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D.W. Smith

Publications and source records attributed to D.W. Smith.

11 recordsLinked to original sources

Nonlinear effects of group size on the success of wolves hunting elk

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.

Behavioral Ecology↗

Estimation of successful breeding pairs for wolves in the Northern Rocky Mountains, USA

Under the Endangered Species Act, documenting recovery and federally mandated population levels of wolves (Canis lupus) in the Northern Rocky Mountains (NRM) requires monitoring wolf packs that successfully recruit young. United States Fish and Wildlife Service regulations define successful breeding pairs as packs estimated to contain an adult male and female, accompanied by ???2 pups on 31 December of a given year. Monitoring successful breeding pairs will become more difficult following proposed delisting of NRM wolves; alternatives to historically intensive methods, appropriate to the different ecological and regulatory context following delisting, are required. Because pack size is easier to monitor than pack composition, we estimated probability a pack would contain a successful breeding pair based on its size for wolf populations inhabiting 6 areas in the NRM. We also evaluated the extent to which differences in demography of wolves and levels of human-caused mortality among the areas influenced the probability of packs of different sizes would contain successful breeding pairs. Probability curves differed among analysis areas, depending primarily on levels of human-caused mortality, secondarily on annual population growth rate, and little on annual population density. Probabilities that packs contained successful breeding pairs were more uniformly distributed across pack sizes in areas with low levels of human mortality and stable populations. Large packs in areas with high levels of human-caused mortality and high annual growth rates had relatively high probabilities of containing breeding pairs whereas those for small packs were relatively low. Our approach can be used by managers to estimate number of successful breeding pairs in a population where number of packs and their sizes are known. Following delisting of NRM wolves, human-caused mortality is likely to increase, resulting in more small packs with low probabilities of containing breeding pairs. Differing contributions of packs to wolf population growth based on their size suggests monitoring successful breeding pairs will provide more accurate insights into population dynamics of wolves than will monitoring number of packs or individuals only.

Journal of Wildlife Management↗

Persistence of canine distemper virus in the Greater Yellowstone Ecosystem's carnivore community

Canine distemper virus (CDV) is an acute, highly immunizing pathogen that should require high densities and large populations of hosts for long-term persistence, yet CDV persists among terrestrial carnivores with small, patchily distributed groups. We used CDV in the Greater Yellowstone ecosystem's (GYE) wolves (Canis lupus) and coyotes (Canis latrans) as a case study for exploring how metapopulation structure, host demographics, and multi-host transmission affect the critical community size and spatial scale required for CDV persistence. We illustrate how host spatial connectivity and demographic turnover interact to affect both local epidemic dynamics, such as the length and variation in inter-epidemic periods, and pathogen persistence using stochastic, spatially explicit susceptible-exposed-infectious-recovered simulation models. Given the apparent absence of other known persistence mechanisms (e.g., a carrier or environmental state, densely populated host, chronic infection, or a vector), we suggest that CDV requires either large spatial scales or multi-host transmission for persistence. Current GYE wolf populations are probably too small to support endemic CDV. Coyotes are a plausible reservoir host, but CDV would still require 50 000-100 000 individuals for moderate persistence (>50% over 10 years), which would equate to an area of 1-3 times the size of the GYE (60000-200000 km2). Coyotes, and carnivores in general, are not uniformly distributed; therefore, this is probably a gross underestimate of the spatial scale of CDV persistence. However, the presence of a second competent host species can greatly increase the probability of long-term CDV persistence at much smaller spatial scales. Although no management of CDV is currently recommended for the GYE, wolf managers in the region should expect periodic but unpredictable CDV-related population declines as often as every 2-5 years. Awareness and monitoring of such outbreaks will allow corresponding adjustments in management activities such as regulated public harvest, creating a smooth transition to state wolf management and conservation after >30 years of being protected by the Endangered Species Act.

Ecological Applications↗

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. ?? 2009 Blackwell Publishing Ltd/CNRS.

Ecology Letters↗

A serological survey of infectious disease in Yellowstone National Park's canid community

Background: Gray wolves (Canis lupus) were reintroduced into Yellowstone National Park (YNP) after a >70 year absence, and as part of recovery efforts, the population has been closely monitored. In 1999 and 2005, pup survival was significantly reduced, suggestive of disease outbreaks. Methodology/Principal Findings: We analyzed sympatric wolf, coyote ( Canis latrans ), and red fox ( Vulpes vulpes ) serologic data from YNP, spanning 1991–2007, to identify long-term patterns of pathogen exposure, identify associated risk factors, and examine evidence for disease-induced mortality among wolves for which there were survival data. We found high, constant exposure to canine parvovirus (wolf seroprevalence: 100%; coyote: 94%), canine adenovirus-1 (wolf pups [0.5–0.9 yr]: 91%, adults [≥1 yr]: 96%; coyote juveniles [0.5–1.5 yrs]: 18%, adults [≥1.6 yrs]: 83%), and canine herpesvirus (wolf: 87%; coyote juveniles: 23%, young adults [1.6–4.9 yrs]: 51%, old adults [≥5 yrs]: 87%) suggesting that these pathogens were enzootic within YNP wolves and coyotes. An average of 50% of wolves exhibited exposure to the protozoan parasite, Neospora caninum , although individuals’ odds of exposure tended to increase with age and was temporally variable. Wolf, coyote, and fox exposure to canine distemper virus (CDV) was temporally variable, with evidence for distinct multi-host outbreaks in 1999 and 2005, and perhaps a smaller, isolated outbreak among wolves in the interior of YNP in 2002. The years of high wolf-pup mortality in 1999 and 2005 in the northern region of the park were correlated with peaks in CDV seroprevalence, suggesting that CDV contributed to the observed mortality. Conclusions/Significance: Of the pathogens we examined, none appear to jeopardize the long-term population of canids in YNP. However, CDV appears capable of causing short-term population declines. Additional information on how and where CDV is maintained and the frequency with which future epizootics might be expected might be useful for future management of the Northern Rocky Mountain wolf population.

PLoS ONE↗

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

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. ?? 2009 British Ecological Society.

Journal of Animal Ecology↗

Landscape heterogeneity shapes predation in a newly restored predator-prey system

Because some native ungulates have lived without top predators for generations, it has been uncertain whether runaway predation would occur when predators are newly restored to these systems. We show that landscape features and vegetation, which influence predator detection and capture of prey, shape large-scale patterns of predation in a newly restored predator-prey system. We analysed the spatial distribution of wolf (Canis lupus) predation on elk (Cervus elaphus) on the Northern Range of Yellowstone National Park over 10 consecutive winters. The influence of wolf distribution on kill sites diminished over the course of this study, a result that was likely caused by territorial constraints on wolf distribution. In contrast, landscape factors strongly influenced kill sites, creating distinct hunting grounds and prey refugia. Elk in this newly restored predator-prey system should be able to mediate their risk of predation by movement and habitat selection across a heterogeneous risk landscape. ?? 2007 Blackwell Publishing Ltd/CNRS.

Ecology Letters↗

A proposed ethogram of large-carnivore predatory behavior, exemplified by the wolf

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. ?? 2007 American Society of Mammalogists.

Journal of Mammalogy↗

Unusual behavior by Bison, Bison bison, toward Elk, Cervus elaphus, and wolves, Canis lupus

Incidents are described of Bison ( Bison bison ) in Yellowstone National Park mauling and possibly killing a young Elk ( Cervus elaphus ) calf, chasing wolves ( Canis lupus ) off Elk they had just killed or were killing, and keeping the wolves away for extended periods. During one of the latter cases, the Bison knocked a wolf-wounded Elk down. Bison were also seen approaching wolves that were resting and sleeping, rousting them, following them to new resting places and repeating this behavior. These behaviors might represent some type of generalized hyper-defensiveness that functions as an anti-predator strategy.

Idaho, Montana, Wyoming↗

Large carnivores response to recreational big game hunting along the Yellowstone National Park and Absaroka-Beartooth Wilderness boundary

The Greater Yellowstone Ecosystem contains the rare combination of an intact guild of native large carnivores, their prey, and differing land management policies (National Park versus National Forest; no hunting versus hunting). Concurrent field studies on large carnivores allowed us to investigate activities of humans and carnivores on Yellowstone National Park's (YNP) northern boundary. Prior to and during the backcountry big-game hunting season, we monitored movements of grizzly bears ( Ursus arctos ), wolves ( Canis lupus ), and cougars ( Puma concolor ) on the northern boundary of YNP. Daily aerial telemetry locations (September 1999), augmented with weekly telemetry locations (August and October 1999), were obtained for 3 grizzly bears, 7 wolves in 2 groups of 1 pack, and 3 cougars in 1 family group. Grizzly bears were more likely located inside the YNP boundary during the pre-hunt period and north of the boundary once hunting began. The cougar family tended to be found outside YNP during the pre-hunt period and moved inside YNP when hunting began. Wolves did not significantly change their movement patterns during the pre-hunt and hunting periods. Qualitative information on elk ( Cervus elaphus ) indicated they moved into YNP once hunting started, suggesting that cougars followed living prey or responded to hunting activity, grizzly bears focused on dead prey (e.g., gut piles, crippled elk), and wolves may have taken advantage of both. Measures of association (Jacob's Index) were positive within carnivore species but inconclusive among species. Further collaborative research and the use of new technologies such as Global Positioning System (GPS) telemetry collars will advance our ability to understand these species, the carnivore community and its interactions, and human influences on carnivores.

Wildlife Society Bulletin↗

Denning behaviour of non-gravid wolves, Canis lupus

Wild wolves (Canis lupus) that had produced pups in earlier years but were not currently pregnant, and ovariectomized captive wolves, dug dens during and after the whelping season even though they produced no pups. These observations suggest that den digging is not a function of pregnancy or of ovarian estrogen or progesterone. We hypothesize that increasing prolactin in spring elicits or mediates den-digging behavior.

Canadian Field-Naturalist↗