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/an/, I am pressed for time and all the local libraries are closed. Do any of you have a complete copy of the following article, or JSTOR access you'd be willing to share?

Mountain Sheep and Coyotes: Patterns of Predator Evasion in a Mountain Ungulate
Vernon C. Bleich
Journal of Mammalogy, Vol. 80, No. 1 (Feb., 1999), pp. 283-289
doi:10.2307/1383228
>> Anonymous
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Long-shot is looooooooong. I'd be surprised if this turns out the way you want it to. Try public veterinary student forums.
>> Anonymous
OP, is this the result of procrastination?
>> Anonymous
Hey OP, I got a copy. Where can I send it? Do you want just the text, or the images as well?
>> Don't worry. Here's the text and image. Anonymous
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Abstract (Document Summary)
I used retrospective analyses to investigate relationships among terrain type, reactions of prey, and age and sex of prey and outcomes of encounters between mountain sheep (Ovis canadensis) and coyotes (Canis latrans).

Copyright American Society of Mammalogists Feb 1999[Headnote]
I used retrospective analyses to investigate relationships among terrain type, reactions of prey, and age and sex of prey and outcomes of encounters between mountain sheep (Ovis canadensis) and coyotes (Canis latrans). Adult females fled from coyotes more often when in open terrain and young always fled regardless of type of terrain. Young were less likely to survive encounters with coyotes than were adults, and no adult females were killed by coyotes. When types of terrain were pooled, females were more apt to flee than were males, but no difference existed in proportion of females and young that fled. When terrain type and sex of adults were pooled, no difference in survival existed between adults that fled and those that did not. When coyotes were encountered in open terrain, female mountain sheep fled more frequently than did female mule deer (Odocoileus hemionus). Ungulates less dependent on terrain for predator evasion than are mountain sheep may employ a more plastic strategy than artiodactyls inhabiting precipitous terrain.
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Predation may elicit overt behavioral changes by ungulates, such as modification of vigilance behavior and foraging efficiency (Berger, 1978a; Berger et al., 1983; Frid, 1997; Molvar and Bowyer, 1994), abandonment of formerly used habitats (Wehausen, 1996), or differential use of habitats by parturient and nonparturient females (Kohlmann et al., 1996). Among prey, reactions to predators may vary from retreat to overt aggression (Berger, 1979; Bowyer, 1987) and, within species, individuals may respond along a continuum from charging the predator to rapid retreat (Berger, 1991). Moreover, body condition of adults or offspring can influence the reaction of maternal females when confronted by a predator (Smith, 1987).

Antipredator mechanisms (Kruuk, 1964) can be either direct (flight or aggression) or indirect (vigilance, social behavior, or use of specific habitats). Both have implications for survival of individuals, although direct responses are more likely to be reported in the literature (Berger, 1991). Nevertheless, observations of interactions between large mammalian predators and their prey are difficult to obtain and frequently are anecdotal because such encounters are observed infrequently and have not been a primary focus of researchers.
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I investigated the ways that habitat may have affected direct antipredator strategies of mountain sheep, the manner in which those strategies influenced outcomes of their encounters with coyotes, and compared responses of female mountain sheep and female mule deer (Odocoileus hemionus) to coyotes encountered in non-mountainous terrain. I hypothesized that 1) responses of mountain sheep to coyotes would differ as a function of habitat type, and the sex and age of the sheep involved, 2) outcomes of encounters with coyotes would be influenced by the direct response of the prey to the predator, and 3) responses of mountain sheep to coyotes would differ from responses of mule deer.
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MATERIALS AND METHODS

I relied on observations (n = 42) of direct interactions between coyotes and mountain sheep compiled by Berger (1991:67, table 1). I classified mountain sheep as males or females (>=1-year-old), and young (<1-year-old). I used the terms open terrain and escape terrain to describe flat or rolling non-rocky areas and rocky or steep mountainous areas, respectively. I based my analyses on the habitat in which interactions occurred and, with one exception, on the age and sex of mountain sheep reported by Berger (1991); I treated one yearling male as an adult because it occupied habitat typically used by adult males during the encounter. Group sizes of mountain sheep or coyotes were not available and, thus, could not be considered in this paper.

I used data from Bowyer (1987:521, fig. 3) on responses of mule deer to elucidate potential differences in responses of these ungulates and mountain sheep to the threat of predation by coyotes. Mule deer are well-adapted to flee and make use of vegetation to hide from predators. In contrast, mountain sheep largely are dependent on terrain features to evade predators. For interspecific analyses, I considered only female sheep, because available data were nearly all for female deer (R. T. Bowyer, pers. comm.); hence, results would not be confounded by intersexual differences in strategies of predator evasion.
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The number of observations of coyotes interacting with mountain sheep males (n = 9), females (n = 23), and young (n = 10) reported by Berger (1991) and the observations of mule deer and coyotes (n = 16) reported by Bowyer (1987) indicated that univariate statistical tests were most appropriate. I used categorical analyses (G-test, Fisher exact test-Zar, 1984) to test for associations (P <= 0.10) between habitat type and age and sex of mountain sheep and responses to coyotes, as well as the outcomes of such encounters, and to test for effects of behavior of mountain sheep on outcomes of encounters. Similarly, I used a G-test to compare responses of female mountain sheep and mule deer to coyotes encountered in open terrain. I tested for differences in responses of mule deer to coyotes with a binomial test (Zar, 1984). Where appropriate for contingency tables, I applied the sequential Bonferroni technique to decrease the probability of Type I error (Rice, 1989). I estimated statistical power (1 - 3) according to Cohen (1969:209-265). Where sample size was small, or beta > 0.70, I presented data without statistical analyses, thereby providing the reader an opportunity to consider the potential importance of the observations. When I failed to reject a null hypothesis, I included the effect size (Delta) with the test statistics (Steidl et al., 1997).
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RESULTS

Effect of terrain on behavior and survival.-When mountain sheep (all categories of age and sex combined) encountered coyotes, they fled more often (G = 6.693, d.f. = 1, P = 0.010) when in open (20 of 25) than when in escape terrain (7 of 17). When young were excluded from this analysis, adults still fled more often (P < 0.001) when in open terrain (17 of 22) than when in escape terrain (0 of 10). Likewise, adult females fled more often (P = 0.001) when in open terrain (15 of 16) than when in escape terrain (0 of 7). Sample size was inadequate to allow a meaningful comparison of males that fled when in open terrain (2 of 6) and escape terrain (0 of 3). All 10 young that encountered coyotes fled, regardless of terrain type.

No adult females were killed in either open (n = 7) or escape (n = 16) terrain. Sample sizes were inadequate to allow meaningful comparisons of the proportions of males killed in either open (2 of 6) or escape (0 of 3) terrain, or of young killed in open (2 of 3) or escape (2 of 7) terrain. Mortality of young in open terrain (67%) was, however, 2.4 times that in steep terrain (28%).
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Effects of age and sex on behavior and survival.-When type of terrain was pooled, there was a significant difference (G = 15.493, df. = 2, P < 0.001) in the proportion of individual young (10 of 10), females (15 of 23), and males (2 of 9) that fled when they encountered coyotes. Pairwise comparisons indicated that young (P = 0.001) and females (P = 0.049) fled more often than males, and that young fled more often than females (P = 0.071).

A significant difference (G = 11.455, df = 2, P = 0.003) also occurred among the proportion of young (3 of 10), females (0 of 23), and males (2 of 9) that died during encounters with coyotes. Pairwise comparisons indicated that proportionately fewer young survived encounters than did females, or adults in general (P = 0.005 and P = 0.021, respectively), and that proportionately more males than females died (P = 0.073). Sample sizes were inadequate to allow a meaningful comparison between young and males.

Effect of behavior on survival.-When sexes of adults were combined and type of terrain was pooled, none of the 15 adults that did not flee were killed, and only 2 of the 12 that fled died (P = 0.188, A - 0.06). All young that encountered coyotes fled, and they were killed more often (4 of 10) than were adults (2 of 27) that fled (P = 0.014).
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Responses of mountain sheep and mule deer to coyotes.-No difference existed in the proportion of mule deer that fled (9 of 16) or stayed (7 of 16) when they encountered coyotes in open terrain (P > 0.50). When I compared responses of female mountain sheep and female mule deer (Fig. 1), mountain sheep fled more often (G = 6.578, df = 1, P = 0.010) than mule deer. In contrast, deer fled more often (G = 8.859, df = 1, P < 0.003) than sheep when the latter encountered coyotes in escape terrain. Proportions of deer (1 of 16) and mountain sheep (0 of 23) killed by coyotes essentially were identical.

DISCUSSION

Consistent with hypothesis 1, responses to coyotes varied with terrain, sex, and age of mountain sheep. Adults fled more frequently when encounters occurred in open terrain that facilitated pursuit by these coursing predators. Young mountain sheep always fled when they encountered coyotes and did so more often than adults. Moreover, young were killed proportionately more often than adults. Young are substantially smaller than adults (Bleich et al., 1997) and likely are more vulnerable to predation by medium sized predators (Berger, 1991; Curio, 1976).
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Regardless of the response to coyotes, higher proportions of young and male mountain sheep were killed than were females. In the eastern Mojave Desert, males occupied areas with more coyotes in which they were especially vulnerable to predators during periods of sexual segregation (Bleich et al., 1997). Because of their small size and inexperience, juveniles may be less adept than adults at escaping from coyotes. For example, Shank (1977) reported that two fleeing young were killed when they were herded away from escape terrain by coyotes but adults apparently learn to not be herded.

No difference existed in the proportion of adults that were killed, whether or not they attempted to escape, leading me to reject hypothesis 2. Nevertheless, young fled in all instances, and were more apt to be killed by coyotes than were adults. The lack of a relationship between the behavior of adults and being killed by coyotes was surprising; among domestic sheep and mule deer, those that fled were attacked or pursued more often than those that did not (Bowyer, 1987; Connolly et al., 1976).
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Models of habitat suitability developed for mountain sheep (Bleich and Holl, 1982; Bleich et al., 1992; Cunningham, 1989; Hansen, 1980) have emphasized the importance of escape terrain as a habitat feature, and Holl (1982) demonstrated a strong relationship between density of females and available escape terrain. Data summarized herein suggest that the terrain in which encounters initially occurred did not affect the probability of adult females being killed by coyotes. Young, however, may be especially vulnerable to predation, and females may enhance their fitness by selecting habitats in which offspring are less apt to encounter and more likely to evade predators (Bleich et al., 1997).
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Mountain ungulates are especially dependent on steep and rugged terrain to avoid predation (Schaller, 1977). The strategy of reducing risk may differ for ungulates adapted to less precipitous terrain where the ability to escape predators is more limited, but hiding (Geist, 1981:171), increasing group size (sensu Hamilton, 1971), or confronting predators (Griffith, 1988) are successful strategies. Mule deer in open terrain demonstrated no propensity to flee (Bowyer, 1987:521, fig. 3), but mountain sheep almost always fled when they encountered coyotes away from escape terrain. In contrast, female mountain sheep never fled when they encountered coyotes in escape terrain-a situation in which they are adept at evading predation. Rachlow and Bowyer (1991) reported low neonatal mortality among Dall's sheep (Ovis dalli), even in a predator-rich environment. Likewise, mortality due to predation by coyotes was not detected among female mountain sheep occupying steep terrain in a desert environment (Bleich et al., 1997). Female mountain sheep employ a general strategy of flight when in open terrain and stand their ground or act aggressively when they encounter coyotes in escape terrain. In contrast, mule deer, and perhaps other cervids occupying less precipitous terrain, employ a more plastic predator evasion strategy that may be affected by the proximity of vegetative cover. As with mountain sheep, female mule deer exhibit active defense of young under certain circumstances (Bowyer, 1987; Griffith, 1988).
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There is a strong effect of habitat, age, and sex on behavioral responses of mountain sheep to predators and the subsequent outcome of such encounters. Additionally, ungulates well-adapted to mountainous terrain respond differently to the threat of predation than those occupying less precipitous terrain. Several aspects of risk, however, are the subject of much uncertainty (Lima and Dill, 1990), and rigorous analyses of antipredator behaviors frequently are confounded by differences among species (Kohlmann et al., 1996).

An integrative approach (Frid, 1997) that simultaneously considers effects of visibility (Risenhoover and Bailey, 1985), availability of escape terrain (Bleich et al., 1997), group size of predators and prey (Bowyer, 1987; Elgar, 1989; Fitzgibbon and Lazarus, 1995), and effects of age and sex will likely be productive in understanding differences in predator-evasion strategies of ungulates occupying mountainous and nonmountainous terrain. Moreover, stalking and coursing predators may elicit differing responses by prey. As a consequence, hunting strategies of different predators also should be considered by researchers studying responses of large mammals to the threat of predation.
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ACKNOWLEDGMENTS

I thank J. Berger, R. T Bowyer, H. B. Ernest, M. M. Grigione, X. Manteca, M. W. Oehler, Sr., B. M. Pierce, J. L. Rechel, E. S. Rubin, R. Schaefer, S. G. Torres, and two anonymous reviewers for helpful comments. Preparation of this paper was supported by a University of California Davis-California Resources Agency Fellowship. This is a contribution from the California Department of Fish and Game Mountain Sheep Conservation Program and is Professional Paper 001 from the Eastern Sierra Center for Applied Population Ecology. I dedicate this paper to the memory of Dr. W. C. Gasaway, from whom I learned much about the ecology of predation and the political realities of wildlife conservation.
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[Reference]
LITERATURE CITED

[Reference]
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[Reference]
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. 1979. "Predator harassment" as a defensive strategy in ungulates. The American Midland Naturalist, 102:197-199.
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[Reference]
BERGER, J., D. DANEKE, J. JOHNSON, AND S. H. BERWICK. 1983. Pronghorn foraging economy and predator avoidance in a desert ecosystem: implications for the conservation of large mammalian herbivores. Biological Conservation, 25:193-208. BLEICH, V. C. 1996. Interactions between coyotes (Canis latrans) and mountain sheep (Ovis canadensis). The Southwestern Naturalist, 41:81-82. BLEICH, V. C., AND S. A. HOLL. 1982. Management of chaparral habitat for mule deer and mountain sheep in southern California. Pp. 247-254, in Proceedings of the Symposium on the Dynamics and Management of Mediterranean-type Ecosystems (C. E. Conrad and W. C. Oechel, tech. coords.). United States Forest Service General Technical Report, PSW58:1637.

[Reference]
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[Reference]
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BLEICH, V. C., J. D. WEHAUSEN, R. R. RAMEY II, AND J. L. RECHEL. 1996. Metapopulation theory and mountain sheep: implications for conservation. Pp. 353-373, in Metapopulations and wildlife conservation (D. R. McCullough, ed). Island Press, Covelo, California.
BOWYER, R. T. 1987. Coyote group size relative to predation on mule deer. Mammalia, 51:515-526. COHEN, J. 1969. Statistical power analysis for the behavioral sciences. Academic Press, New York. CONNOLLY, G. E., R. M. TIMM, W. E. HOWARD, AND W. M LONGHURST. 1976. Sheep killing behavior of captive coyotes. The Journal of Wildlife Management, 40:400-407.
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[Reference]
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CuRio, E. 1976. The ethology of predation. Zoophysiology and Ecology, Springer-Verlag, Berlin, Germany, 7:1-250.
DEFoRGE, J. R., AND J. E. ScoTT. 1982. Ecological investigations into high lamb mortality. Desert Bighorn Council Transactions, 26:65-76. ELGAR, M. A. 1989. Predator vigilance and group size in mammals and birds: a critical review of the empirical evidence. Biological Review, 64:13-33.

[Reference]
FESTA-BIANCHET, M. 1988. Seasonal range selection in bighorn sheep: conflicts between forage quality, forage quantity, and predator avoidance. Oecologia, 75: 500-506.
FITZGIBBON, C. D., AND J. LAZARUS. 1995. Anti-predator behaviour of Serengeti ungulates: individual differences and population consequences. Pp. 274296, in Serengeti II: research, management and conservation of an ecosystem (A. R. E. Sinclair and P Arcese, eds.). The University of Chicago Press, Chicago, Illinois.
>> Anonymous
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I thought I was too slow for a combo breaker.

Well, here it is.
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[Reference]
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. 1971. Geometry for the selfish herd. Journal of Theoretical Biology, 31:295-311. HANSEN, C. G. 1980. Habitat evaluation. Pp. 320-335, in The desert bighorn: its life history, ecology and management (G. Monson and L. Sumner, eds.). University of Arizona Press, Tucson. HASS, C. C. 1989. Bighorn lamb mortality: predation, inbreeding, and population effects. Canadian Journal of Zoology, 67:699-705.
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[Reference]
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male bighorn sheep. Journal of Mammalogy, 50:128. KELLY, W. E. 1980. Predator relationships. Pp. 186196, in The desert bighorn: its life history, ecology and management (G. Monson and L. Sumner, eds.). University of Arizona Press, Tucson. KOHLMANN, S. G., D. M. MULLER, AND P U. ALKON. 1996. Antipredator constraints on lactating Nubian ibexes. Journal of Mammalogy, 77:1122-1131. KRUUK, H. 1964. Predators and anti-predator behaviour of the black-headed gull (Larus ridibundus L.). Behaviour, 11 supplement:l-129.
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RICE, W. R. 1989. Analyzing tables of statistical tests.
Evolution, 43:223-225.
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[Reference]
sheep and goats of the Himalaya. The University of Chicago Press, Chicago, Illinois. SHANK, C. C. 1977. Cooperative defense by bighorn
sheep. Journal of Mammalogy, 58:243-244. SMITH, W. P. 1987. Maternal defense in Columbian white-tailed deer: when is it worth it? The American Naturalist, 130:310-316.
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[Reference]
WEHAUSEN, J. D. 1996. Effects of mountain lion predation on bighorn sheep in the Sierra Nevada and Granite Mountains of California. Wildlife Society Bulletin, 24:471-479.
WEHAUSEN, J. D., V. C. BLEICH, B. BLONG, AND T. L. Russi. 1987. Recruitment dynamics in a southern California mountain sheep population. The Journal of Wildlife Management, 51:86-98. WooLF, A., AND T. O'SHEA. 1968. Two bighorn sheep
coyote encounters. Journal of Mammalogy, 49:770. ZAR, J. H. 1984. Biostatistical analysis. Second ed. Prentice-Hall, Inc., Englewood Cliffs, New Jersey. Submitted 25 September 1997. Accepted 16 April 1998.
Associate Editor was C. Renn Tumlison.

[Author Affiliation]
VERNON C. BLEICH

[Author Affiliation]
California Department of Fish and Game, 407 W. Line St, Bishop, CA 93514 and Institute of Arctic Biology and Department of Biology and Wildlife, University of Alaska Fairbanks, Fairbanks, AK 99775
>> Anonymous
>>105700
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>> Anonymous
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>> Anonymous
>>105744

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>> Anonymous
>>105667

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>> Anonymous
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>>105812
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>>105818

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