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      Differential Habitat Use or Intraguild Interactions: What Structures a Carnivore Community?

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          Abstract

          Differential habitat use and intraguild competition are both thought to be important drivers of animal population sizes and distributions. Habitat associations for individual species are well-established, and interactions between particular pairs of species have been highlighted in many focal studies. However, community-wide assessments of the relative strengths of these two factors have not been conducted. We built multi-scale habitat occupancy models for five carnivore taxa of New York’s Adirondack landscape and assessed the relative performance of these models against ones in which co-occurrences of potentially competing carnivore species were also incorporated. Distribution models based on habitat performed well for all species. Black bear ( Ursus americanus) and fisher ( Martes pennanti) distribution was similar in that occupancy of both species was negatively associated with paved roads. However, black bears were also associated with larger forest fragments and fishers with smaller forest fragments. No models with habitat features were more supported than the null habitat model for raccoons ( Procyon lotor). Martens ( Martes americana) were most associated with increased terrain ruggedness and elevation. Weasel ( Mustela spp.) occupancy increased with the cover of deciduous forest. For most species dyads habitat-only models were more supported than those models with potential competitors incorporated. The exception to this finding was for the smallest carnivore taxa (marten and weasel) where habitat plus coyote abundance models typically performed better than habitat-only models. Assessing this carnivore community as whole, we conclude that differential habitat use is more important than species interactions in maintaining the distribution and structure of this carnivore guild.

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          Predator interactions, mesopredator release and biodiversity conservation.

          There is growing recognition of the important roles played by predators in regulating ecosystems and sustaining biodiversity. Much attention has focused on the consequences of predator-regulation of herbivore populations, and associated trophic cascades. However apex predators may also control smaller 'mesopredators' through intraguild interactions. Removal of apex predators can result in changes to intraguild interactions and outbreaks of mesopredators ('mesopredator release'), leading in turn to increased predation on smaller prey. Here we provide a review and synthesis of studies of predator interactions, mesopredator release and their impacts on biodiversity. Mesopredator suppression by apex predators is widespread geographically and taxonomically. Apex predators suppress mesopredators both by killing them, or instilling fear, which motivates changes in behaviour and habitat use that limit mesopredator distribution and abundance. Changes in the abundance of apex predators may have disproportionate (up to fourfold) effects on mesopredator abundance. Outcomes of interactions between predators may however vary with resource availability, habitat complexity and the complexity of predator communities. There is potential for the restoration of apex predators to have benefits for biodiversity conservation through moderation of the impacts of mesopredators on their prey, but this requires a whole-ecosystem view to avoid unforeseen negative effects. 'Nothing has changed since I began. My eye has permitted no change. I am going to keep things like this.' From 'Hawk Roosting', by Ted Hughes.
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            Does interference competition with wolves limit the distribution and abundance of coyotes?

            Interference competition with wolves Canis lupus is hypothesized to limit the distribution and abundance of coyotes Canis latrans, and the extirpation of wolves is often invoked to explain the expansion in coyote range throughout much of North America. We used spatial, seasonal and temporal heterogeneity in wolf distribution and abundance to test the hypothesis that interference competition with wolves limits the distribution and abundance of coyotes. From August 2001 to August 2004, we gathered data on cause-specific mortality and survival rates of coyotes captured at wolf-free and wolf-abundant sites in Grand Teton National Park (GTNP), Wyoming, USA, to determine whether mortality due to wolves is sufficient to reduce coyote densities. We examined whether spatial segregation limits the local distribution of coyotes by evaluating home-range overlap between resident coyotes and wolves, and by contrasting dispersal rates of transient coyotes captured in wolf-free and wolf-abundant areas. Finally, we analysed data on population densities of both species at three study areas across the Greater Yellowstone Ecosystem (GYE) to determine whether an inverse relationship exists between coyote and wolf densities. Although coyotes were the numerically dominant predator, across the GYE, densities varied spatially and temporally in accordance with wolf abundance. Mean coyote densities were 33% lower at wolf-abundant sites in GTNP, and densities declined 39% in Yellowstone National Park following wolf reintroduction. A strong negative relationship between coyote and wolf densities (beta = -3.988, P < 0.005, r(2) = 0.54, n = 16), both within and across study sites, supports the hypothesis that competition with wolves limits coyote populations. Overall mortality of coyotes resulting from wolf predation was low, but wolves were responsible for 56% of transient coyote deaths (n = 5). In addition, dispersal rates of transient coyotes captured at wolf-abundant sites were 117% higher than for transients captured in wolf-free areas. Our results support the hypothesis that coyote abundance is limited by competition with wolves, and suggest that differential effects on survival and dispersal rates of transient coyotes are important mechanisms by which wolves reduce coyote densities.
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              Competition and intraguild predation among three sympatric carnivores.

              We examined the relative roles of dominance in agonistic interactions and energetic constraints related to body size in determining local abundances of coyotes (Canis latrans, 8-20 kg), gray foxes (Urocyon cinereoargenteus, 3-5 kg) and bobcats (Felis rufus, 5-15 kg) at three study sites (hereafter referred to as NP, CP, and SP) in the Santa Monica Mountains of California. We hypothesized that the largest and behaviorally dominant species, the coyote, would exploit a wider range of resources (i.e., a higher number of habitat and/or food types) and, consequently, would occur in higher density than the other two carnivores. We evaluated our hypotheses by quantifying their diets, food overlap, habitat-specific abundances, as well as their overall relative abundance at the three study sites. We identified behavioral dominance of coyotes over foxes and bobcats in Santa Monica because 7 of 12 recorded gray fox deaths and 2 of 5 recorded bobcat deaths were due to coyote predation, and no coyotes died as a result of their interactions with bobcats or foxes. Coyotes and bobcats were present in a variety of habitats types (8 out of 9), including both open and brushy habitats, whereas gray foxes were chiefly restricted to brushy habitats. There was a negative relationship between the abundances of coyotes and gray foxes (P=0.020) across habitats, suggesting that foxes avoided habitats of high coyote predation risk. Coyote abundance was low in NP, high in CP, and intermediate in SP. Bobcat abundance changed little across study sites, and gray foxes were very abundant in NP, absent in CP, and scarce in SP; this suggests a negative relationship between coyote and fox abundances across study sites, as well. Bobcats were solely carnivorous, relying on small mammals (lagomorphs and rodents) throughout the year and at all three sites. Coyotes and gray foxes also relied on small mammals year-round at all sites, though they also ate significant amounts of fruit. Though there were strong overall interspecific differences in food habits of carnivores (P<0.0001), average seasonal food overlaps were high due to the importance of small mammals in all carnivore diets [bobcat-gray fox: 0.79±0.09 (SD), n=4; bobcat-coyote: 0.69±0.16, n=6; coyote-gray fox: 0.52±0.05, n=4]. As hypothesized, coyotes used more food types and more habitat types than did bobcats and gray foxes and, overall, coyotes were the most abundant of the three species and ranged more widely than did gray foxes. We propose that coyotes limit the number and distribution of gray foxes in Santa Monica Mountains, and that those two carnivores exemplified a case in which the relationship between their body size and local abundance is governed by competitive dominance of the largest species rather than by energetic equivalences. However, in the case of the intermediate-sized bobcat no such a pattern emerged, likely due to rarity or inconsistency of agonistic interactions and/or behavioral avoidance of encounters by subordinate species.
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                Author and article information

                Contributors
                Role: Editor
                Journal
                PLoS One
                PLoS ONE
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, CA USA )
                1932-6203
                5 January 2016
                2016
                : 11
                : 1
                : e0146055
                Affiliations
                [1 ]Department of Fisheries & Wildlife Sciences, University of Missouri, Columbia, Missouri, United States of America
                [2 ]USDA Forest Service, Pacific Northwest Research Station, Forestry Sciences Laboratory, Corvallis, Oregon, United States of America
                [3 ]Wildlife Conservation Society Canada, Toronto, Ontario, Canada
                [4 ]Faculty of Forestry, University of Toronto, Toronto, Ontario, Canada
                [5 ]North Carolina Museum of Natural Sciences and North Carolina State University, Raleigh, North Carolina, United States of America
                University of Missouri Kansas City, UNITED STATES
                Author notes

                Competing Interests: The authors have declared that no competing interests exist.

                Conceived and designed the experiments: MEG JCR RK. Performed the experiments: MEG DBL JCR JRM RK. Analyzed the data: JRM DBL. Contributed reagents/materials/analysis tools: MEG DBL JCR RK. Wrote the paper: MEG DBL JCR JRM RK.

                Article
                PONE-D-15-33540
                10.1371/journal.pone.0146055
                4711579
                26731404
                5e8a74b9-5cd7-4db3-9136-c6e978014dd5

                This is an open access article, free of all copyright, and may be freely reproduced, distributed, transmitted, modified, built upon, or otherwise used by anyone for any lawful purpose. The work is made available under the Creative Commons CC0 public domain dedication.

                History
                : 31 July 2015
                : 11 December 2015
                Page count
                Figures: 2, Tables: 5, Pages: 18
                Funding
                Financial support for this work came from the Wildlife Conservation Society, the Geraldine R. Dodge Foundation, the National Geographic Society, the New York State Museum, the New York Biodiversity Research Institute, the New York Department of Environmental Conservation, and the National Science Foundation (DEB 0347609).
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