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Habitat Use of Sympatric Prey Suggests Divergent Anti-Predator Responses

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Habitat Use of Sympatric Prey Suggests Divergent Anti-Predator Responses Oecologia https://doi.org/10.1007/s00442-018-4323-z COMMUNITY ECOLOGY – ORIGINAL RESEARCH Habitat use of sympatric prey suggests divergent anti‑predator responses to recolonizing gray wolves Justin A. Dellinger1,2 · Carolyn R. Shores1 · Apryle Craig1 · Michael R. Heithaus3 · William J. Ripple4 · Aaron J. Wirsing1 Received: 15 January 2018 / Accepted: 8 December 2018 © Springer-Verlag GmbH Germany, part of Springer Nature 2018 Abstract The non-consumptive efects of predators on prey are now widely recognized, but the need remains for studies identifying the factors that determine how particular prey species respond behaviorally when threatened with predation. We took advantage of ongoing gray wolf (Canis lupus) recolonization in eastern Washington, USA, to contrast habitat use of two sympatric prey species—mule (Odocoileus hemionus) and white-tailed (O. virginianus) deer—at sites with and without established wolf packs. Under the hypothesis that the nature and scale of responses by these ungulates to wolf predation risk depend on their divergent fight tactics (i.e., modes of feeing from an approaching predator), we predicted that (1) mule deer would respond to wolves with coarse-scale spatial shifts to rugged terrain favoring their stotting tactic; (2) white-tailed deer would manage wolf risk with fne-scale shifts toward gentle terrain facilitating their galloping tactic within their current home range. Resource selection functions based on 61 mule deer and 59 white-tailed deer equipped with GPS radio-collars from 2013 to 2016 revealed that habitat use for each species was altered by wolf presence, but in divergent ways that supported our predictions. Our fndings add to a growing literature highlighting fight behavior as a viable predictor of prey responses to predation risk across multiple ecosystem types. Consequently, they suggest that predators could initiate multiple indirect non-consumptive efects in the same ecosystem that are transmitted by divergent responses of sympatric prey with diferent fight tactics. Keywords Canis lupus · Galloping · Mule deer · Non-consumptive efects · Odocoileus hemionus · O. virginianus · Predation risk · Stotting · White-tailed deer Introduction Behavioral responses of prey to predators often take the form of shifts in habitat use, which in turn may infuence Communicated by Anders Angerbjörn. how prey exploit resources and interact with co-occurring Electronic supplementary material The online version of this species (Lima and Dill 1990; Fortin et al. 2005; Thaker article (https​://doi.org/10.1007/s0044​2-018-4323-z) contains et al. 2011; Latombe et al. 2014). Prey individuals are supplementary material, which is available to authorized users. typically assumed to avoid their predators (Laundré et al. 2010), diminishing their ecological efects (e.g., forag- * Justin A. Dellinger [email protected] ing pressure) where predators are more numerous, while increasing their impacts where predators are relatively 1 School of Environmental and Forest Sciences, University scarce (Ripple and Beschta 2012). A growing literature, of Washington, Seattle, WA, USA however, suggests that prey responses to predators hinge 2 Wildlife Investigations Lab, California Department of Fish on key features of the interaction such as predator hunting and Wildlife, Rancho Cordova, CA, USA mode and landscape context and, as a result, that anti- 3 Department of Biological Sciences, Florida International predator habitat shifts and their consequences will not University, North Miami, FL, USA always follow this pattern (Schmitz 2008; Heithaus et al. 4 Global Trophic Cascades Program, Department of Forest 2009; Wirsing et al. 2010). For example, Schmitz (2008) Ecosystems and Society, Oregon State University, Corvallis, found that the hunting mode of sympatric spider predators OR, USA Vol.:(0123456789)1 3 Oecologia (sit-and-wait versus active) dictated whether herbivorous and white-tailed deer (O. virginianus)—with diferent fight grasshoppers (Melanoplus femurrubrum) foraged in refuge tactics. or exposed grassland habitat and, as a result, promoted In 2008, gray wolves began naturally recolonizing Wash- either increased or reduced plant diversity. Working in ington from southern British Columbia, Canada, and north- a coral reef ecosystem, Catano et al. (2016) observed ern Idaho, and there are now 20 confrmed packs in the state that landscape heterogeneity mediated avoidance of a (Jimenez and Becker 2016). At present, these packs are dis- predator decoy by herbivorous fshes, with reef complex- tributed heterogeneously across eastern Washington, setting ity apparently enhancing avoidance, because it impedes the stage for natural experiments examining the efects of predator detection and facilitates prey escape. By impli- gray wolf recolonization on prey populations, including cation, eforts to identify the factors that determine how habitat use patterns, by contrasting areas with and without prey use space when threatened with predation are crucial resident wolves. In this region, mule deer and white-tailed to the development of a general framework for predict- deer dominate the ungulate guild (Robinson et al. 2002), ing the efects of predators on their prey and, ultimately, and gray wolves are known to primarily consume both spe- ecosystems. cies (making up ≥ 50% of wolf diet depending on the pack) Flight behavior, or the means by which prey fee from locally (Spence 2017). Mule deer and white-tailed deer may approaching predators (Ydenberg and Dill 1986), has been difer in their responses to the presence of wolves, however, highlighted as an important driver of context dependence in because their respective fight behaviors are believed to be predator–prey relationships that can interact with predator most efective in diferent habitat types (Lingle and Pellis hunting mode and landscape features to determine prey spa- 2002). Specifcally, mule deer stot (i.e., bound in a forward tial responses to predation risk (Lingle 2002; Heithaus et al. direction with all four legs touching the ground simultane- 2009; Wirsing et al. 2010). Overall predation risk can be ously) when approached by canid predators, and the ef- deconstructed into the probability of encountering a preda- cacy of this running gait as a means of escaping an attack tor (pre-encounter risk) and the probability of death as a is thought to be facilitated by uneven terrain (Geist 1981; result of the encounter (post-encounter risk) (Lima and Dill Lingle 2002). White-tailed deer, by contrast, are believed 1990; Hebblewhite et al. 2005). Thus, prey individuals with to be better equipped to evade canids on relatively gentle certain fight tactics might actually beneft from minimizing terrain, because they sprint away from predators and rely post-encounter risk using space where predators are rela- on the early detection (Kunkel and Pletscher 2001; Lingle tively abundant but less lethal, whereas prey species with and Pellis 2002; Kittle et al. 2008). As coursing predators, fight tactics that do not facilitate surviving an encounter gray wolves tend to select for relatively gentle terrain while with a predator should seek to minimize pre-encounter risk hunting (see ESM 1; Mech and Peterson 2003; Oakleaf by avoiding the predator altogether (Heithaus et al. 2009; et al. 2006). Thus, we might expect mule deer to exhibit Wirsing et al. 2010). In a given ecosystem, the hunting mode coarse-scale habitat shifts from gentle to rugged terrain of the predator and the structure of the landscape will deter- that reduce the likelihood of encounters with gray wolves mine the spatial pattern of predator lethality experienced (i.e., pre-encounter risk) and avoid substrates that hinder the by each prey species and, consequently, which prey species efectiveness of stotting in the event of an attack. Conversely, might beneft from managing pre- versus post-encounter risk white-tailed deer exposed to wolves should be more likely to (Heithaus et al. 2009; Wirsing et al. 2010). This framework remain in areas dominated by gentle terrain, given that their has implications for the spatial scale at which prey species running gait is not well suited to rugged areas, and exhibit should respond to the risk of predation. Namely, prey spe- fner-scale shifts in habitat use that facilitate sprinting in the cies whose fight behavior confers a high likelihood of sur- event of a predator encounter. viving an encounter should be better able to manage risk Here, taking advantage of spatial heterogeneity in wolf through fne-scale (i.e., within home range) habitat shifts presence, we contrasted both coarse- and fne-scale habi- when exposed to the threat of predation, whereas those less tat use patterns of adult mule deer and white-tailed deer able to survive an attack should be more likely to manage in areas with and without established packs. We hypoth- pre-encounter risk via coarse-scale shifts in habitat use (i.e., esized that the nature and scale of anti-predator responses changes to home range location; Kaufman et al. 2007). To of these two deer species to wolves are mediated by fight date, a few studies have assessed the relationship between behavior. Under this hypothesis, we expected mule deer fight behavior and habitat use patterns of multiple prey spe- to seek to avoid wolf encounters by increasing their use of cies at multiple spatial scales in response to predation risk rugged terrain and, consequently, that diferences in habi- from a shared predator. Accordingly, we examined the pos-
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