The Behavior and Ecology of Cursorial Predators and Dangerous Prey: Integrating Behavioral Mechanisms with Population-level Patterns in Large Mammal Systems

Driving into Yellowstone National Park for the first time is a moving experience. Gazing over the sweeping landscapes, seeing a geyser erupt 80 feet into the air, and having your first ‘wildlife encounter’, whether that be a 2 ton bull bison aggressively wallowing on his dirt mound, snorting and kicking up dust, or watching a pack of 6 wolves move through a valley off in the distance, pausing to howl in search of their companions. Yellowstone staff wishes to manage our park in a way that preserves these remarkable experiences. In order to effectively manage this dynamic ecosystem, it is critical to thoroughly understand how different animal and plant species interact with each other and their environment. Wolves were reintroduced to Yellowstone in 1995-1997 and park researchers and managers are still trying to understand how their presence impacts the ecosystem. In Yellowstone, wolves primarily prey on elk; however, predation on bison has started to increase in recent years. We still know little about how wolves hunt bison and what impacts wolves have had on how bison use their environment. The objective of this study was to better understand the behavioral and ecological interactions of wolves and bison, the most dangerous prey for wolves in North America. Since reintroduction, researchers have collected data on how wolves hunt both elk and bison. I used these data to understand 1) the conditions that allow wolves to capture their most dangerous prey, bison, 2) whether wolves have started preying on bison more often as the bison population increased, and 3) whether wolf reintroduction has limited bison use of Yellowstone’s most extreme high-elevation winter range. Finally, I collaborated with ecologists in Scandinavia to determine how wolf predation was affected by a competitor, the brown bear. My study adds to the current body of work addressing the effects of wolf reintroduction in Yellowstone. This research is unique because it focuses on wolf bison interactions, about which little is known in this system. This research also sheds light on the behavioral relationships at play in a special type of predator-prey interaction: predators that hunt dangerous prey

Threatened and Invasive Reptiles Are Not Two Sides of the Same Coin

The ‘two sides of the same coin’ hypothesis posits that biological traits that predispose species to extinction and invasion lie on opposite ends of a continuum. Conversely, anthropogenic factors may have similar effects on extinction and invasion risk. We test these two hypotheses using data on more than 1000 reptile species

Scavenging patterns of an inbred wolf population in a landscape with a pulse of human-provided carrion

Scavenging is an important part of food acquisition for many carnivore species that switch between scavenging and predation. In landscapes with anthropogenic impact, humans provide food that scavenging species can utilize. We quantified the magnitude of killing versus scavenging by gray wolves (Canis lupus) in Scandinavia where humans impact the ecosystem through hunter harvest, land use practices, and infrastructure. We investigated the cause of death of different animals utilized by wolves, and examined how the proportion of their consumption time spent scavenging was influenced by season, wolf social affiliation, level of inbreeding, density of moose (Alces alces) as their main prey, density of brown bear (Ursus arctos) as an intraguild competitor, and human density. We used data from 39 GPS-collared wolves covering 3198 study days (2001-2019), including 14,205 feeding locations within space-time clusters, and 1362 carcasses utilized by wolves. Most carcasses were wolf-killed (80.5%) while a small part had died from other natural causes (1.9%). The remaining had either anthropogenic mortality causes (4.7%), or the cause of death was unknown (12.9%). Time spent scavenging was higher during winter than during summer and autumn. Solitary wolves spent more time scavenging than pack-living individuals, likely because individual hunting success is lower than pack success. Scavenging time increased with the mean inbreeding coefficient of the adult wolves, possibly indicating that more inbred individuals resort to scavenging, which requires less body strength. There was weak evidence for competition between wolves and brown bears as well as a positive relationship between human density and time spent scavenging. This study shows how both intrinsic and extrinsic factors drive wolf scavenging behavior, and that despite a high level of inbreeding and access to carrion of anthropogenic origin, wolves mainly utilized their own kills

Scavenging patterns of an inbred wolf population in a landscape with a pulse of human-provided carrion

Scavenging is an important part of food acquisition for many carnivore species that switch between scavenging and predation. In landscapes with anthropogenic impact, humans provide food that scavenging species can utilize. We quantified the magnitude of killing versus scavenging by gray wolves (Canis lupus) in Scandinavia where humans impact the ecosystem through hunter harvest, land use practices, and infrastructure. We investigated the cause of death of different animals utilized by wolves, and examined how the proportion of their consumption time spent scavenging was influenced by season, wolf social affiliation, level of inbreeding, density of moose (Alces alces) as their main prey, density of brown bear (Ursus arctos) as an intraguild competitor, and human density. We used data from 39 GPS-collared wolves covering 3198 study days (2001–2019),including 14,205 feeding locations within space–time clusters, and 1362 carcasses utilized by wolves. Most carcasses were wolf-killed (80.5%) while a small part had died from other natural causes (1.9%). The remaining had either anthropogenic mortality causes (4.7%), or the cause of death was unknown (12.9%). Time spent scavenging was higher during winter than during summer and autumn. Solitary wolves spent more time scavenging than pack-living individuals, likely because individual hunting success is lower than pack success. Scavenging time increased with the mean inbreeding coefficient of the adult wolves, possibly indicating that more inbred individuals resort to scavenging, which requires less body strength. There was weak evidence for competition between wolves and brown bears as well as a positive relationship between human density and time spent scavenging. This study shows how both intrinsic and extrinsic factors drive wolf scavenging behavior, and that despite a high level of inbreeding and access to carrion of anthropogenic origin, wolves mainly utilized their own kills. Canis lupus, consumption time, human density, inbreeding, intraguild competition, prey density, social affiliationpublishedVersio

Brown bear predation on semi-domesticated reindeer and depredation compensations

The recovery and conservation of large carnivores can negatively impact the economy of traditional pastoralist societies, including indigenous reindeer (Rangifer tarandus) herding communities. Quantifying the magnitude of predation on livestock is critical to evaluating governmental carnivore compensation schemes. We collaborated with two Sami herding communities in northern Sweden (2010-2012) to examine brown bear (Ursus arctos) predation patterns on semi-domesticated reindeer and quantify the economic impact of bear predation. Predation patterns were estimated by following 21 GPS proximity-collared bears and similar to 2500 transmitter-collared female reindeer during calving season. We calculated economic impact by multiplying the monetary value of reindeer by the expected number lost to bears. On average, bears killed 10.2 [8.6, 11.5] calves per bear, accounting for 39-62% of all calf mortality, while few adult reindeer were killed. Bear kill rates increased with time spent in the calving area, and varied widely by individual and reproductive status, e.g., females with cubs-of-the-year did not kill calves. Kill intervals increased over the parturition season, and were larger for sub-adults than adults. The mean reindeer calf predation rate was 16-27%, which resulted in an annual loss between similar to(sic)50,000 and similar to(sic)62,000 per herding group. Current compensation schemes for herding communities in Sweden are calculated as a fixed rate based on herding community land-area. The herding groups in our study were reimbursed for similar to 2% of realized monetary loss. Compensation schemes based on herding community area, rather than realized predation patterns, may be less effective at mitigating the economic impact of living with large carnivores

Large carnivore conservation and traditional pastoralism: A case study on bear–reindeer predation mitigation measures

While wildlife and cultural preservation goals can be either complimentary or counteractive, the goals of large carnivore conservation and traditional pastoralist lifestyles are often at odds. Livestock depredation can negatively impact the economies of livestock herders, while subsequent lethal removals contribute to local carnivore population declines. Here, we collaborated with two Sámi reindeer herding communities (2010–2016) situated in Sweden's boreal forest to evaluate the efficacy and economic feasibility of three brown bear predation mitigation measures: corralling pregnant reindeer during parturition, lethal bear management removals, and public bear-license hunting. Calving corrals increased survival for reindeer calves born to average-sized females by 7%–15%, and by 14%–30% for calves born to small females. However, the realized cost of implementing calving corrals outweighed the financial gain for both our study areas (net losses ranged between €1111 and €6210 per calf saved from bear predation per year when using the updated 2021 calf value; 1€ [Euro] = US$1.1), as well as for almost every theoretical scenario we explored (net losses €234 and €13,995 per calf saved from bear predation). The exception was the theoretical scenario where small herding communities overlapped large bear populations, which crossed the breakeven efficacy bear/reindeer ratio of 13.5 bears/100 reindeer and had a potential net gain of €36 per saved calf. Similarly, the cost of lethal management removals of bears outweighed the potential financial gain from saved calves, with net losses between €75 and €239 per calf. License hunting, where the hunters voluntarily incur the monetary costs of removing bears, is in most cases the only economically viable mitigation measure where the cost of mitigation did not outweigh the financial gain from increased reindeer survival. While the annual public license hunt was the most cost-effective mitigation measure, it may be less biologically effective, that is, bear hunting occurs in the fall and reindeer parturition the following spring which leaves time for the empty niche of harvested bears to be filled by survivors. Economically and biologically effective predation mitigation measures are key for promoting coexistence, and we suggest that potential mitigation measures should be studied in collaboration with local people

Brown bear predation on semi-domesticated reindeer and depredation compensations

The recovery and conservation of large carnivores can negatively impact the economy of traditional pastoralist societies, including indigenous reindeer (Rangifer tarandus) herding communities. Quantifying the magnitude of predation on livestock is critical to evaluating governmental carnivore compensation schemes. We collaborated with two Sami herding communities in northern Sweden (2010–2012) to examine brown bear (Ursus arctos) predation patterns on semidomesticated reindeer and quantify the economic impact of bear predation. Predation patterns were estimated by following 21 GPS proximity- collared bears and ~2500 transmitter-collared female reindeer during calving season. We calculated economic impact by multiplying the monetary value of reindeer by the expected number lost to bears. On average, bears killed 10.2 [8.6, 11.5] calves per bear, accounting for 39− 62% of all calf mortality, while few adult reindeer were killed. Bear kill rates increased with time spent in the calving area, and varied widely by individual and reproductive status, e.g., females with cubs-of-the- year did not kill calves. Kill intervals increased over the parturition season, and were larger for sub-adults than adults. The mean reindeer calf predation rate was 16–27%, which resulted in an annual loss between ~€50,000 and ~€62,000 per herding group. Current compensation schemes for herding communities in Sweden are calculated as a fixed rate based on herding community land-area. The herding groups in our study were reimbursed for ~2% of realized monetary loss. Compensation schemes based on herding community area, rather than realized predation patterns, may be less effective at mitigating the economic impact of living with large carnivores. Compensation Depredation Economic impact Proximity collars Indigenous communities PastoralismpublishedVersio

Of wolves and bears: Seasonal drivers of interference and exploitation competition between apex predators

This is an open access article under the terms of the Creative Commons Attribution-NonCommercial-NoDerivs License, which permits use and distribution in any medium, provided the original work is properly cited, the use is non-commercial and no modifications or adaptations are made. © 2022 The Authors. Ecological Monographs published by Wiley Periodicals LLC on behalf of Ecological Society of America.Competition between apex predators can alter the strength of top-down forcing, yet we know little about the behavioral mechanisms that drive competition in multipredator ecosystems. Interactions between predators can be synergistic (facilitative) or antagonistic (inhibitive), both of which are widespread in nature, vary in strength between species and across space and time, and affect predation patterns and predator–prey dynamics. Recent research has suggested that gray wolf (Canis lupus) kill rates decrease where they are sympatric with brown bears (Ursus arctos), however, the mechanisms behind this pattern remain unknown. We used data from two long-term research projects in Scandinavia (Europe) and Yellowstone National Park (North America) to test the role of interference and exploitation competition from bears on wolf predatory behavior, where altered wolf handling and search time of prey in the presence of bears are indicative of interference and exploitation competition, respectively. Our results suggest the mechanisms driving competition between bears and wolves were dependent on the season and study system. During spring in Scandinavia, interference competition was the primary mechanism driving decreased kill rates for wolves sympatric with bears; handling time increased, but search time did not. In summer, however, when both bear and wolf predation focused on neonate moose, the behavioral mechanism switched to exploitation competition; search time increased, but handling time did not. Alternartively, interference competition did affect wolf predation dynamics in Yellowstone during summer, where wolves prey more evenly on neonate and adult ungulates. Here, bear presence at a carcass increased the amount of time wolves spent at carcasses of all sizes and wolf handling time for small prey, but decreased handling time for the largest prey. Wolves facilitate scavenging opportunities for bears, however, bears alter wolf predatory behavior via multiple pathways and are primarily antagonistic to wolves. Our study helps to clarify the behavioral mechanisms driving competition between apex predators, illustrating how interspecific interactions can manifest into population-level predation patterns.publishedVersio

Of wolves and bears: Seasonal drivers of interference and exploitation competition between apex predators

Competition between apex predators can alter the strength of top-down forcing, yet we know little about the behavioral mechanisms that drive competition in multipredator ecosystems. Interactions between predators can be synergistic (facilitative) or antagonistic (inhibitive), both of which are widespread in nature, vary in strength between species and across space and time, and affect predation patterns and predator–prey dynamics. Recent research has suggested that gray wolf (Canis lupus) kill rates decrease where they are sympatric with brown bears (Ursus arctos), however, the mechanisms behind this pattern remain unknown. We used data from two long-term research projects in Scandinavia (Europe) and Yellowstone National Park (North America) to test the role of interference and exploitation competition from bears on wolf predatory behavior, where altered wolf handling and search time of prey in the presence of bears are indicative of interference and exploitation competition, respectively. Our results suggest the mechanisms driving competition between bears and wolves were dependent on the season and study system. During spring in Scandinavia, interference competition was the primary mechanism driving decreased kill rates for wolves sympatric with bears; handling time increased, but search time did not. In summer, however, when both bear and wolf predation focused on neonate moose, the behavioral mechanism switched to exploitation competition; search time increased, but handling time did not. Alternartively, interference competition did affect wolf predation dynamics in Yellowstone during summer, where wolves prey more evenly on neonate and adult ungulates. Here, bear presence at a carcass increased the amount of time wolves spent at carcasses of all sizes and wolf handling time for small prey, but decreased handling time for the largest prey. Wolves facilitate scavenging opportunities for bears, however, bears alter wolf predatory behavior via multiple pathways and are primarily antagonistic to wolves. Our study helps to clarify the behavioral mechanisms driving competition between apex predators, illustrating how interspecific interactions can manifest into population-level predation patterns.publishedVersio