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Conservation and management of ungulates in North America



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.
Conservation and management of ungulates in
North America
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:
International Journal of Environmental Studies, 2013
Vol. 70, No. 3, 372382,
Ó2013 Taylor & Francis
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(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 Dalls 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 mouon 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
As a result, there are numerous problems for native species including competition for
natural and nancial 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 disappearin 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 benets 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 societys 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
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remaining stocks. The nal 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 inuence
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, 500,0002,000,000 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 nally 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 ries, 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 rst 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
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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 modication [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 scientic 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
Mexicos 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 sh [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 Diversication in the Rural Sector, initiated in 1997, is the most ambitious wildlife
management initiative in Mexico to date [15]. Mexicos new conservation ethic is bene-
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 indenitely [17:17].
Conservation is, perhaps, best dened as what Roosevelt referred to as wise useof those
natural resources [9:20], a denition that has persisted in the eld of wildlife management
for many decades. Gabrielson [18] identied 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), Nelsons 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
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(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 Canadas 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.
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-
tic discipline of wildlife management, Aldo Leopold [17] identied 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 inuences 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) articial 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 dened 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-specic 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
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The third step in the evolution of game management (i.e. reservations of lands) was
the rst attempt to establish refuges and manage habitats in the western USA, speci-
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 rst 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 moderntheory 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
predatorprey 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 inuences 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
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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 inuence of ungulate population size relative to Kwhen assessing the potential benets
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 protecta 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 benet(s) to the population under consideration
may be achieved [23]. When invoking predator removal(s) specically 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
predatorprey 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
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considered in the management of ungulates, resulting in broad public involvement. Public
hearings, surveys and meetings with interested parties have become integral to twenty-rst
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 sportsmens 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 signicant overbrowsing of extensive ranges, and
highly problematic ecosystem damage. Most subspecies of caribou, and musk ox, moose
and Dalls 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 identied 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
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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 inuence. 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 inuences 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 [2939]. Further, musk ox, moose and Dalls sheep
can be negatively inuenced by increased parasitism caused by increases in temperature
[4042], 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 signicant 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 inuence migration, the use of corridors, winter and summer ranges, competi-
tion with livestock and a host of other biological factors.
Healthy and viable populations of native ungulates exist in North America because of
active and visionary management. The ax, plow, rie and cow contributed signicantly 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
disappearin their lifetimes (much as we are seeing habitat disappeartoday) and took
action to reverse the trend. The result was the beginning of scientic 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
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increased more and different segments of society demanded a voice in wildlife issues.
Sportsmen still contribute signicantly 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.
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.
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... Therefore, while for regions such as Europe, Oceania and South America, a higher amount of data are available, there are regions that are not well represented, such as Africa. For the USA, we integrated the information in GRIIS with additional data extracted from Feldhamer et al. (2003) and Krausman and Bleich (2013). ...
... In Argentina, New Zealand and Cuba 19, 17 and 15 invasive alien mammals with impacts on health are reported (Fig. 10.2). Feldhamer et al. (2003) and Krausman and Bleich (2013).) Fig. 10.2. ...
... (From and Feldhamer et al. (2003) and Krausman and Bleich (2013).) Australia, Brazil, Mexico, the Bahamas and the Czech Republic are characterized by the highest proportions of alien mammals impacting human health out of the total number of invasive mammals reported in the country (range from 93% to 96%). ...
... Environmental Evidence *Correspondence: 1 National Climate Adaptation Science Center, U.S. Geological Survey, Reston, VA, USA Full list of author information is available at the end of the article conditions [8,9], and predator communities [8] are of increasing concern to ungulate managers [10]. Of these, an improved understanding of the effects of changing climate conditions has been highlighted as a key information need [11][12][13]. Climatic variation occurs across multiple spatial and temporal scales, and understanding the impacts of both short-and long-term changes will provide valuable information to wildlife and land managers. Climate is an important driver of ungulate life-history characteristics, population dynamics, and migratory behaviors and changes in climate can directly or indirectly affect the growth, development, fecundity, dispersal, demographic trends, and long-term viability of populations [9,13] as well as the timing and locations of migratory movements [14,15]. ...
... This systematic map will focus on the fifteen ungulate species of the Order Artiodactyla native to North America [11]. We will describe the abundance and distribution of evidence relating to the impacts of climate variability and change on the life-history characteristics, population dynamics, and migratory behaviors of these species by gathering evidence on the topic from across the continent. ...
Full-text available
Background Climate is an important driver of ungulate life-histories, population dynamics, and migratory behaviors, and can affect the growth, development, fecundity, dispersal, and demographic trends of populations. Changes in temperature and precipitation, and resulting shifts in plant phenology, winter severity, drought and wildfire conditions, invasive species distribution and abundance, predation, and disease have the potential to directly or indirectly affect ungulates. However, ungulate responses to climate variability and change are not uniform and vary by species and geography. Here, we present a systematic map protocol aiming to describe the abundance and distribution of evidence on the effects of climate variability and change on ungulate life-histories, population dynamics, and migration in North America. This map will help to identify knowledge gaps and clusters of evidence, and can be used to inform future research directions and adaptive management strategies. Methods We will catalogue evidence on how climate variability and change affect the life-histories, population dynamics, and migration patterns of the fifteen ungulate species native to North America. We will search both academic and grey literature, using academic journal databases and specialist websites. Articles will be screened for inclusion at the title/abstract and full-text levels, and data will be extracted from articles that pass the full-text review. These data will be summarized quantitatively, visually, and with a narrative review to describe the distribution and abundance of evidence on the effects of climate variability and change on ungulates in North America.
... In the context of increasing populations of wild ungulates throughout northern America and Europe (Krausman & Bleich, 2013;Ramirez et al., 2018;Rooney, 2001), how much could these populations impact the stoichiometry of forests and opened landscapes? Would this impact reinforce or buffer the general trend of Nsaturation observed in many forests of the northern hemisphere (Perakis & Hedin, 2002)? ...
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Biological control of nutrient cycles is well documented in aquatic ecosystems, where consumer‐driven recycling by herbivores can significantly impact ecosystem stoichiometry. In contrast, little is known in terrestrial ecosystems, where there is evidence that herbivores can also impact ecosystem stoichiometry. I studied a stoichiometric model of the soil‐plant‐herbivore system. The model shows that herbivores influence the ecosystem stoichiometry mainly through the direct and indirect controls of ecosystem inputs and losses, in a more complex way than predicted by the classic consumer‐driven recycling theory. Overall, it shows that herbivores affect nutrient ratios in terrestrial ecosystems mostly independently of their own stoichiometric ratios, and that their impact may be different in forest versus grassland. The results highlight the sensitivity of terrestrial ecosystems to elusive actors, negligible in biomass but capable of modifying nutrient loss rates with major impacts on nutrient cycles and ecosystem stoichiometry. I addressed the impact of herbivores on nitrogen and phosphorus recycling in terrestrial ecosystems (forest and grassland) with a stoichiometric model. My results suggest that herbivores impact ecosystem stoichiometry mostly independently of their own stoichiometry, and may promote either N or P limitation depending on the propensity for loss of soil organic and mineral nitrogen relative to phosphorus, and on the N:P ratio exported by the herbivores from the ecosystem.
... For example, shorebird populations are decreasing (Butchart et al., 2010), half of the world's ungulate and carnivore species are at higher risk of extinction than in the 1970s (Bowyer et al., 2019;Di Marco et al., 2014), and 43% of amphibian species have experienced recent declines with 33% being at risk of extinction (Stuart et al., 2004). The challenges facing wildlife populations include loss and fragmentation of habitat (Bowyer et al., 2019;Knick et al., 2010;Van der Ree et al., 2011), global climate change (Brown & Thorpe, 2008;Hoegh-Guldberg & Bruno, 2010), overexploitation (Ludwig, 2001;Stuart et al., 2004) and increasing human-wildlife conflict due to growing human populations (Hammerson et al., 2017;Krausman & Bleich, 2013;Singh et al., 2010). While much is being done to mitigate these losses, these challenges will likely increase in the future. ...
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Abstract Many global wildlife populations are experiencing unprecedented declines. Estimates of population abundance are needed to effectively manage common species and to conserve vulnerable species. Camera traps have advanced as wildlife monitoring tools for ungulates and can provide improved methods of estimating population abundance. Little is known, however, about how camera traps set for ungulates compare with traditional methods (e.g., ground and aerial surveys) used simultaneously. From 2012 to 2014, we captured and radio collared 34 female and 32 male bighorn sheep (Ovis canadensis) in a closed population in Utah, USA. Each collar had a unique letter and number combination. We then estimated number of young, females and yearlings, males and population abundance using multiple methods simultaneously: helicopter surveys, resight surveys performed from the ground, camera trap surveys using marked but not individually identifiable individuals and camera trap surveys using marked and individually identifiable animals. All methods estimated similar abundance. Across years, ages and sexes, however, camera trap surveys produced the most consistent and precise estimates of abundance for adult females and yearlings, lambs and the population. That method was less intrusive and safer than helicopter surveys. Our results indicate that camera trap surveys using photographs of marked animals in which the majority of the population visits a specific resource can produce precise estimates of abundance that are safer, as well as less intrusive and expensive than traditional methods. Using camera traps also creates a permanent record of photographs that can be archived and reanalyzed to answer future ecological and population questions. Finally, this method of estimating abundance can be used in other areas with ungulates that congregate around resources (e.g., watering sites or mineral licks).
... Strong density-dependence, in body growth and recruitment, is typical (Eberhardt, 2002;Bonenfant et al., 2009). Those patterns, and the environments inhabited by ungulates, have been at the forefront of important discoveries in ecology (McCullough, 1979;Bleich et al., 1997), evolution (Boyce, 1988), and conservation (Cain et al., 2008;Krausman and Bleich, 2013). This Research Topic provides an overview and expansion of those advances, especially movement ecology (Mysterud et al., 2011), sociality and mating systems (Bowyer et al., 2020), the role of individual heterogeneity in population biology (Plard et al., 2012), responses to predation (Fortin et al., 2009), and evolutionary tradeoffs among biological functions (Gaillard et al., 2000). ...
... Threats like hunting, habitat loss, accidents and pollution (Sheikh and Molur 2004), population explosion (Komers andCurman 2000, Krausman andBleich 2013), genetic troubles (Jnawali et al. 2011, Purvis et al. 2000, stress and illegal killing have caused the species to be 'Extinct in the Wildlife' in Pakistan (Sheikh and Molur 2004;Nemat et al. 2013). Therefore, different conservation strategies like protection laws (Ali et al. 2011), captive breeding, hormone-mediated conservation, artificial insemination (Sontakke et al. 2009;Sagar and Antoney 2017) and religious affiliation (Kankane 2013;Kankane 2014;Mohapatra 2014) were adopted. ...
Full-text available
Blackbuck is the existing members of genus Antilope which is most elegant and graceful among all Antelopes of Asia with distinct sexual dimorphism. Blackbuck show endemism in Pakistan, Nepal and India occupying mainly the semi-arid grassland areas. The name of species attributes towards the dark brownish to blackish coat color of male species. While female and the young ones are tawny or yellow. Chin undersides of legs and chest are white in both male and female individuals. Males have whorled horns with 79cm length with absence in females. The average body length of this animal is 100-150cm with the tail length of 10-17cm. The average body weight for male is 20-57kg and for female 19-33kg. Thin grassy forests, open and semi-desert areas are good habitat for it. Being diurnal and herbivorous, it acts as both grazers and browsers. It is the fastest animal with an average speed of 80km/h. It remains reproductively functional whole year. Reproductive disorders including dystocia and different infectious diseases due to ectoparasites, endoparasites, bacteria and viruses affect these species. Habitat loss, stress, illegal killing and genetic troubles cause the species to be ‘Extinct in the Wildlife’ in Pakistan so conservation strategies are underway for species protection. It is indispensable for zoologists and conservational biologists to observe the species for its conservation and confronting threats. Review article highlighted necessary information about species, which will clear the way for further research on species.
... The successful conservation of large mammals (e.g. ungulates) has been achieved in part by a good knowledge of its biology and how they use habitats Krausman & Bleich [19]. In this regard, animals participate actively in the natural restauration of vegetation. ...
The use of georeferenced information on the presence of a species to predict its distribution across a geographic area is one of the most common tools in management and conservation. The collection of high-quality presence-absence data through structured surveys is, however, expensive, and managers usually have more abundant low-quality presence-only data collected by citizen scientists, opportunistic observations, and culling returns for game species. Integrated Species Distribution Models (ISDMs) have been developed to make the most of the data available by combining the higher-quality, but usually less abundant and more spatially restricted presence-absence data, with the lower quality, unstructured, but usually more extensive and abundant presence-only data. Joint-likelihood ISDMs can be run in a Bayesian context using INLA (Integrated Nested Laplace Approximation) methods that allow the addition of a spatially structured random effect to account for data spatial autocorrelation. These models, however, have only been applied to simulated data so far. Here, for the first time, we apply this approach to empirical data, using presence-absence and presence-only data for the three main deer species in Ireland: red, fallow and sika deer. We collated all deer data available for the past 15 years and fitted models predicting distribution and relative abundance at a 25 km2 resolution across the island. Models' predictions were associated to spatial estimate of uncertainty, allowing us to assess the quality of the model and the effect that data scarcity has on the certainty of predictions. Furthermore, we validated the three species-specific models using independent deer hunting returns. Our work clearly demonstrates the applicability of spatially-explicit ISDMs to empirical data in a Bayesian context, providing a blueprint for managers to exploit unused and seemingly unusable data that can, when modelled with the proper tools, serve to inform management and conservation policies.
Shed antler hunting (i.e., collecting cast cervid antlers) has increased in popularity during the past decade, but little is known about how this recreational activity affects ungulate movements and space use. We placed geographic positioning system (GPS)‐collars on 133 female and male bighorn sheep (Ovis canadensis), bison (Bison bison), and mule deer (Odocoileus hemionus) to quantify their movements and space use during shed antler hunts compared with those behaviors during helicopter surveys in Utah, USA, from 2012 to 2015. For each species, we calculated means and 95% confidence intervals for distance moved during 90‐minute segments (16 points/day) pre‐event (control, 7 consecutive days prior to event), event (1–2 days), and post‐event (7 consecutive days after event) for shed antler hunts and helicopter surveys. We also compared use of space for each species during these events. Female bighorn sheep did not increase distance moved or substantially change space use during shed antler hunts and helicopter surveys. Male bighorn sheep increased distance moved 41% on average during shed antler hunts and by 2.02 times during helicopter surveys but did not change space use during those events. Female bison increased distance moved 15% on average during shed antler hunts and 30% during helicopter surveys. Mule deer increased distance moved and altered space use the most during shed antler hunts; females increased distance moved 97%, and 54% of females moved a mean distance of 742 ± 642 (SD) m (range = 9–3,778 m) outside of their home ranges during those hunts for a mean of 9.2 ± 9.4 hours (range = 1.5 to 41 hr). Male mule deer increased distance moved by 2.10 times on average during shed antler hunts, and 82% of males moved a mean distance of 1,264 ± 732 m (range = 131–3,637 m) outside of their home ranges during those hunts for a mean of 12.6 ± 7.6 hours (range = 4.5–33 hr). Our results provide timely information about how legal shed antler hunting affects movements and space use of female and male ungulates, especially mule deer, and can guide the conservation of ungulate populations and their habitat. © 2021 The Wildlife Society. During legal shed antler hunts, mule deer moved the farthest and were displaced the most and for the longest time from their home ranges. Our results will help wildlife managers understand the effects of shed antler hunting on movement and space use of ungulates and provide timely information that can help conserve ungulate populations and their habitat.
Full-text available
Biologists often must use incomplete information to make recommendations concerning harvest of large mammals. Consequently, those recommendations must draw on a firm understanding of the ecology of the species in question, along with selection of the most applicable population characteristics on which to base harvest-both essential components for prudent management. Density-dependent processes, which are ubiquitous among populations of large mammals, may be counterintuitive because of unexpected patterns in recruitment coincident with changes in population size. Misconceptions concerning population dynamics of ungulates also can occur when demo-graphics are based solely on correlations with environmental factors. Further, the concept of a harvestable surplus can be misleading for managing ungulate populations, because of the parabolic relationship between population size and number of recruits-harvest determines the surplus rather than vice versa. Understanding consequences of mortality, especially relative components of compensatory or additive mortality, also is necessary. Knowledge of the proximity of an ungulate population to ecological carrying capacity (K) is required to fully assess whether most mortality is compensatory or additive. We describe selected life-history traits and population characteristics of ungulates useful in parametrizing where populations are in relation to K, thereby allowing for a reasonable harvest despite some uncertainty in population size. We advocate an adaptive-management approach while monitoring those life-history traits to evaluate the suitability of a particular harvest strategy. ALCES VOL. 56: 15-38 (2020)
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Enhancing the preparation of students for careers in wildlife conservation has been an objective of The Wildlife Society for many years. Among recent graduates, we have detected a general decline in knowledge of natural history and evolutionary biology. We suggest that additional formal training in those disciplines will result in personnel that are better prepared to contribute in meaningful ways to wildlife conservation in the future. Agency personnel and university faculty share responsibility for ensuring that adequately trained individuals are available to meet the needs of conservation agencies.
Full-text available
Global warming will initiate a cascade of changes throughout the arctic ecosystems; predicting their magnitude and direction becomes more complex at the higher trophic levels. For top herbivores such as reindeer and caribou Rangifer tarandus, we have to consider the effects of climate change on lower trophic levels (forage), as well as the more direct effects on the animals themselves.
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Woodland caribou (Rangifer tarandus caribou) range occupancy and populations have declined in northwestern (NW) Ontario over the last 100 years primarily due to human-induced factors. Recovery efforts are underway to halt this decline by reducing risk factors. Climate forecasts suggest a 4—5 oC increase in May—August mean temperature over the next century with little change in precipitation. Resulting increases in extreme weather events and increased fire weather severity will likely increase the amount of forest burned, reduce the area of older forest, alter distribution and abundance of forest tree species and plant communities, and increase abundance of alternate prey. The reduced amount of older forest preferred by caribou will be in greater demand by the forest industry leading to more conflict over ecological and economic values. Most of these factors will increase risk to caribou survival. Although forests may experience enhanced productivity, forest management practices will try to adapt harvest, regeneration, silviculture and fire management practices to both maintain economic benefits and increase the ability of forests to sequester carbon. The interaction of climate-induced forest change and forest management practices adds uncertainty to caribou conservation efforts at the southern edge of its current range. This uncertainty reinforces the need for a precautionary approach to forest management, increased research and monitoring effort, sustained emphasis on caribou recovery, and careful rationalization of restoration efforts where greatest opportunities for success may be realized.