Daniel R. MacNulty

Winter severity and wolf predation on a formerly wolf-free elk herd

We studied wolf (Canis lupus) predation on elk (Cervus elaphus) in Yellowstone National Park from 17 March to 15 April 1997 (severe winter conditions) and from 2 to 31 March 1998 (mild winter conditions) 2-3 years after wolves were reintroduced to the park. Elk composed 91% of 117 kills. Data comparisons for 1997 versus 1998 were: hunting success rate, 26% versus 15%; kill rate, 17.1 kg/wolf/day versus 6.1; percent of kill consumed in first day 7 versus 86; percent femur marrow fat of adult kills, 27 versus 70; calf:adult ratios of kills, 2:33 versus 17:23; sex ratio of kills, 14M:19F versus 17M:6F; mean age of elk killed, males 6.1 years, females 15.2 versus males, 4.8, females 13.0. Winter severity influenced the wolf-elk relationship more than the naivete of the elk herd to predation by wolves.

Modeling Effects of Environmental Change on Wolf Population Dynamics, Trait Evolution, and Life History

Analysis of a 20-year data set on Yellowstone wolves reveals that environmental change generates eco-evolutionary responses. Environmental change has been observed to generate simultaneous responses in population dynamics, life history, gene frequencies, and morphology in a number of species. But how common are such eco-evolutionary responses to environmental change likely to be? Are they inevitable, or do they require a specific type of change? Can we accurately predict eco-evolutionary responses? We address these questions using theory and data from the study of Yellowstone wolves. We show that environmental change is expected to generate eco-evolutionary change, that changes in the average environment will affect wolves to a greater extent than changes in how variable it is, and that accurate prediction of the consequences of environmental change will probably prove elusive.

Predatory senescence in ageing wolves

It is well established that ageing handicaps the ability of prey to escape predators, yet surprisingly little is known about how ageing affects the ability of predators to catch prey. Research into long-lived predators has assumed that adults have uniform impacts on prey regardless of age. Here we use longitudinal data from repeated observations of individually-known wolves (Canis lupus) hunting elk (Cervus elaphus) in Yellowstone National Park to demonstrate that adult predatory performance declines with age and that an increasing ratio of senescent individuals in the wolf population depresses the rate of prey offtake. Because this ratio fluctuates independently of population size, predatory senescence may cause wolf populations of equal size but different age structure to have different impacts on prey populations. These findings suggest that predatory senescence is an important, though overlooked, factor affecting predator-prey dynamics.

Body size and predatory performance in wolves: is bigger better?

1. Large body size hinders locomotor performance in ways that may lead to trade-offs in predator foraging ability that limit the net predatory benefit of larger size. For example, size-related improvements in handling prey may come at the expense of pursuing prey and thus negate any enhancement in overall predatory performance due to increasing size. 2. This hypothesis was tested with longitudinal data from repeated observations of 94 individually known wolves (Canis lupus) hunting elk (Cervus elaphus) in Yellowstone National Park, USA. Wolf size was estimated from an individually based sex-specific growth model derived from body mass measurements of 304 wolves. 3. Larger size granted individual wolves a net predatory advantage despite substantial variation in its effect on the performance of different predatory tasks; larger size improved performance of a strength-related task (grappling and subduing elk) but failed to improve performance of a locomotor-related task (selecting an elk from a group) for wolves > 39 kg. 4. Sexual dimorphism in wolf size also explained why males outperformed females in each of the three tasks considered (attacking, selecting, and killing). 5. These findings support the generalization that bigger predators are overall better hunters, but they also indicate that increasing size ultimately limits elements of predatory behaviour that require superior locomotor performance. We argue that this could potentially narrow the dietary niche of larger carnivores as well as limit the evolution of larger size if prey are substantially more difficult to pursue than to handle.

A PROPOSED ETHOGRAM OF LARGE-CARNIVORE PREDATORY BEHAVIOR, EXEMPLIFIED BY THE WOLF

Abstract Although predatory behavior is traditionally described by a basic ethogram composed of 3 phases (search, pursue, and capture), behavioral studies of large terrestrial carnivores generally use the concept of a “hunt” to classify and measure foraging. This approach is problematic because there is no consensus on what behaviors constitute a hunt. We therefore examined how the basic ethogram could be used as a common framework for classifying large-carnivore behavior. We used >2,150 h of observed wolf (Canis lupus) behavior in Yellowstone National Park, including 517 and 134 encounters with elk (Cervus elaphus) and American bison (Bison bison), respectively, to demonstrate the functional importance of several frequently described, but rarely quantified, patterns of large-carnivore behavior not explicitly described by the basic ethogram (approaching, watching, and attacking groups). To account for these additionally important behaviors we propose a modified form of the basic ethogram (search, approach, watch, attack-group, attack-individual, and capture). We tested the applicability of this ethogram by comparing it to 31 previous classifications and descriptions involving 7 other species and 5 other wolf populations. Close correspondence among studies suggests that this ethogram may provide a generally useful scheme for classifying large-carnivore predatory behavior that is behaviorally less ambiguous than the concept of a hunt.

The adaptive value of morphological, behavioural and life-history traits in reproductive female wolves

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.

Density‐dependent intraspecific aggression regulates survival in northern Yellowstone wolves (Canis lupus)

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.

Influence of Group Size on the Success of Wolves Hunting Bison

An intriguing aspect of social foraging behaviour is that large groups are often no better at capturing prey than are small groups, a pattern that has been attributed to diminished cooperation (i.e., free riding) in large groups. Although this suggests the formation of large groups is unrelated to prey capture, little is known about cooperation in large groups that hunt hard-to-catch prey. Here, we used direct observations of Yellowstone wolves (Canis lupus) hunting their most formidable prey, bison (Bison bison), to test the hypothesis that large groups are more cooperative when hunting difficult prey. We quantified the relationship between capture success and wolf group size, and compared it to previously reported results for Yellowstone wolves hunting elk (Cervus elaphus), a prey that was, on average, 3 times easier to capture than bison. Whereas improvement in elk capture success levelled off at 2–6 wolves, bison capture success levelled off at 9–13 wolves with evidence that it continued to increase beyond 13 wolves. These results are consistent with the hypothesis that hunters in large groups are more cooperative when hunting more formidable prey. Improved ability to capture formidable prey could therefore promote the formation and maintenance of large predator groups, particularly among predators that specialize on such prey.

Multi-robot system based on model of wolf hunting behavior to emulate wolf and elk interactions

Wolves are one of the most successful large predators on earth. Their success is made apparent by their presence in most northern ecosystems. They owe much of this success to their generalized hunting behavior which allows them to quickly and effectively adjust to different species of prey. The success of this hunting behavior for wolves is the inspiration for a project to bestow this behavior onto a system of robots with the hopes that they might utilize the apparent strengths of the behavior to achieve their own success.

FIV Diversity: FIVPle Subtype Composition May Influence Disease Outcome in African Lions

Feline immunodeficiency virus (FIV) infects domestic cats and at least 20 additional species of non-domestic felids throughout the world. Strains specific to domestic cat (FIV(Fca)) produce AIDS-like disease progression, sequelae and pathology providing an informative model for HIV infection in humans. Less is known about the immunological and pathological influence of FIV in other felid species although multiple distinct strains of FIV circulate in natural populations. As in HIV-1 and HIV-2, multiple diverse cross-species infections may have occurred. In the Serengeti National Park, Tanzania, three divergent subtypes of lion FIV (FIV(Ple)) are endemic, whereby 100% of adult lions are infected with one or more of these strains. Herein, the relative distribution of these subtypes in the population are surveyed and, combined with observed differences in lion mortality due to secondary infections based on FIV(Ple) subtypes, the data suggest that FIV(Ple) subtypes may have different patterns of pathogenicity and transmissibility among wild lion populations.