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Deer-Predator Relationships: a Review of Recent North American Studies with Emphasis on Mule and Black-Tailed Deer

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Deer-Predator Relationships: a Review of Recent North American Studies with Emphasis on Mule and Black-Tailed Deer Deer-Predator Relationships: A Review of Recent North American Studies with Emphasis on Mule and Black-Tailed Deer Warren B. Ballard; Daryl Lutz; Thomas W. Keegan; Len H. Carpenter; James C. deVos, Jr. Wildlife Society Bulletin, Vol. 29, No. 1. (Spring, 2001), pp. 99-115. Stable URL: http://links.jstor.org/sici?sici=0091-7648%28200121%2929%3A1%3C99%3ADRAROR%3E2.0.CO%3B2-I Wildlife Society Bulletin is currently published by Alliance Communications Group. Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/about/terms.html. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/journals/acg.html. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is an independent not-for-profit organization dedicated to and preserving a digital archive of scholarly journals. For more information regarding JSTOR, please contact [email protected]. http://www.jstor.org Wed Apr 18 12:38:59 2007 Predation B DEER-PREDATOR RELATIONSHIPS 99 Deer-predator relationships: a review of recent North American studies with emphasis on mule and black-tailed deer lirrren B. Ballarc(. Dnr7.l Lutz, Thomas ll: Keega11, Lerr H. Carpenter. ccnd Jarnos C. de Vo.s,Jr. Abstract In recent years mule (Odocoileus hemionus) and black-tailed (0.h, columbianus) deer appear to have declined in many areas of the western United States and Canada, causing concern for population welfare and continued uses of the deer resource. Causes of the decline have not been identified, but predation by coyotes (Canis latrans), mountain lions (Puma concolor), and wolves (Canis lupus) has been proposed as one of many factors. We reviewed results of published studies conducted since the mid-1 970s concerning predator-deer relationships to determine whether predation could be a factor in the apparent deer population declines and whether there was evidence that predator control could be a viable management tool to restore deer populations. We reviewed 17 published studies concerning mule deer. We found only 4 pub- lished studies of the effects of predation on black-tailed deer. A larger database exist- ed for white-tailed deer (0. virginianus), with 19 studies examining effects of preda- tion on white-tailed deer. Study results were confounded by numerous factors. A deer population's relation- ship to habitat carrying capacity was crucial to the impacts of predation. Deer popu- lations at or near carrying capacity did not respond to predator removal experiments. When deer populations appeared limited by predation and such populations were well below forage carrying capacity, deer mortality was reduced significantly when predator populations were reduced. Only one study, however, demonstrated that deer population increases resulted in greater harvests, although considerable data indicat- ed that wolf control resulted in greater harvests of moose (Alces alces) and caribou (Rangifer tarandus). The most convincing evidence for deer population increases occurred when small enclosures (2-39 km2) were used. Our review suggests that predation by coyotes, mountain lions, or wolves may be a significant mortality factor in some areas under certain conditions. Relation to habi- tat carrying capacity, weather, human use patterns, number and type of predator species, and habitat alterations all affect predator-prey relationships. Only through intensive radiotelemetry and manipulative studies can predation be rdentified as a major limiting factor. When it is identified, deer managers face crucial decisions. Address for LVarren B. Ballard: Department of Range, Wilcllife, and Fisheries Management, Texas Tech University; Box 12125, Lubbock, TX 79409. USA; e-mail: [email protected]. Address for Daryl Lutz: Wyoming Game and Fish Department, 3030 Energy Lane, Suite 100, Caper, LVY 82604. USA. Address ior Thomas LV. Keegan: Oregon Department oi Fish and Ll'ildlife, P. 0. Box 59, Port- land, OR 97207, USA. Address for Len H. Carpenter: Wildlife Management Institute, 4015 Cheney Drive, Fort Collins, CO 80526, USA. Address for James C. deVos, Ir.: Research Branch, Arizona Game and Fish Department, 2221 West Greenway Road, Phoenix, AZ 85023, USA. Present atldress ior Keegan: LVashington Department of Fish and L21ildlife. 600 Capitol North, Olympia, LVA 98.501, USA. Wildlife Society Bulletin 2001, 29(1):99-115 Peer edited 100 Wtrnlife Society Bulletin 2001,29(1):99-1 1 5 Reductions in predator densities have occurred only on relatively small study areas (2-1 80 km*) where predators were identified as a major limiting factor and deer pop- ulations were \yell below forage carrying capacity (an important criterion). Thus a problem of scale, methods used to kill predators, benefit:cost ratios, results to hunters, and public acceptance are primary considerations. Methods of predator control avail- able to deer managers have been severely restricted and current methods may not be feasible over large areas when and if predation becon~esa problem. Public accept- ance of predator reduction programs is essential for predator-prey management, but may not be achievable given current public attitudes toward predators. We identified several recommendations and research needs based on our review of the literature given current social and political limitations. Key words black-tailed deer, carrying capacity, coyote, mountain lion, mule deer, population man- agement, predation management, predators, wolf Wildlife management agencies in the western dation on ungulates, indicated that a selective IJnited States and Canada are concerned about an review of the literature could reinforce almost any apparent decline of mule deer (Ollocoileus view 011 the role of predation. He reviewed 3 1 stud- henzionus) and black-tailed deer (0, h. colum- ies that indicated predation was a limiting or regu- bium~~)populations over large portions of western lating influence and 27 studies indicating that pre- North America (Western Association of Fish and dation was not limiting (Connolly 19-8). However, Wildlife Agencies, mule Deer Committee. 1998 degree of docun~entation varied widely among unpublished report). Western deer populations studies. He concluded that predators acting in con- have been described as very volatile, with major cert with weather, disease, and habitat changes cycles of high and low populations (Denney 1976). could have important effects on prey numbers. Herds apparently began increasing in the 1920s. Since Connollj-'s review, scientists ha\-e continued peaked in the late 1940s to early 1960s, declined to debate whether predation is a significant regu- during the 1960s to mid-l970s, increased during lating factor on ungulate populations (Messier the 1980s. and then declined during the 1990s 1991, Sinclair 1991. Skogland 1991, Boutin 1992, (Denney 1976. Hurley and Unsworth 1998). Solne Van Ballenberghe and Ballartl 1994). Because of investigators indicated that deer populations in increased interest in relationships between preda- some areas have been declining since the 1960s tion and deer populations. we reviewed available (Workman and Low 1976. Schneegas and Bumstead literature concerning deer-predator relationships 1977, Bleich and Taylor 1998). and sought to draw conclusions regarding effects of Numerous factors could be responsible for deer predation on mule and black-tailed deer popula- declines. iilcluding habitat loss or change, severe tions and, based upon our assessment. make appro- weather (e.g., drought, deep snow), starvation, priate recommendations for additional research changes in age and sex structure, disease, predation. and management. competition with livestock and wildlife species such as elk (Cervus eluphus), hunting, and interactions of these factors (Wallmo 1981, Halls 1984, Whittaker Methods and Lindzey 1999). Recently, some members of the We focused our review on studies conducted public (e.g.. Barsness 1998) and some biologists since the mid-1970s and, where applicable to deer. (Gasaway et al. 1992) indicated that predation maj- included some studies summarized by Connolly be largely responsible for declines or lack of ungu- (1 9'8,198 1). We used selected abstracting services late population recovery and that predator control and searched for literature pertaining to deer-pred- may be necessary to restore sollle populations to ator relationships. We searched all major biological greater levels. However, empirical evidence exists and wildlife journals and reliewed literature cita- only for moose (Alces nlces). caribou (Rangfer tions within articles for additional references We tamndus), and one black-tailed deer population, and purposefully excluded predator diet studies this hypothesis has not been tested for mule deer. because these do not allow assessment of effects of Connolly (1978), in his review of effects of pre- predation on prej populations. Deer-predator relationships Ballard et al. 101 Several authors indicated that confusion exists in one cause of mortality or introduction of a new the predator-prey literature because biologists cause may or may not increase
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