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MacNulty D.R.,University of Minnesota | Smith D.W.,Yellowstone Center for Resources | Mech L.D.,U.S. Geological Survey | Packer C.,University of Minnesota
Behavioral Ecology | Year: 2012

Despite the popular view that social predators live in groups because group hunting facilitates prey capture, the apparent tendency for hunting success to peak at small group sizes suggests that the formation of large groups is unrelated to prey capture. Few empirical studies, however, have tested for nonlinear relationships between hunting success and group size, and none have demonstrated why success trails off after peaking. Here, we use a unique dataset of observations of individually known wolves (Canis lupus) hunting elk (Cervus elaphus) in Yellowstone National Park to show that the relationship between success and group size is indeed nonlinear and that individuals withholding effort (free riding) is why success does not increase across large group sizes. Beyond 4 wolves, hunting success leveled off, and individual performance (a measure of effort) decreased for reasons unrelated to interference from inept hunters, individual age, or size. But performance did drop faster among wolves with an incentive to hold back, i.e., nonbreeders with no dependent offspring, those performing dangerous predatory tasks, i.e., grabbing and restraining prey, and those in groups of proficient hunters. These results suggest that decreasing performance was free riding and that was why success leveled off in groups with >4 wolves that had superficially appeared to be cooperating. This is the first direct evidence that nonlinear trends in group hunting success reflect a switch from cooperation to free riding. It also highlights how hunting success per se is unlikely to promote formation and maintenance of large groups. © 2011 The Author. Source


Vucetich J.A.,Michigan Technological University | Hebblewhite M.,University of Montana | Smith D.W.,Yellowstone Center for Resources | Peterson R.O.,Michigan Technological University
Journal of Animal Ecology | Year: 2011

1.Predation rate (PR) and kill rate are both fundamental statistics for understanding predation. However, relatively little is known about how these statistics relate to one another and how they relate to prey population dynamics. We assess these relationships across three systems where wolf-prey dynamics have been observed for 41years (Isle Royale), 19years (Banff) and 12years (Yellowstone). 2.To provide context for this empirical assessment, we developed theoretical predictions of the relationship between kill rate and PR under a broad range of predator-prey models including predator-dependent, ratio-dependent and Lotka-Volterra dynamics. 3.The theoretical predictions indicate that kill rate can be related to PR in a variety of diverse ways (e.g. positive, negative, unrelated) that depend on the nature of predator-prey dynamics (e.g. structure of the functional response). These simulations also suggested that the ratio of predator-to-prey is a good predictor of prey growth rate. That result motivated us to assess the empirical relationship between the ratio and prey growth rate for each of the three study sites. 4.The empirical relationships indicate that PR is not well predicted by kill rate, but is better predicted by the ratio of predator-to-prey. Kill rate is also a poor predictor of prey growth rate. However, PR and ratio of predator-to-prey each explained significant portions of variation in prey growth rate for two of the three study sites. 5.Our analyses offer two general insights. First, Isle Royale, Banff and Yellowstone are similar insomuch as they all include wolves preying on large ungulates. However, they also differ in species diversity of predator and prey communities, exploitation by humans and the role of dispersal. Even with the benefit of our analysis, it remains difficult to judge whether to be more impressed by the similarities or differences. This difficulty nicely illustrates a fundamental property of ecological communities. Second, kill rate is the primary statistic for many traditional models of predation. However, our work suggests that kill rate and PR are similarly important for understanding why predation is such a complex process. © 2011 The Authors. Journal of Animal Ecology © 2011 British Ecological Society. Source


Peterson R.O.,Michigan Technological University | Vucetich J.A.,Michigan Technological University | Bump J.M.,Michigan Technological University | Smith D.W.,Yellowstone Center for Resources
Annual Review of Ecology, Evolution, and Systematics | Year: 2014

Questions of whether trophic cascades occur in Isle Royale National Park (IRNP) or Yellowstone National Park's northern range (NR) cannot lead to simple, precise, or definitive answers. Such answers are limited especially by multicausality in the NR and by complex temporal variation in IRNP. Spatial heterogeneity, contingency, and nonequilibrium dynamics also work against simple answers in IRNP and NR. The existence of a trophic cascade in IRNP and NR also depends greatly on how it is defined. For example, some conceive of trophic cascades as entailing essentially any indirect effect of predation. This may be fine, but the primary intellectual value of such a conception may be to assess an important view in community ecology that most species are connected to most other species in a food web through a network of complicated, albeit weak, indirect effects. These circumstances that work against simple answers likely apply to many ecosystems. Despite the challenges of assessing the existence of trophic cascades in IRNP and NR, such assessments result in considerable insights about a more fundamental question: What causes population abundance to fluctuate? © 2014 by Annual Reviews. All rights reserved. Source


Cubaynes S.,University of Oxford | Macnulty D.R.,Utah State University | Stahler D.R.,Yellowstone Center for Resources | Quimby K.A.,Yellowstone Center for Resources | And 2 more authors.
Journal of Animal Ecology | Year: 2014

Understanding the population dynamics of top-predators is essential to assess their impact on ecosystems and to guide their management. Key to this understanding is identifying the mechanisms regulating vital rates. Determining the influence of density on survival is necessary to understand the extent to which human-caused mortality is compensatory or additive. In wolves (Canis lupus), empirical evidence for density-dependent survival is lacking. Dispersal is considered the principal way in which wolves adjust their numbers to prey supply or compensate for human exploitation. However, studies to date have primarily focused on exploited wolf populations, in which density-dependent mechanisms are likely weak due to artificially low wolf densities. Using 13 years of data on 280 collared wolves in Yellowstone National Park, we assessed the effect of wolf density, prey abundance and population structure, as well as winter severity, on age-specific survival in two areas (prey-rich vs. prey-poor) of the national park. We further analysed cause-specific mortality and explored the factors driving intraspecific aggression in the prey-rich northern area of the park. Overall, survival rates decreased during the study. In northern Yellowstone, density dependence regulated adult survival through an increase in intraspecific aggression, independent of prey availability. In the interior of the park, adult survival was less variable and density-independent, despite reduced prey availability. There was no effect of prey population structure in northern Yellowstone, or of winter severity in either area. Survival was similar among yearlings and adults, but lower for adults older than 6 years. Our results indicate that density-dependent intraspecific aggression is a major driver of adult wolf survival in northern Yellowstone, suggesting intrinsic density-dependent mechanisms have the potential to regulate wolf populations at high ungulate densities. When low prey availability or high removal rates maintain wolves at lower densities, limited inter-pack interactions may prevent density-dependent survival, consistent with our findings in the interior of the park. © 2014 The Authors. Source


Stahler D.R.,Yellowstone Center for Resources | Macnulty D.R.,Utah State University | Wayne R.K.,University of California at Los Angeles | vonHoldt B.,University of California at Los Angeles | Smith D.W.,Yellowstone Center for Resources
Journal of Animal Ecology | Year: 2013

Reproduction in social organisms is shaped by numerous morphological, behavioural and life-history traits such as body size, cooperative breeding and age of reproduction, respectively. Little is known, however, about the relative influence of these different types of traits on reproduction, particularly in the context of environmental conditions that determine their adaptive value. Here, we use 14 years of data from a long-term study of wolves (Canis lupus) in Yellowstone National Park, USA, to evaluate the relative effects of different traits and ecological factors on the reproductive performance (litter size and survival) of breeding females. At the individual level, litter size and survival improved with body mass and declined with age (c. 4-5 years). Grey-coloured females had more surviving pups than black females, which likely contributed to the maintenance of coat colour polymorphism in this system. The effect of pack size on reproductive performance was nonlinear as litter size peaked at eight wolves and then declined, and litter survival increased rapidly up to three wolves, beyond which it increased more gradually. At the population level, litter size and survival decreased with increasing wolf population size and canine distemper outbreaks. The relative influence of these different-level factors on wolf reproductive success followed individual > group > population. Body mass was the primary determinant of litter size, followed by pack size and population size. Body mass was also the main driver of litter survival, followed by pack size and disease. Reproductive gains because of larger body size and cooperative breeding may mitigate reproductive losses because of negative density dependence and disease. These findings highlight the adaptive value of large body size and sociality in promoting individual fitness in stochastic and competitive environments. © 2012 The Authors. Journal of Animal Ecology © 2012 British Ecological Society. Source

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