Multi-prey System Research Articles

Predator home range size mediates indirect interactions between prey species in an arctic vertebrate community.

Indirect interactions are widespread among prey species that share a common predator, but the underlying mechanisms driving these interactions are often unclear, and our ability to predict their outcome is limited. Changes in behavioural traits that impact predator space use could be a key proximal mechanism mediating indirect interactions, but there is little empirical evidence of the causes and consequences of such behavioural-numerical response in multispecies systems. Here, we investigate the complex ecological relationships between seven prey species sharing a common predator. We used a path analysis approach on a comprehensive 9-year data set simultaneously tracking predator space use, prey densities and prey mortality rate on key species of a simplified Arctic food web. We show that high availability of a clumped and spatially predictable prey (goose eggs) leads to a twofold reduction in predator (arctic fox) home range size, which increases local predator density and strongly decreases nest survival of an incidental prey (American golden plover). On the contrary, a scattered cyclic prey with potentially lower spatial predictability (lemming) had a weaker effect on fox space use and an overall positive impact on the survival of incidental prey. These contrasting effects underline the importance of studying behavioural responses of predators in multiprey systems and to explicitly integrate behavioural-numerical responses in multispecies predator-prey models.

Mountain lion predation in a multi-prey system on private lands in Texas

Mountain lion predation in a multi-prey system on private lands in Texas

Predicting kill sites of an apex predator from GPS data in different multiprey systems.

Kill rates are a central parameter to assess the impact of predation on prey species. An accurate estimation of kill rates requires a correct identification of kill sites, often achieved by field-checking GPS location clusters (GLCs). However, there are potential sources of error included in kill-site identification, such as failing to detect GLCs that are kill sites, and misclassifying the generated GLCs (e.g., kill for nonkill) that were not field checked. Here, we address these two sources of error using a large GPS dataset of collared Eurasian lynx (Lynx lynx), an apex predator of conservation concern in Europe, in three multiprey systems, with different combinations of wild, semidomestic, and domestic prey. We first used a subsampling approach to investigate how different GPS-fix schedules affected the detection of GLC-indicated kill sites. Then, we evaluated the potential of the random forest algorithm to classify GLCs as nonkills, small prey kills, and ungulate kills. We show that the number of fixes can be reduced from seven to three fixes per night without missing more than 5% of the ungulate kills, in a system composed of wild prey. Reducing the number of fixes per 24 h decreased the probability of detecting GLCs connected with kill sites, particularly those of semidomestic or domestic prey, and small prey. Random forest successfully predicted between 73%-90% of ungulate kills, but failed to classify most small prey in all systems, with sensitivity (true positive rate) lower than 65%. Additionally, removing domestic prey improved the algorithm's overall accuracy. We provide a set of recommendations for studies focusing on kill-site detection that can be considered for other large carnivore species in addition to the Eurasian lynx. We recommend caution when working in systems including domestic prey, as the odds of underestimating kill rates are higher.

Combining mirid predators to reduce crop damage and sustain biocontrol in multi-prey systems

Combining mirid predators to reduce crop damage and sustain biocontrol in multi-prey systems

Wolf spatial behavior promotes encounters and kills of abundant prey.

Predators use different spatial tactics to track the prey on the landscape. Three hypotheses describe spatial tactics: prey abundance for prey that are aggregated in space; prey habitat for uniformly distributed prey; and prey catchability for prey that are difficult to catch and kill. The gray wolf (Canis lupus) is a generalist predator that likely employs more than one spatial hunting tactic to match their diverse prey with distinct distributions and behavior that are available. We conducted a study on 17 GPS collared wolves in 6 packs in Riding Mountain National Park, Manitoba, Canada where wolves prey on moose (Alces alces) and elk (Cervus canadensis). We evaluated wolf selection for prey density, habitat selection and catchability on the landscape through within-territory habitat selection analysis. We reveal support for both the prey habitat and prey catchability hypotheses. For moose, their primary prey, wolves employed a mixed habitat and catchability tactic. Wolves used spaces described by the intersection of moose habitat and moose catchability. Wolves selected for the catchability of elk, their secondary prey, but not elk habitat. Counter to our predictions, wolves avoided areas of moose and elk density, likely highlighting the ongoing space race between predator and prey. We illustrate that of the three hypotheses the primary driver was prey catchability, where the interplay of both prey habitat with catchability culminate in predator spatial behaviour in a multiprey system.

What drives wolf preference towards wild ungulates? Insights from a multi-prey system in the Slovak Carpathians.

The wolf is a generalist-opportunistic predator that displays diverse and remarkably adaptable feeding strategies across its range with local adaptations to certain prey species depending on their availability and vulnerability. The multi-prey system of the Slovak Carpathians supports important portion of the European wolf population; however, it has been markedly understudied. We evaluated winter diet composition and prey selection of Slovak wolves based on 321 scat samples collected between September–April within four different study areas during 2015–2017. The winter diet of wolves in the Slovak Carpathians was characterized by a 98% occurrence of wild large-sized and medium-sized ungulates with red deer occurring in wolf scats most often, consistent with their highest density among other wild ungulates. However, by comparing the consumption with availability of wild prey, we found that wolves in fact selected for wild boar especially in areas with higher altitudinal range, while selected for red deer in areas with low altitudinal range where this prey species was more spatially predictable. Although wolves showed the potential to switch between red deer and wild boar when their density increases, we found that this variation can be rather linked to changing prey vulnerability, which is dependent on particular environmental conditions at local scale such as topography and snow accumulation. The present study provides valuable insights into the winter foraging ecology of Slovak wolves in a multi-prey system of the Carpathians and allows for practical implications in the management of the rapidly increasing populations of wild ungulates across Europe.

The Role of Weather and Long-Term Prey Dynamics as Drivers of Wolf Population Dynamics in a Multi-Prey System

As climate change accelerates in northern latitudes, there is an increasing need to understand the role of climate in influencing predator-prey systems. We investigated wolf population dynamics and numerical response in Denali National Park and Preserve in Alaska, United States from 1986 to 2016 under a long-term range of varying climatic conditions and in the context of prey vulnerability, abundance, and population structure using an integrated population modeling approach. We found that wolf natality, or the number of wolves added to packs, increased with higher caribou population size, calf:cow ratio, and hare numbers, responding to a 1-year lag. Apparent survival increased in years with higher calf:cow ratios and cumulative snowfall in the prior winter, indicators of a vulnerable prey base. Thus, indices of prey abundance and vulnerability led to responses in wolf demographics, but we did not find that the wolf population responded numerically. During recent caribou and moose population increases wolf natality increased yet wolf population size declined. The decline in wolf population size is attributed to fewer packs in recent years with a few very large packs as opposed to several packs of comparable size. Our results suggest that territoriality can play a vital role in our study area on regulating population growth. These results provide a baseline comparison of wolf responses to climatic and prey variability in an area with relatively low levels of human disturbance, a rare feature in wolf habitat worldwide.

Gray wolf (Canis lupus) predation patterns following recent recolonization in a multi-predator, multi-prey system

Predator–prey interactions are among the most fundamental of ecological relationships. Recolonizing gray wolf (Canis lupus Linnaeus, 1758) populations present new challenges for wildlife management in multi-prey, multi-carnivore systems. We documented diet composition and kill rates for wolves in a recently recolonized area over winter and summer seasons (2014–2015). Elk (Cervus canadensis Erxleben, 1777) were the primary ungulate prey (63%) located at wolf kill sites. Deer (mule deer (Odocoileus hemionus (Rafinesque, 1817)) and white-tailed deer (Odocoileus virginianus (Zimmermann, 1780))) were less prevalent than elk in wolf diets, but the amount of deer in diets (40%–50%) varied by pack and season. Juvenile elk were the most prevalent class of prey in wolf diets during summer (63.3%) and winter (36.3%), with adult elk (32.5%) observed nearly as often as juveniles in winter. Kill rates varied by season, with rates 2.3 times higher in summer (mean = 3.5 ungulates/week per pack) than winter (mean = 1.5 ungulates/week per pack), consistent with increased availability and use of neonate prey. Prey biomass acquisition did not vary by pack or season (summer = 243 kg/week per pack; winter = 182 kg/week per pack). Our study quantified predation patterns for a recolonizing wolf population, and patterns that we documented were similar to other multi-prey systems in North America.

Interspecific prey neighborhoods shape risk of predation in a savanna ecosystem.

The vulnerability of an individual to predation depends on the availability of other prey items in the surrounding environment. Interspecific prey aggregations or "neighborhoods" may therefore affect an individual's vulnerability to predation. We examined the influence of prey neighborhood structure (i.e., the densities and identities of prey neighborhoods) on spatial variation in predation in a multi-prey system with a primary apex predator. We combined GPS locations of lions (Panthera leo), kill-site surveys, and spatially explicit density estimates of five species of ungulates for which a significant level of predation was attributable to lions. In addition to the dual influence of predator activity and vegetation, predation risk was attributable to the structure of prey neighborhoods for at least two of the five species of prey. Along with traditionally recognized components of predation (the rate of predator-prey encounters and prey catchability), we encourage ecologists to consider how prey neighborhood structure influences spatial variation in predation risk.

A spatial theory for characterizing predator-multiprey interactions in heterogeneous landscapes.

Trophic interactions in multiprey systems can be largely determined by prey distributions. Yet, classic predator-prey models assume spatially homogeneous interactions between predators and prey. We developed a spatially informed theory that predicts how habitat heterogeneity alters the landscape-scale distribution of mortality risk of prey from predation, and hence the nature of predator interactions in multiprey systems. The theoretical model is a spatially explicit, multiprey functional response in which species-specific advection-diffusion models account for the response of individual prey to habitat edges. The model demonstrates that distinct responses of alternative prey species can alter the consequences of conspecific aggregation, from increasing safety to increasing predation risk. Observations of threatened boreal caribou, moose and grey wolf interacting over 378 181 km(2) of human-managed boreal forest support this principle. This empirically supported theory demonstrates how distinct responses of apparent competitors to landscape heterogeneity, including to human disturbances, can reverse density dependence in fitness correlates.

Functional responses of the rough-legged buzzard in a multi-prey system

The functional response is a key element of predator-prey interactions. Basic functional response theory explains foraging behavior of individual predators, but many empirical studies of free-ranging predators have estimated functional responses by using population-averaged data. We used a novel approach to investigate functional responses of an avian predator (the rough legged-buzzard Buteo lagopus Pontoppidan, 1763) to intra-annual spatial variation in rodent density in subarctic Sweden, using breeding pairs as the sampling unit. The rough-legged buzzards responded functionally to Norwegian lemmings (Lemmus lemmus L. 1758), grey-sided voles (Myodes rufocanus Sundevall, 1846) and field voles (Microtus agrestis L. 1761), but different rodent prey were not utilised according to relative abundance. The functional response to Norwegian lemmings was a steep type II curve and a more shallow type III response to grey-sided voles. The different shapes of these two functional responses were likely due to combined effects of differences between lemmings and grey-sided voles in habitat utilisation, anti-predator behaviour and size-dependent vulnerability to predation. Diet composition changed less than changes in relative prey abundance, indicating negative switching, with high disproportional use of especially lemmings at low relative densities. Our results suggest that lemmings and voles should be treated separately in future empirical and theoretical studies in order to better understand the role of predation in this study system.

Seasonal foraging ecology of non-migratory cougars in a system with migrating prey.

We tested for seasonal differences in cougar (Puma concolor) foraging behaviors in the Southern Yellowstone Ecosystem, a multi-prey system in which ungulate prey migrate, and cougars do not. We recorded 411 winter prey and 239 summer prey killed by 28 female and 10 male cougars, and an additional 37 prey items by unmarked cougars. Deer composed 42.4% of summer cougar diets but only 7.2% of winter diets. Males and females, however, selected different proportions of different prey; male cougars selected more elk (Cervus elaphus) and moose (Alces alces) than females, while females killed greater proportions of bighorn sheep (Ovis canadensis), pronghorn (Antilocapra americana), mule deer (Odocoileus hemionus) and small prey than males. Kill rates did not vary by season or between males and females. In winter, cougars were more likely to kill prey on the landscape as: 1) elevation decreased, 2) distance to edge habitat decreased, 3) distance to large bodies of water decreased, and 4) steepness increased, whereas in summer, cougars were more likely to kill in areas as: 1) elevation decreased, 2) distance to edge habitat decreased, and 3) distance from large bodies of water increased. Our work highlighted that seasonal prey selection exhibited by stationary carnivores in systems with migratory prey is not only driven by changing prey vulnerability, but also by changing prey abundances. Elk and deer migrations may also be sustaining stationary cougar populations and creating apparent competition scenarios that result in higher predation rates on migratory bighorn sheep in winter and pronghorn in summer. Nevertheless, cougar predation on rare ungulates also appeared to be influenced by individual prey selection.

Wolves, white‐tailed deer, and beaver: implications of seasonal prey switching for woodland caribou declines

Population increases of primary prey can negatively impact alternate prey populations via demographic and behavioural responses of a shared predator through apparent competition. Seasonal variation in prey selection patterns by predators also can affect secondary and incidental prey by reducing spatial separation. Global warming and landscape changes in Alberta's bitumen sands have resulted in prey enrichment, which is changing the large mammal predator–prey system and causing declines in woodland caribou Rangifer tarandus caribou populations. We assessed seasonal patterns of prey use and spatial selection by wolves Canis lupus in two woodland caribou ranges in northeastern Alberta, Canada, that have undergone prey enrichment following recent white‐tailed deer Odocoileus virginianus invasion. We determined whether risk of predation for caribou (incidental prey) and the proportion of wolf‐caused‐caribou mortalities varied with season. We found that wolves showed seasonal variation in primary prey use, with deer and beaver Castor canadensis being the most common prey items in wolf diet in winter and summer, respectively. These seasonal dietary patterns were reflected in seasonal wolf spatial resource selection and resulted in contrasting spatial relationships between wolves and caribou. During winter, wolf selection for areas used by deer maintained strong spatial separation between wolves and caribou, whereas wolf selection for areas used by beaver in summer increased the overlap with caribou. Changing patterns in wolf resource selection were reflected by caribou mortality patterns, with 76.2% of 42 adult female caribou mortalities occurring in summer. Understanding seasonal patterns of predation following prey enrichment in a multiprey system is essential when assessing the effect of predation on an incidental prey species. Our results support the conclusion that wolves are proximately responsible for woodland caribou population declines throughout much of their range.

The effects of puma prey selection and specialization on less abundant prey in Patagonia

Populations of generalist foragers may in fact be composed of individuals that select different prey. We monitored 9 pumas (Puma concolor) in Chilean Patagonia using Argos–global positioning system (Argos-GPS) technology for a mean of 9.33 months 6 5.66 SD. We investigated 694 areas where puma location data were spatially aggregated, called GPS clusters, at which we identified 433 kill sites and 6 acts of scavenging. Pumas as a population specialized upon guanacos (Lama guanicoe), whereas only 7 of 9 individual pumas specialized upon guanacos. One puma specialized upon domestic sheep (Ovis aries) and 1 upon European hares (Lepus europaeus) in terms of numbers of prey killed. Male and female pumas selected different distributions of prey and pumas exhibited prey selection at both the individual and population level. Three of 9 pumas exhibited selection when we compared individual prey use to prey availability within individual pumas’ home ranges. One puma selected endangered huemul (Hippocamelus bisulcus) and 2 selected sheep. When we compared individual prey use to prey use at the population level, 5 of 9 pumas differed from the population norm. Whereas pumas did not select huemul at the population level, 2 individuals did select huemul. Two individuals also selected domestic sheep, and the influence of these 2 pumas was substantial enough to result in a population-level effect. Our research highlights the need to determine whether pumas exhibit individual foraging variation throughout their range, the extrinsic factors associated with (and possibly influencing) such variation, and how pumas that select rare and less abundant species in multiprey systems impact recovering prey populations.

Separating spatial search and efficiency rates as components of predation risk

Predation risk is an important driver of ecosystems, and local spatial variation in risk can have population-level consequences by affecting multiple components of the predation process. I use resource selection and proportional hazard time-to-event modelling to assess the spatial drivers of two key components of risk--the search rate (i.e. aggregative response) and predation efficiency rate (i.e. functional response)--imposed by wolves (Canis lupus) in a multi-prey system. In my study area, both components of risk increased according to topographic variation, but anthropogenic features affected only the search rate. Predicted models of the cumulative hazard, or risk of a kill, underlying wolf search paths validated well with broad-scale variation in kill rates, suggesting that spatial hazard models provide a means of scaling up from local heterogeneity in predation risk to population-level dynamics in predator-prey systems. Additionally, I estimated an integrated model of relative spatial predation risk as the product of the search and efficiency rates, combining the distinct contributions of spatial heterogeneity to each component of risk.

Foraging strategies by omnivores: are black bears actively searching for ungulate neonates or are they simply opportunistic predators?

Omnivores feed on animals with dynamic distributions and on plants with static distributions. The search tactics they adopt will not only define the risk for the targeted prey, but also for other prey that may be consumed when encountered. The potential impact of omnivores on the dynamics of multi-prey systems thus depends on resource selection and on the tactics used to find their prey. We present an approach that can clarify the foraging decisions of omnivores by combining analyses of habitat selection, local residency time, and interpatch movements. We use this framework to evaluate whether predation by omnivorous black bears on ungulate neonates resulted from an active search or from incidental encounters. We monitored 12 bears, 22 forest-dwelling caribou, and 36 moose during calving seasons. We estimated the spatial patterns in relative occurrence probability of ungulate neonates using Resource Selection Functions (RSFs). We also mapped plant abundance from vegetation surveys. RSF were then built to assess the link between bear distribution and the distribution of these three food types (vegetation, moose calves, caribou fawns). We further evaluated the search tactic used by bears that led to this spatial dependency by exploring patterns of residency times and interpatch movements. Bears did not select areas with a high probability of encounter with neonates, but selected areas with abundant vegetation. Surprisingly, bears displayed shorter residency times in vegetation-rich areas. The selection for vegetation-rich areas was therefore achieved by moving preferentially, but frequently, between areas offering abundant vegetation. Such frequent interpatch movements could result in high rates of fortuitous encounters with neonates, even if bears are not actively searching for them. To mitigate the impacts of forest harvesting on threatened caribou populations, vegetation-rich areas selected by bears (e.g. roadsides) should be segregated from large patches of mature conifer forest suitable for caribou.

Habitat selection by a focal predator (Canis lupus) in a multiprey ecosystem of the northern Rockies

Abstract Large predators respond to land cover and physiography that maximize the likelihood of encountering prey. Using locations from global positioning system–collared wolves (Canis lupus), we examined whether land cover, vegetation productivity or change, or habitat-selection value for ungulate prey species themselves most influenced patterns of selection by wolves in a large, intact multiprey system of northern British Columbia. Selection models based on land cover, in combination with topographical features, consistently outperformed models based on indexes of vegetation quantity and quality (using normalized difference vegetation index) or on selection value to prey species (moose [Alces americanus], elk [Cervus elaphus], woodland caribou [Rangifer tarandus], and Stone's sheep [Ovis dalli stonei]). Wolves generally selected for shrub communities and high diversity of cover across seasons and avoided conifer stands and non-vegetated areas and west aspects year-round. Seasonal selection strategies we...

Building a mechanistic understanding of predation with GPS-based movement data

Quantifying kill rates and sources of variation in kill rates remains an important challenge in linking predators to their prey. We address current approaches to using global positioning system (GPS)-based movement data for quantifying key predation components of large carnivores. We review approaches to identify kill sites from GPS movement data as a means to estimate kill rates and address advantages of using GPS-based data over past approaches. Despite considerable progress, modelling the probability that a cluster of GPS points is a kill site is no substitute for field visits, but can guide our field efforts. Once kill sites are identified, time spent at a kill site (handling time) and time between kills (killing time) can be determined. We show how statistical models can be used to investigate the influence of factors such as animal characteristics (e.g. age, sex, group size) and landscape features on either handling time or killing efficiency. If we know the prey densities along paths to a kill, we can quantify the 'attack success' parameter in functional response models directly. Problems remain in incorporating the behavioural complexity derived from GPS movement paths into functional response models, particularly in multi-prey systems, but we believe that exploring the details of GPS movement data has put us on the right path.

Predator‐mediated Allee effects in multi‐prey systems

Allee effects can have significant consequences for small populations and understanding the causal mechanisms for such effects is important for guiding conservation actions. One proposed mechanism is through predation, in which a type II functional response leads to increasing predation rates as prey numbers decline. However, models to support this mechanism have incorporated only a single declining prey species in the functional response, which is probably an oversimplification. We reevaluated the potential for predator-mediated Allee effects in a multi-prey system using Holling's disc equation. We also used empirical data on a large herbivore to examine how grouping behavior may influence the potential for predation-mediated Allee effects. Results based on a multi-prey expression of the functional response predict that Allee effects caused by predation on relatively rare secondary prey may not occur because handling time of the abundant prey dominates the functional response such that secondary prey are largely "bycatch". However, a predator-mediated Allee effect can occur if secondary prey live in groups and if, as the population declines, their average group size declines (a relationship seen in several species). In such a case, the rate at which the number of groups declines is less than the rate at which the population declines. Thus the rate at which a predator encounters a group remains relatively stable, but when a predator kills one animal from smaller groups, the predation rate increases. These results highlight the need to evaluate risks associated with potential changes in group size as populations decline.

FACTORS CORRELATED WITH FORAGING BEHAVIOR OF WOLVES IN AND NEAR GLACIER NATIONAL PARK, MONTANA

We examined prey selection, search distance (measured as km traveled/kill), and spatial use of recolonizing wolves (Canis lupus) in a multi-prey system in northwestern Montana, USA, and southeastern British Columbia, Canada, from 1986 to 1996. Our objective was to explore factors affecting these parameters to better understand wolf–prey relationships of recolonizing wolves. Within white-tailed deer (Odocoileus virginianus) winter ranges, wolves selectively killed elk (Cervus elaphus) over deer. Number of wolves (r = 0.67, P = 0.03), year (r = 0.68, P = 0.02), and possibly human hunter-days/elk harvested (r = 0.55, P = 0.08) were positively correlated with variation in proportion of deer killed by wolves annually. Outside of severe winters, white-tailed deer, elk, and moose (Alces alces) appeared to be equally vulnerable to wolf predation. Search distance of wolves varied by up to 12 times annually. Snow depth (r = 0.73, P = 0.03) and proportion of total kills by wolves that were deer (r = 0.66, P = 0.06) were negatively correlated with the annual variation in the total search distance of wolves. Search distance per wolf was correlated negatively with year (r = 0.66, P = 0.06) and exponentially with hunter-days/elk harvested (r = 0.70, P = 0.04). Space use by wolves may have been in response to local changes in deer abundance. Wolves appeared to select the most profitable prey species. Severe winters and wolf selection for deer, coinciding with a decrease in elk numbers, increased wolf hunting efficiency by reducing search distance. Further research is needed to determine whether reduced search distance equates to increased kill rates by wolves in this system. Based on the time, expense, and difficulty of gathering data on wolf search distance in this sytem, however, we recommend against assessing impacts of wolves on prey via measuring kill rate. Rather, we suggest monitoring impacts of recolonizing wolves by directly assessing cause-specific mortality and recruitment rates of prey species.