This paper summarizes information from the literature and personal observation to present an overview of the effects of predators on big game populations. Specifically it examines control of prey populations, culling of inferior prey, and stimulation of prey productivity. Food habit investigations were omitted because of the impossibility of evaluating the impact of predation by this approach because of a lack of population data

Beasom, Samuel L.

English

DigitalCommons@University of Nebraska

University of Nebraska - Lincoln DigitalCommons@University of Nebraska - Lincoln Great Plains Wildlife Damage Control Workshop Proceedings Wildlife Damage Management, Internet Center for December 1977 PREDATION ON BIG GAME AND ITS MANAGEMENT Samuel L. Beasom USDA Forest Service, Rocky Mountain Forest and Range Experiment Station Follow this and additional works at: https://digitalcommons.unl.edu/gpwdcwp  Part of the Environmental Health and Protection Commons Beasom, Samuel L., "PREDATION ON BIG GAME AND ITS MANAGEMENT" (1977). Great Plains Wildlife Damage Control Workshop Proceedings. 236. https://digitalcommons.unl.edu/gpwdcwp/236 This Article is brought to you for free and open access by the Wildlife Damage Management, Internet Center for at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Great Plains Wildlife Damage Control Workshop Proceedings by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. 259PREDATION ON BIG GAME AND ITS MANAGEMENTSamuel L. BeasomUSDA Forest ServiceRocky Mountain Forest and Range Experiment StationThis paper summarizes information from the literature and personalobservation to present an overview of the effects of predators on big gamepopulations. Specifically it examines control of prey populations, cullingof inferior prey, and stimulation of prey productivity. Food habit investigationswere omitted because of the impossibility of evaluating the impact of predationby this approach because of a lack of population data.Control of Prey PopulationsA population is controlled (whether by predation or another means) when? total deaths equal or exceed births. This control stabilizes populations and.keeps them within the limits imposed by their habitats.Populations of most species of native ungulates have periodically irruptedand then quickly declined the past 100 years (Keith 1974). This was a result ofdecimation of predator populations (Leopold 1943) and/or a major discrepancybetween existing population levels and food supplies and cover (Riney 1964).Keith (1974) indicated interactions between these two forces make it difficultto determine which is the most important, but he cited numerous examples thatimplicate a lack of predation as the chief cause of the irruptions.The effect of predation on populations of large ungulates is mostobjectively evaluated by predator control, but this method has producedvarying results. Jones (1949), in Texas, was unable to demonstrate a difference.in pronghorn antelope (Antilocapra americana) fawn survival between treatedL'and untreated areas. He attributed the lack of treatment effect to ineffectivepredator control. Later investigations in the same general area (Bailey 1973,Uzzell  1973) revealed a marked increase in pronghorn net productivity where260coyotes (Canis latrans) were intensively removed. Similar investigations inArizona (Arrington and Edwards 1951), Utah (Udy 1953),  and Wyoming (Yoakum1968 as interpreted by Knowlton 1968) attributed increases in net reproductionand in total pronghorn populations to intensive coyote control.Predator control varied in effectiveness of alleviating depredations onyoung deer (Odocoileus spp.) and caribou (Rangifer tarandus). Longhurstet al. (1952) found no evidence that fawn survival increased where coyoteswere trapped heavily compared to the control area. Beasom (1974a, 1974b)reported an approximately 300% and 400% increase in white-tailed deer(Odocoileus virginianus) net fawn production during a 2-year period on a studyarea in Texas. The deer population density increased by about 28% on thetreated area and declined by about 12% on the untreated area. Daniel (1975)found an inverse correlation between coyote abundance and percent fawnproduction. Although he noted a higher net fawn production on the treatedareas, deer population density did not increase above that on the untreated Lareas each year. It is not known where the density level of the deer herdsin these two Texas studies were located in relation to the carrying capacitiesof the respective habitats, so it is difficult to relate the reported densitychange differences to this important factor. In Newfoundland, Bergerud (1971)found that intensive lynx (Lynx canadensis) control on one study area wasfollowed by an approximately 100% increase in caribou calf survival by fallcompared to an untreated area. Data on the change in caribou populationdensity between the areas were not available.In general, predators have the potential to curtail most ungulate populationswhen proper circumstances prevail. Errington (1946) suggested that canid.qpredators can control ungulate populations by preying mostly on the young,because of the relatively low productivity of large herbivores. Allee  et al.(1949) generalied that, to control any population, the net productivity input261must be curtailed. The level of predation on big game populations in anyarea is doubtless a result of the density of the predator population, thebehavioral reactions of the game animals, the food predilections of thepredator, and the abundance and quality of alternative foods for the predator.Although predators apparently influence ungulate populations, unlimitedpopulation growth in the absence of predation has not been demonstrated. Whenthe population reaches or surpasses the habitat carrying capacity, furthergrowth is slowed by other mortality factors. Controlling predatory animalsto increase an already high ungulate population would be fruitless.Culling of Inferior PreyA common view is that predators serve to maintain healthy prey populationsrby culling the sick and weak or otherwise inferior individuals. Mech (1970)included young prey animals as biologically inferior. This discussion, however,does not consider all young animals to be biologically inferior.Selection of strong or weak prey depends on the hunting habits of thepredator. For example, stalking predators, such as cats, usually kill bysurprise attack, which is relatively independent of the condition of the prey(Hornocker 1970, Hirst 1965). Predators which chase, and elicit a flight responsefrom their prey, such as wolves (Canis spp.) and spotted hyenas (Crocuta crocuta),tend to take a disproportionate number of poor condition animals as prey (Mech1970, Kruk 1972). The coyote cannot be assigned either category because ofits diverse hunting patterns and feeding habits. It probably hunts big game bythe chase-kill method (Murie 1940). This hunting behavior killed a dispropor-tionate number of old, injured or young deer. Although reports of coyoteskilling full grown, healthy wild ungulates are common, they are probably theexception. Size alone, since they rarely hunt in groups larger than two (Fox1971), would preclude a coyote's preying on adults of most native North Amc~ricnnungulates. Coyotes feed mostly on small mammals and birds but will sometimeskill deer and other large animals, usually young or weakened individuals(Mech 1970).Coyotes reportedly prey heavily on adult deer in winter when prey bodycondition presumably declines (Murie 1940, Knowlton 1964, Knowles 1976).Coyote predation at this time may be disproportionate toward adult malesbecause they are physiologically exhausted from the rut as well as the poorerfood conditions (Knowlton 1964, Beasom unpublished data). Hood (personalcommunication), while conducting a telemetry study to investigate deermovements in South Texas, noted a similar disproportionate coyote predationon collared adult bucks from late winter to early spring.Coyote predation on pronghorns indicates that this predator does not takedisproportionately more unfit ungulates (Bruns 1970). Murie (1940) presentedseveral accounts of pronghorns easily outdistancing coyotes on unfenced rangeswithin Yellowstone National Park. However, coyotes apparently have learned Ito use fences (especially net wire) to capture even healthy pronghorns(Knowlton 1968).Stimulation of Prey ProductivitySome predation can stimulate productivity and actually lead to largerpopulations of prey (Howard 1974). The idea is that populations regulatedwithin the limits imposed by available food supply are more productive, interms of young produced per adult female, than unregulated populations.Errington (1946) showed that reproductive success was inversely proportionalto population size. White-tailed deer also may be more productive at lowdensities than at high densities (Teer et al. 1965).However, there is little data to establish the importance of predationin stimulating productivity in big game. In a California study, Longhurstet al. (1952) reported slightly more mule deer fawns produced on an area wherecoyotes were uncontrolled than where they were controlled intensively. These263results are unique to the literature; all similar studies have shown nochange or more young produced under a predator control regime.The different findings about effects of coyote predation at stimulatingproductivity in game Iprobably result from three factors. First, the extentof control achieved was not assessed. Second, a distinction should be madebetween gross and net productivity. The predator may well stimulate grossproductivity (number born) by predation on adults but reduce net productivity.^(number surviving through some point in time) by predation on young. Third,carrying capacity invariably was not assessed. Predation would stimulate grossproductivity more if population density is at or above carrying capacity thanbelow.Stimulation of gross productivity in big game by predation would be aselective advantage to the predator which preys most heavily on the young.Predation on the young probably would create an older population than if thepredation were distributed proportionately or equally across all age classes.Since the most productive deer herds are comprised largely of the older ageclasses (Teer et al. 1965) and on good range (Short 1972) (both of which arecaused by predation), coyote predation could be a cause. Obviously, enoughyoung must survive to replenish older age classes or this system could notbe perpetuated.Summary and ConclusionsLoss of habitat and habitat degradation have the greatest effect on gamepopulations. Although habitat improvement may provide the greatest long termbenefit to game populations, habitatalteration is not always feasible. Insuch situations, management should be able to apply feasible alternatives,including predator control, if justifiable.Intensive population control of certain predatory animals can lead to atleast short term population increases of most big game. The primary function of264management is to develop ways to producing a harvestable surplus of animalseach year rather than stockpiling animals. The potential effectiveness ofpredator control in a game management program should be evaluated in relationto this fall surplus. The harvestable surplus need not be increased if ancexcess already exists or is not being properly utilized.LITERATURE CITEDAllee, W. C., H. E. Emerson, 0. Park, T. Park, and K. T. Schmidt. 1949.Principles of animal ecology. W. B. Saunders, Philadelphia, Pa. 837~~.Arrington, 0. N., and A. E. Edwards. 1951. Predator control as a factor inantelope management. Trans. North Am. Wildl. Conf. 16:179-193.Beasom, S. L. 1974a. Intensive short-term predator removal as a game managementtool. Trans. North Am. Wildl. and Nat. Resour. Conf. 39:230-240.. 1974b. Relationships between predator removal and white-tailed deernet productivity. J. Wildl. Manage. 38(4):854-859.Bergerud, A. T. 1971. The population dynamics of Newfoundland caribou.Wildl. Monogr. 25. 55PP.Bruns, E. H. 1970. Winter predation of golden eagles and coyotes on pronghornantelopes. Canadian Field Nat. 84:301-304.Errington, P. L. 1946. Predation and vertebrate populations. Q. Rev. Biol.21:144-177,  221-245.Hailey, T. L. 1973. Causes and degree of mortality of the Trans-Pecosantelope herd. Tex. Parks Wildl. Dep. Prog. Rep. Fed. Aid Proj. W-57-R-20.4PP. (multilith).Hirst, S. M. 1965. Ecological aspects of big game predation. Fauna and Flora. 516:3-15.Hornocker, M. G. 1970. An analysis of mountain lion predation upon mule deerand elk in the Idaho Primitive Area. Wildl. Monogr. 21. 39PP...Howard, W. E. 1974. The biology of predator control. Addison-Wesley Pub.Co., Inc. 48~~.Jones, P. V., Jr. 1949. Antelope management. Tex. Game Fish 7(12):4-5,18-20,  24-25,  28-29.Keith, L. B. 1 9 7 4 . Some features of population dynamics in mammals. Proc.Intl. Cong. of Game.Biol. 11:17-58.Knowles, C. J. 1 9 7 6 . Observations of coyote predation on mule deer andwhite-tailed deer in the Missouri River Breaks, 1975-1976. Pages 117-138in Montana deer studies. Montana Dep. Fish Game. 170pp. (lithograph).Knowlton, F. F. 1 9 6 4 . Aspects of coyote predation in South Texas withspecial reference to white-tailed deer. Ph.D. Thesis. Purdue Univ.,Lafayette, Ind. 198~~.. 1968. Coyote predation as a factor in management of antelope in fencedpastures. Proc. Biennial Antelope States Workshop. 3:64-73.Kruk, H. 1972. The spotted hyena: A study of predation and social behavior.Univ. of Chicago Press, Chicago. 335pp.Leopold A. 1 9 4 3 . Deer irruptions. Wise. Conserv. Bull. 8(8):3-11.Longhurst, W. M., A. S. Leopold, and R. F. Dasmann. 1952. A survey ofCalifornia deer herds, their ranges and management problems. Calif.Dep. of Fish Game Bull. 6. 136~~.Mech, L. D. 1970. The wolf: The ecology and behavior of an endangered species.The Natural History Press, New York. 384~~.Murie, A. 1940. Ecology of the coyote in the Yellowstone. U. S. Nat. ParkServ. Fauna Series 4. 206~~.Riney, T. 1964. The impact of i.ntroduction  of large herbivores on thetropical environment. I.U.C.N. Pub., New Series 4:261-273.Short, H. L. 1972. Ecological framework for deer management. J. Forestry72:200-203.266Teer,.J. G., J. W. Thomas, and E. A. Walker. 1965. Ecology and managementof white-tailed deer in the Llano Basin of Texas. Wildl. Monogr. 15. 62pp.Udy, J. R. 1953. Effects of predator control on antelope populations. tUtah State Dep. Fish Game, Fed. Aid Div. Publ. 5. 48pp.Uzzell,  P. B. 1973. Antelope management in Texas. Tex. Parks Wildl. Dep.LlPP. (multilith).1

PREDATION ON BIG GAME AND ITS MANAGEMENT

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University of Nebraska - Lincoln 
DigitalCommons@University of Nebraska - Lincoln 
Great Plains Wildlife Damage Control Workshop 
Proceedings 
Wildlife Damage Management, Internet Center 
for 
December 1977 
PREDATION ON BIG GAME AND ITS MANAGEMENT 
Samuel L. Beasom 
USDA Forest Service, Rocky Mountain Forest and Range Experiment Station 
Follow this and additional works at: https://digitalcommons.unl.edu/gpwdcwp 
 Part of the Environmental Health and Protection Commons 
Beasom, Samuel L., "PREDATION ON BIG GAME AND ITS MANAGEMENT" (1977). Great Plains Wildlife 
Damage Control Workshop Proceedings. 236. 
https://digitalcommons.unl.edu/gpwdcwp/236 
This Article is brought to you for free and open access by the Wildlife Damage Management, Internet Center for at 
DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Great Plains Wildlife Damage 
Control Workshop Proceedings by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. 
259
PREDATION ON BIG GAME AND ITS MANAGEMENT
Samuel L. Beasom
USDA Forest Service
Rocky Mountain Forest and Range Experiment Station
This paper summarizes information from the literature and personal
observation to present an overview of the effects of predators on big game
populations. Specifically it examines control of prey populations, culling
of inferior prey, and stimulation of prey productivity. Food habit investigations
were omitted because of the impossibility of evaluating the impact of predation
by this approach because of a lack of population data.
Control of Prey Populations
A population is controlled (whether by predation or another means) when
? total deaths equal or exceed births. This control stabilizes populations and
.
keeps them within the limits imposed by their habitats.
Populations of most species of native ungulates have periodically irrupted
and then quickly declined the past 100 years (Keith 1974). This was a result of
decimation of predator populations (Leopold 1943) and/or a major discrepancy
between existing population levels and food supplies and cover (Riney 1964).
Keith (1974) indicated interactions between these two forces make it difficult
to determine which is the most important, but he cited numerous examples that
implicate a lack of predation as the chief cause of the irruptions.
The effect of predation on populations of large ungulates is most
objectively evaluated by predator control, but this method has produced
varying results. Jones (1949), in Texas, was unable to demonstrate a difference
.
in pronghorn antelope (Antilocapra americana) fawn survival between treated
L'
and untreated areas. He attributed the lack of treatment effect to ineffective
predator control. Later investigations in the same general area (Bailey 1973,
Uzzell  1973) revealed a marked increase in pronghorn net productivity where
260
coyotes (Canis latrans) were intensively removed. Similar investigations in
Arizona (Arrington and Edwards 1951), Utah (Udy 1953),  and Wyoming (Yoakum
1968 as interpreted by Knowlton 1968) attributed increases in net reproduction
and in total pronghorn populations to intensive coyote control.
Predator control varied in effectiveness of alleviating depredations on
young deer (Odocoileus spp.) and caribou (Rangifer tarandus). Longhurst
et al. (1952) found no evidence that fawn survival increased where coyotes
were trapped heavily compared to the control area. Beasom (1974a, 1974b)
reported an approximately 300% and 400% increase in white-tailed deer
(Odocoileus virginianus) net fawn production during a 2-year period on a study
area in Texas. The deer population density increased by about 28% on the
treated area and declined by about 12% on the untreated area. Daniel (1975)
found an inverse correlation between coyote abundance and percent fawn
production. Although he noted a higher net fawn production on the treated
areas, deer population density did not increase above that on the untreated L
areas each year. It is not known where the density level of the deer herds
in these two Texas studies were located in relation to the carrying capacities
of the respective habitats, so it is difficult to relate the reported density
change differences to this important factor. In Newfoundland, Bergerud (1971)
found that intensive lynx (Lynx canadensis) control on one study area was
followed by an approximately 100% increase in caribou calf survival by fall
compared to an untreated area. Data on the change in caribou population
density between the areas were not available.
In general, predators have the potential to curtail most ungulate populations
when proper circumstances prevail. Errington (1946) suggested that canid
.q
predators can control ungulate populations by preying mostly on the young,
because of the relatively low productivity of large herbivores. Allee  et al.
(1949) generalied that, to control any population, the net productivity input
261
must be curtailed. The level of predation on big game populations in any
area is doubtless a result of the density of the predator population, the
behavioral reactions of the game animals, the food predilections of the
predator, and the abundance and quality of alternative foods for the predator.
Although predators apparently influence ungulate populations, unlimited
population growth in the absence of predation has not been demonstrated. When
the population reaches or surpasses the habitat carrying capacity, further
growth is slowed by other mortality factors. Controlling predatory animals
to increase an already high ungulate population would be fruitless.
Culling of Inferior Prey
A common view is that predators serve to maintain healthy prey populations
r
by culling the sick and weak or otherwise inferior individuals. Mech (1970)
included young prey animals as biologically inferior. This discussion, however,
does not consider all young animals to be biologically inferior.
Selection of strong or weak prey depends on the hunting habits of the
predator. For example, stalking predators, such as cats, usually kill by
surprise attack, which is relatively independent of the condition of the prey
(Hornocker 1970, Hirst 1965). Predators which chase, and elicit a flight response
from their prey, such as wolves (Canis spp.) and spotted hyenas (Crocuta crocuta),
tend to take a disproportionate number of poor condition animals as prey (Mech
1970, Kruk 1972). The coyote cannot be assigned either category because of
its diverse hunting patterns and feeding habits. It probably hunts big game by
the chase-kill method (Murie 1940). This hunting behavior killed a dispropor-
tionate number of old, injured or young deer. Although reports of coyotes
killing full grown, healthy wild ungulates are common, they are probably the
exception. Size alone, since they rarely hunt in groups larger than two (Fox
1971), would preclude a coyote's preying on adults of most native North Amc~ricnn
ungulates. Coyotes feed mostly on small mammals and birds but will sometimes
kill deer and other large animals, usually young or weakened individuals
(Mech 1970).
Coyotes reportedly prey heavily on adult deer in winter when prey body
condition presumably declines (Murie 1940, Knowlton 1964, Knowles 1976).
Coyote predation at this time may be disproportionate toward adult males
because they are physiologically exhausted from the rut as well as the poorer
food conditions (Knowlton 1964, Beasom unpublished data). Hood (personal
communication), while conducting a telemetry study to investigate deer
movements in South Texas, noted a similar disproportionate coyote predation
on collared adult bucks from late winter to early spring.
Coyote predation on pronghorns indicates that this predator does not take
disproportionately more unfit ungulates (Bruns 1970). Murie (1940) presented
several accounts of pronghorns easily outdistancing coyotes on unfenced ranges
within Yellowstone National Park. However, coyotes apparently have learned I
to use fences (especially net wire) to capture even healthy pronghorns
(Knowlton 1968).
Stimulation of Prey Productivity
Some predation can stimulate productivity and actually lead to larger
populations of prey (Howard 1974). The idea is that populations regulated
within the limits imposed by available food supply are more productive, in
terms of young produced per adult female, than unregulated populations.
Errington (1946) showed that reproductive success was inversely proportional
to population size. White-tailed deer also may be more productive at low
densities than at high densities (Teer et al. 1965).
However, there is little data to establish the importance of predation
in stimulating productivity in big game. In a California study, Longhurst
et al. (1952) reported slightly more mule deer fawns produced on an area where
coyotes were uncontrolled than where they were controlled intensively. These
263
results are unique to the literature; all similar studies have shown no
change or more young produced under a predator control regime.
The different findings about effects of coyote predation at stimulating
productivity in game Iprobably result from three factors. First, the extent
of control achieved was not assessed. Second, a distinction should be made
between gross and net productivity. The predator may well stimulate gross
productivity (number born) by predation on adults but reduce net productivity.^
(number surviving through some point in time) by predation on young. Third,
carrying capacity invariably was not assessed. Predation would stimulate gross
productivity more if population density is at or above carrying capacity than
below.
Stimulation of gross productivity in big game by predation would be a
selective advantage to the predator which preys most heavily on the young.
Predation on the young probably would create an older population than if the
predation were distributed proportionately or equally across all age classes.
Since the most productive deer herds are comprised largely of the older age
classes (Teer et al. 1965) and on good range (Short 1972) (both of which are
caused by predation), coyote predation could be a cause. Obviously, enough
young must survive to replenish older age classes or this system could not
be perpetuated.
Summary and Conclusions
Loss of habitat and habitat degradation have the greatest effect on game
populations. Although habitat improvement may provide the greatest long term
benefit to game populations, habitatalteration is not always feasible. In
such situations, management should be able to apply feasible alternatives,
including predator control, if justifiable.
Intensive population control of certain predatory animals can lead to at
least short term population increases of most big game. The primary function of
264
management is to develop ways to producing a harvestable surplus of animals
each year rather than stockpiling animals. The potential effectiveness of
predator control in a game management program should be evaluated in relation
to this fall surplus. The harvestable surplus need not be increased if an
c
excess already exists or is not being properly utilized.
LITERATURE CITED
Allee, W. C., H. E. Emerson, 0. Park, T. Park, and K. T. Schmidt. 1949.
Principles of animal ecology. W. B. Saunders, Philadelphia, Pa. 837~~.
Arrington, 0. N., and A. E. Edwards. 1951. Predator control as a factor in
antelope management. Trans. North Am. Wildl. Conf. 16:179-193.
Beasom, S. L. 1974a. Intensive short-term predator removal as a game management
tool. Trans. North Am. Wildl. and Nat. Resour. Conf. 39:230-240.
. 1974b. Relationships between predator removal and white-tailed deer
net productivity. J. Wildl. Manage. 38(4):854-859.
Bergerud, A. T. 1971. The population dynamics of Newfoundland caribou.
Wildl. Monogr. 25. 55PP.
Bruns, E. H. 1970. Winter predation of golden eagles and coyotes on pronghorn
antelopes. Canadian Field Nat. 84:301-304.
Errington, P. L. 1946. Predation and vertebrate populations. Q. Rev. Biol.
21:144-177,  221-245.
Hailey, T. L. 1973. Causes and degree of mortality of the Trans-Pecos
antelope herd. Tex. Parks Wildl. Dep. Prog. Rep. Fed. Aid Proj. W-57-R-20.
4PP. (multilith).
Hirst, S. M. 1965. Ecological aspects of big game predation. Fauna and Flora. 5
16:3-15.
Hornocker, M. G. 1970. An analysis of mountain lion predation upon mule deer
and elk in the Idaho Primitive Area. Wildl. Monogr. 21. 39PP.
..
Howard, W. E. 1974. The biology of predator control. Addison-Wesley Pub.
Co., Inc. 48~~.
Jones, P. V., Jr. 1949. Antelope management. Tex. Game Fish 7(12):4-5,
18-20,  24-25,  28-29.
Keith, L. B. 1 9 7 4 . Some features of population dynamics in mammals. Proc.
Intl. Cong. of Game.Biol. 11:17-58.
Knowles, C. J. 1 9 7 6 . Observations of coyote predation on mule deer and
white-tailed deer in the Missouri River Breaks, 1975-1976. Pages 117-138
in Montana deer studies. Montana Dep. Fish Game. 170pp. (lithograph).
Knowlton, F. F. 1 9 6 4 . Aspects of coyote predation in South Texas with
special reference to white-tailed deer. Ph.D. Thesis. Purdue Univ.,
Lafayette, Ind. 198~~.
. 1968. Coyote predation as a factor in management of antelope in fenced
pastures. Proc. Biennial Antelope States Workshop. 3:64-73.
Kruk, H. 1972. The spotted hyena: A study of predation and social behavior.
Univ. of Chicago Press, Chicago. 335pp.
Leopold A. 1 9 4 3 . Deer irruptions. Wise. Conserv. Bull. 8(8):3-11.
Longhurst, W. M., A. S. Leopold, and R. F. Dasmann. 1952. A survey of
California deer herds, their ranges and management problems. Calif.
Dep. of Fish Game Bull. 6. 136~~.
Mech, L. D. 1970. The wolf: The ecology and behavior of an endangered species.
The Natural History Press, New York. 384~~.
Murie, A. 1940. Ecology of the coyote in the Yellowstone. U. S. Nat. Park
Serv. Fauna Series 4. 206~~.
Riney, T. 1964. The impact of i.ntroduction  of large herbivores on the
tropical environment. I.U.C.N. Pub., New Series 4:261-273.
Short, H. L. 1972. Ecological framework for deer management. J. Forestry
72:200-203.
266
Teer,.J. G., J. W. Thomas, and E. A. Walker. 1965. Ecology and management
of white-tailed deer in the Llano Basin of Texas. Wildl. Monogr. 15. 62pp.
Udy, J. R. 1953. Effects of predator control on antelope populations. t
Utah State Dep. Fish Game, Fed. Aid Div. Publ. 5. 48pp.
Uzzell,  P. B. 1973. Antelope management in Texas. Tex. Parks Wildl. Dep.
LlPP. (multilith).
1


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