through modelling and statistics, understanding the influences of human beings on
ungulates, and how they are affected by public opinion, cooperative management between
all parties interested in ungulates, structured decision making (that allows for alternate
management) and a host of other topics have been added to the techniques ungulate scien-
tists work with. None, however, replaces an understanding of natural history, basic biology
and evolutionary processes that are so critically important to wildlife conservation [20].
Managing habitat for ungulates
Wildlife management in North America progressed through six stages that elevated
ungulates from the status of unmanaged and overexploited to successfully managed and
conserved: (1) the creation of laws and regulations; (2) appropriate predator control; (3)
the creation of reservation land refuges; (4) artificial replenishment, mainly through reintro-
ductions; (5) environmental controls (i.e. control of disease); and (6) habitat management
[5,7,9,17,18]. Habitat has been defined in numerous ways but it constitutes the resources
and conditions present in an area that produce occupancy, provide for reproduction and
allow survival for a given organism. Habitat is species-specific and is the sum of the
resources needed by a particular organism. These resources include food, cover, water and
special factors that are necessary for reproduction and survival, and habitat includes the
corridors and migratory routes that are necessary for those processes even if they are only
used during short periods annually.
376 P.R. Krausman and V.C. Bleich
Downloaded by [98.143.143.254] at 05:00 27 June 2013
The third step in the evolution of game management (i.e. reservations of lands) was
the first attempt to establish refuges and manage habitats in the western USA, specifi-
cally for ungulates (e.g. Yellowstone National Park in 1872) [9]. Early conservationists
realized that wildlife would not exist without habitat. The National Forest Reserves
Act of 1897 followed the establishment of refuges and provided administration, con-
servation and use of large areas of federal lands, which became national forests in
1905. Most of these lands were habitat for big game, including ungulates, in the
USA [4].
In subsequent years, conservation observers reported the poor condition of these lands
due to overuse by livestock. The western USA was open and grazing by domestic live-
stock was uncontrolled prior to the establishment of the US Forest Service in the early
1900s and the US Bureau of Land Management in 1946. Decades of unrestricted use had
negative consequences for landscapes and the wildlife that depended on them, and so
restrictions were put in place to maintain and enhance some ranges (e.g. Gallatin National
Forest, Montana; Kaibab National Forest, Arizona). These measures were the first recorded
instances of habitat management for ungulate species by resource managers in North
America [4]. As wildlife science progressed, it became apparent that available forage was
one of the most important factors limiting populations of deer. This became the basis of
the idea of density dependence and carrying capacity, or the number of deer or other wild-
life a unit of range could sustain. As a result, many early ungulate habitat investigations
centred on forage conditions and range management practises to improve quality and avail-
ability of forage [4]. As new data and results of studies accumulated, the management of
native ungulates evolved around three major objectives: management of ungulates in con-
cert with other land uses and available habitat, provision for sustainable use of annual sur-
pluses of ungulates (i.e. hunting) and provision of maximum recreation possible for the
public (i.e. photography and viewing). The application of science was primarily directed to
white-tailed deer, mule deer and elk. Rising abundance of ungulates demanded manage-
ment attention and although habitat management programmes for most species were slow
to develop in North America [21], they were established in many state and federal agen-
cies in the USA by the 1950s. Studies of ungulate range and forage production (including
habitat use strategies), use, and condition of animals and their forage became increasingly
common as methods to evaluate range carrying capacity, animal population levels and
human harvest recommendations. As ungulate species were restored to their native land-
scapes, however, the ‘modern’theory that they were controlled by forage supplies did not
always work. Questions arose about the ‘… ability of existing knowledge of deer-habitat
relationships to explain population trends, plus the lack of close or consistent correlation
between population characteristics and trend and measures of browse utilization and condi-
tion, resulted in reduced emphasis on browse and other range surveys as bases for estimat-
ing carrying capacity and determining harvest and other management needs by the late
1970s and 1980s …’ [4:315]. As more data were accumulated, biologists realized that
ungulates had complex relationships with predators, which led to increased interest in
predator–prey relationships.
Understanding ungulate-habitat relationships was greatly enhanced in the 1960s and
1970s with the advent of radiocollar transmitters, telemetry receivers and computers. This
technology increased our understanding of how ungulates use habitat, and how anthropo-
genic influences alter the use of landscapes by ungulates. As technology continues to
increase, investigators will continue to understand these relationships better. This will be
instrumental in allowing human beings and big game to coexist.
Conservation and management of North American ungulates 377
Downloaded by [98.143.143.254] at 05:00 27 June 2013
Predator management
Predation can be an important limiting factor for ungulate populations, although it infre-
quently is viewed in the context of environmental carrying capacity (K; [22]). Predator
control sometimes is desirable, or even necessary, to ensure the establishment, recovery or
persistence of populations of large mammals. It is important, however, to be mindful of
the influence of ungulate population size relative to Kwhen assessing the potential benefits
of removing predators to achieve management objectives. Indeed, determination of whether
a top-down (i.e. predation) or bottom-up (i.e. nutritional resources) process is limiting a
population is important to any decision regarding predator removal. For example, removal
of predators to ‘protect’a population of ungulates that demonstrate the life-history charac-
teristics associated with ungulate populations that are regulated from the bottom up (i.e. by
nutrient availability; [22]) would yield little in terms of allowing such a population to
escape from a low-level equilibrium [23].
Conversely, in populations that exhibit life-history characteristics typical of those regu-
lated by bottom-up processes [22] some benefit(s) to the population under consideration
may be achieved [23]. When invoking predator removal(s) specifically to achieve ungulate
management objectives, an understanding of the role of predation and the prey population
relative to Kis critically important, and holds greater promise than simple consideration of
predator–prey ratios or kill rates [22]. Even in the absence of such knowledge, however,
predator removal programmes sometimes have been invoked, and some have been success-
ful in helping to speed recovery of endangered taxa [24] or to achieve other management
objectives. Predator management programmes will remain an important option for conser-
vationists to have available for their use in the future.
Human dimensions in ungulate management
For many years, managers were mainly concerned with the animal and its habitat as the
two main legs of wildlife management, but by the 1970s the issues faced by managers
were increasingly complex. Indeed, most ungulate populations were stable or were increas-
ing. White-tailed deer increased dramatically altering ecosystems and encroaching into
urban areas, exotic feral horses and burros in the western USA expanded their range and
competed with native ungulates. After the passage of the ESA in the USA and the SARA
in Canada, a wide range of threatened and endangered species had to be considered, and
management moved away from single-species emphases towards management at the eco-
system level. This obviously involved a wider range of public opinion and society became
more vocal with respect to the management of wildlife; hunters were no longer the only
group with a stake in the management of the ungulate resources.
Because wildlife in the USA and Canada belongs to the citizens of each country, it
became important to understand better how those citizens viewed the management of these
large mammals. Thus, human dimensions (i.e. views and attitudes of humans towards
wildlife and wildlife policy) became the third leg of the wildlife management triad, with
the result that interests of non-hunters were incorporated into ungulate management, there
was a new emphasis on non-hunted wildlife, and there was recognition that broad public
interest existed for wildlife, habitats, and most aspects of management. As a result,
throughout the USA and Canada the viewpoints of more and diverse stakeholders were
378 P.R. Krausman and V.C. Bleich
Downloaded by [98.143.143.254] at 05:00 27 June 2013
considered in the management of ungulates, resulting in broad public involvement. Public
hearings, surveys and meetings with interested parties have become integral to twenty-first
century wildlife management.
Wildlife was assumed to be a part of the human environment, and major wildlife and
habitat management issues, including the impacts of changes in grazing management, log-
ging and road development, and mineral development, were subjected to intense public
and professional management scrutiny. Many sportsmen’s groups and others …became
more involved in management decision making and …challenge[d] programmes or data
supporting programmes. Analysis and reporting of habitat and population impacts or the
reviewing of environmental impact statements rapidly became a major task of biologists,
especially in federal agencies [4:316].
Current status of ungulates in North America
The current status of most ungulates in North America is encouraging, and the literature is
replete with examples of programmes that have resulted in the successful restoration or
recovery of many species [5]. Indeed, many species including bison and pronghorn, which
at one time were on the verge of extirpation or even extinction, have recovered substan-
tially in numbers and distribution as a result of intensive conservation efforts. Other spe-
cies, including white-tailed deer and elk, continue to increase in numbers and expand in
distribution as a result of changes in habitat quality or availability; in some cases reaching
numbers that make their management with habitat constraints almost impossible. In the
case of white-tailed deer, this has led to significant overbrowsing of extensive ranges, and
highly problematic ecosystem damage. Most subspecies of caribou, and musk ox, moose
and Dall’s sheep have not yet been impacted greatly by anthropogenic forces and continue
to thrive in large numbers. Their distributions include vast tracts of habitat not currently
subject to intensive human exploitation and, as a result, those ungulates will likely con-
tinue to thrive.
Some other ungulates, however, have required special protection and intensive efforts to
recover them from the threat of extinction. Among these are two subspecies of pronghorn
(A. americana sonoriensis,A. a. penninsularis), one distinct population segment of
bighorn sheep (O. canadensis nelsoni in the peninsular ranges of California) and one
subspecies (O. c. sierrae) of mountain sheep, one subspecies of the mountain ecotype of
woodland caribou (R. tarandus caribou) and a population segment of Columbian white-
tailed deer. All of these are listed as endangered by the US Fish and Wildlife Service and
subject to provisions of the ESA. In Canada, the woodland caribou is a nationally
threatened species and subject to provisions of the Federal SARA.
Challenges for the future
Mahoney and Cobb [19], recently identified a variety of threats to wildlife conservation in
North America. Among these are an increasing human population, climate change, global
economics, urbanization, the creation of novel ecosystems, abundance and superabundance
of certain species, the human-nature divide, the perception that wildlife are vermin,
changes in public perceptions, commercialization of wildlife, funding, and lack of public
Conservation and management of North American ungulates 379
Downloaded by [98.143.143.254] at 05:00 27 June 2013
education in conservation. All of these are important to understand for the conservation of
ungulates in North America, especially funding and climate change.
Regulated hunting, for consumption and sport, has played an important role in these
success stories directly (through the manipulation of populations; [25]) and indirectly
(through the provision of funding for wildlife conservation; [3]), but hunters alone can no
longer assure a sustainable future for wildlife in the USA [26,27] and North America in
general. In the absence of funds to augment the direct use of wildlife as a result of an
interest in hunting for food or sport, funds available for conservation will decline. Indeed,
the continued viability of wildlife populations requires new and additional levels of fund-
ing, and it is appropriate that all citizens contribute to the cost of wildlife conservation
[27]. The limited funds available will be stretched further as increased human population
growth and climate change create new challenges to management.
Predictions have been made that modern climate change will alter habitats and life his-
tory characteristics of ungulate species across North America, yet only limited data are
available that document its influence. Managers and scientists are actively studying climate
change and its effect on wildlife to enhance and predict management and conservation for
the future. Even though research in the arena of climate change effects is in its infancy,
the predicted influences are serious. Climate change has the potential to alter habitats, food
availability, energy expenditure, physical condition, reproduction, parasite loads and sur-
vival for some species [28]. Indeed, caribou are likely to be impacted by a variety of fac-
tors associated with climate change [29–39]. Further, musk ox, moose and Dall’s sheep
can be negatively influenced by increased parasitism caused by increases in temperature
[40–42], and shifts in habitat and populations among some species (e.g. elk, collared pec-
caries and mountain sheep) can be expected with changing climate [43].
Changes in temperature alone are significant for ungulates. Muskox respond negatively
to warm, snowy winters, which have resulted in population declines [44]. Moose popula-
tions have declined due to increased thermoregulatory costs, lower survival and depressed
fecundity with increases in mean summer temperatures [41]. Parturition in some moose
populations is tied to long-term climatic trends and shifts in climate could impact calf sur-
vival [45]. Furthermore, mountain sheep populations at lower elevations with lower precip-
itation and higher temperatures appear more likely to go extinct than those occurring at
higher elevations [46]. Clearly more work needs to be done to understand how climate
change will influence migration, the use of corridors, winter and summer ranges, competi-
tion with livestock and a host of other biological factors.
Conclusions
Healthy and viable populations of native ungulates exist in North America because of
active and visionary management. The ax, plow, rifle and cow contributed significantly to
the demise of ungulates in North America [17] and those are the same tools being used to
reverse the downward trend. The difference has been science-based management. In the
late 1800s, there were few game laws and less enforcement, which resulted in unregulated
harvesting. Visionaries like Theodore Roosevelt and concerned sportsmen saw ungulates
‘disappear’in their lifetimes (much as we are seeing habitat ‘disappear’today) and took
action to reverse the trend. The result was the beginning of scientific wildlife management,
which led to the North American Model of Wildlife Conservation. Initially, the charge for
conservation and management was led by sportsmen but as the human population
380 P.R. Krausman and V.C. Bleich
Downloaded by [98.143.143.254] at 05:00 27 June 2013
increased more and different segments of society demanded a voice in wildlife issues.
Sportsmen still contribute significantly to the active management of ungulates in North
America but other segments of society need to step up and contribute to the funding
efforts. Human population growth, limited funding, climate change and other challenges
will continue to force managers to reevaluate and alter management of big game in North
America. With the wildlife management triad of biological understanding, knowledge of
habitats and human dimension studies, ungulates in North America will continue to thrive.
Acknowledgements
Earlier drafts of this paper were reviewed by J.A. Bissonette and R.D. Brown. Support for
writing the paper was provided by the Department of Biological Sciences, Idaho State
University, the Boone and Crockett Program in Wildlife Conservation, University of
Montana, and the Eastern Sierra Center for Applied Population Ecology (ESCAPE). This
is Professional Paper 098 from the Eastern Sierra Center for Applied Population Ecology.
References
[1] Nowak, R.M., 1999, Walker’s Mammals of the World, 6th Edn, Vol. 2 (Baltimore, MD: John Hopkins
University Press), p. 1051.
[2] Mungall, E.C., 2000, Exotics. In: S. Demarais and P.R. Krausman (Eds.) Ecology and Management of Large
Mammals in North America (Upper Saddle River, NJ: Prentice Hall), pp. 736–764.
[3] Williamson, L.L., 1987, Evolution of a landmark law. In: H. Kallman, C.P. Agee, W.R. Goforth and J.P.
Linduska (Eds.) Restoring Amerca’s Wildlife 1937–1987 (Washington, DC: U.S. Department of the Interior),
pp. 1–17.
[4] Mackie, R.J., 2000, History and management of large mammals in North America. In: S. Demarais and P.R.
Krausman (Eds.) Ecology and Management of Large Mammals in North America (Upper Saddle River, NJ:
Prentice Hall), pp. 292–320.
[5] Kallman, H., Agee, C.P., Goforth, W.R. and Linduska, J.P. (Eds.), 1987, Restoring America’s Wildlife 1937–
1987 (Washington, DC: U.S. Department of the Interior).
[6] Valdez, R. and Krausman, P.R., 1999, Mountain Sheep of North America (Tucson, AZ: University of
Arizona Press), pp. 1–353.
[7] Krausman, P.R., 2002, Introduction to Wildlife Management; the Basics (Upper Saddle River, NJ; Prentice
Hall), pp. 1–478.
[8] Hornaday, W.T., 1889, The Extermination of the American Bison (Washington, DC: U.S. Government Print-
ing Office), pp. 464–465.
[9] Reiger, J.F., 1975, American Sportsmen and the Origins of Conservation (New York: Winchester Press).
[10] Sanderson, E.W., Redford, K.H., Weber, B., Aune, K., Baldes, D., Berger, J., Carter, D., Curtin, C., Derr, J.,
Dobrott, S., Fearn, E., Fleener, C., Forrest, S., Gerlach, C., Gates, C.C., Gross, J.E., Gogan, P., Grassel, S.,
Hilty, J.A., Jensen, M., Kunkel, K., Lammers, D., List, R., Minkowski, K., Olson, T., Pague, C., Robertson,
P.B. and Stephenson, B., 2008, The ecological future of the North American bison: conceiving long-term,
large-scale conservation of wildlife. Conservation Biology, 22, 252–266.
[11] Gates, C.C., Freese, C.H., Gogan, P.J.P. and Kotzman, M. (Eds.), 2010, American Bison: Status Survey and
Conservation Guidelines 2010 (Gland: International Union for the Conservation of Nature).
[12] Cadieux, C.L., 1987, Pronghorn antelope: great plains rebound. In: H. Kallman, C.P. Agee, W.R. Goforth
and J.P. Linduska (Eds.) Restoring Amerca’s Wildlife 1937–1987 (Washington, DC: U.S. Department of the
Interior), pp. 133–143.
[13] McCabe, R.E. and McCabe, T.R., 1984, Of slings and arrows: an historical retrospection. In: L.K. Halls
(Ed.) White-Tailed Deer: Ecology and Management (Harrisburg: Stackpole Books), pp. 19–72.
[14] Mackie, R.J., 1987, Mule deer. In: H. Kallman, C.P. Agee, W.R. Goforth and J.P. Linduska (Eds.) Restoring
Amerca’s Wildlife 1937–1987 (Washington, DC: U.S. Department of the Interior), pp. 265–271.
[15] Valdez, R., Guzman-Aranda, J.C., Abarca, F.J., Tarango-Arámbula, L.A. and Sánchez, F.C., 2006, Wildlife
conservation and management in Mexico. Wildlife Society Bulletin, 34, 270–282.
[16] Tinker, B.,1978, Mexican Wilderness and Wildlife (Austin: University of Texas Press).
[17] Leopold, A., 1933, Game Management (New York: Charles’s Scribners Sons), pp. 1–481.
[18] Gabrielson, I.N., 1941, Wildlife Conservation (New York: The Macmillan Company).
[19] Mahoney, S.P. and Cobb, D., 2010, Future challenges to the model: why collapse is possible and alteration
inevitable. The Wildlife Professional, 4(3), 83–85.
Conservation and management of North American ungulates 381
Downloaded by [98.143.143.254] at 05:00 27 June 2013
[20] Bleich, V.C. and Oehler, M.W.Sr., 2000, Wildlife education in the United States: thoughts from agency biol-
ogists. Wildlife Society Bulletin, 28, 542–545.
[21] Harmon, K.W., 1987, Managing habitats. In: H. Kallman, C.P. Agee, W.R. Goforth and J.P. Linduska (Eds.)
Restoring Amerca’s Wildlife 1937–1987 (Washington, DC: U.S. Department of the Interior), pp. 79–85.
[22] Bowyer, R.T., Person, D.K. and Pierce, B. M., 2005, Detecting top-down versus bottom-up regulation of
ungulates by large carnivores: implications for conservation of biodiversity. In: J.C. Ray, K.H. Redford, R.S.
Steneck and J. Berger (Eds.) Large Carnivores and the Conservation of Biodiversity (Washington, DC:
Island Press), pp. 342–361.
[23] Bleich, V.C. and Pierce, B.M., 2005, Management of mountain lions in California. In: E.L. Buckner and J.
Reneau (Eds.) Records of North American Big Game, 12 Edn (Missoula, MT: Boone and Crockett Club),
pp. 63–69.
[24] Rominger, E.M., Bleich, V.C. and Goldstein, E.J., 2006, Bighorn sheep, mountain lions, and the ethics of
conservation. Conservation Biology, 20, 1041.
[25] White, G.C. and Bishop, C.J., 2010, A bountiful harvest for science. The Wildlife Professional, 4,42–46.
[26] Williams, S., 2010, Wellspring of wildlife funding: how hunter and angler dollars fuel wildlife conservation.
The Wildlife Professional, 4,35–38.
[27] Regan, R.J., 2010, Priceless, but not free: why all nature lovers should contribute to conservation. The Wild-
life Professional, 4,39–41.
[28] Gunn, A. and Skogland, T., 1997, Responses of caribou and reindeer to global warming. Ecological Studies,
124, 189–200.
[29] Chapin, F.S., Shaver, G.R., Giblin, A.E., Nadelhoffer, K.J. and Laundre, J.A., 1995, Responses of arctic
tundra to experimental and observed changes in climate. Ecology, 76, 694–711.
[30] Brotton, J. and Wall, G., 1997, Climate change and the Bathurst caribou herd in the Northwest Territories,
Canada. Climate Change, 35,35–52.
[31] Mörschel, F.M. and Klein, D.R., 1997, Effects of weather and parasitic insects on behavior and group
dynamics of caribou of the Delta Herd, Alaska. Canadian Journal of Zoology, 75, 1659–1670.
[32] Walsh, N.E., McCabe, T.R., Welker, J.M. and Parsons, A.N., 1997, Experimental manipulations of snow-
depth: effects on nutrient content of caribou forage. Global Change Biology, 3, 158–164.
[33] Lenart, E.A., Bowyer, R.T., Ver Hoef, J. and Ruess, R.W., 2002, Climate change and caribou: effects of
summer weather on forage. Canadian Journal of Zoology, 80, 664–678.
[34] Harding, L.E., 2004, The future of Peary caribou (Rangifer tarandus pearyi) in a changing climate.
Proceeding of the Species at Risk 2004 Recovery Conference 1,1
–9.
[35] McNeil, P., Russell, D.E., Griffith, B., Gunn, A. and Kofinas, G.P., 2005, Where the wild things are:
seasonal variation in caribou distribution in relation to climate change. Rangifer, 16,51–63.
[36] Racey, G.D., 2005, Climate change and woodland caribou in northwestern Ontario: a risk analysis. Rangifer,
16, 123–136, Special Issue.
[37] Rupp, T.S., Olson, M., Adams, L.G., Dale, B.W., Joly, K., Henkelman, J., Collins, W.B. and Starfield, A.M.,
2006, Simulating the influences of various fire regimes on caribou winter habitat. Ecological Applications,
16, 1730–1743.
[38] Joly, K., Cole, M.J. and Jandt, R.R., 2007, Diets of overwintering caribou, Rangifer tarandus, track decadal
changes in arctic tundra vegetation. Canadian Field Naturalist, 121, 379–383.
[39] Tews, J.M., Ferguson, A.D. and Fahrig, L., 2007, Modeling density dependence and climatic disturbances in
caribou: a case study from the Bathurst Island complex, Canadian High Artic. Journal of Zoology, 272,
209–217.
[40] Kutz, S.J., Hoberg, E.P., Nagy, J., Polley, L. and Elkin, B., 2004, “Emerging”parasitic infections in Arctic
ungulates. Integrative and Comparative Biology, 44, 109–118.
[41] Murray, D.L., Cox, E.W., Ballard, W.B., Whitlaw, H.A., Lenarz, M.S., Custer, T.W., Barnett, T. and Fuller,
T.K., 2006, Pathogens, nutritional deficiency, and climate influences on a declining moose population. Wild-
life Monographs, 166,1–30.
[42] Jenkins, E.J., Veitch, A.M., Kutz, S.J., Hoberg, E.P. and Polley, L., 2006, Climate change and the epidemiol-
ogy of protostrongylid nematodes in northern ecosystems: Parelaphostrongylus odocoilei and Potostrongylus
stilesi in Dall’s sheep (Ovid d. dalli). Parasitology, 132, 387–401.
[43] Albert, S., Ramotnik, C.A. and Schmitt, C.G., 2004, Collared peccary range expansion in northwestern New
Mexico. Southwestern Naturalist, 49, 524–528.
[44] Forchhammer, M.C., Post, E., Stenseth, N.C. and Boertmann, D.M., 2002, Long-term responses in arctic
ungulate dynamics to changes in climatic and trophic processes. Population Ecology, 44,113–120.
[45] Bowyer, R.T., Van Ballenberghe, V. and Kie, J.G., 1998, Timing and synchrony of parturition in Alaskan
moose: long-term versus proximal effects of climate. Journal of Mammalogy, 79, 1332–1344.
[46] Epps, C.W., McCullough, D.R., Wehausen, J.D., Bleich, V.C. and Rechel, J.L., 2004, Effects of climate
change on population persistence of desert-dwelling mountain sheep in California. Conservation Biology, 18,
102–113.
382 P.R. Krausman and V.C. Bleich
Downloaded by [98.143.143.254] at 05:00 27 June 2013
