Evidence for top predator control of a grazing ecosystem

The importance of top predators in controlling ecological processes in large, intact ecosystems is unclear. In grasslands that support abundant ungulates, top–down control by predators may be particularly important, because of the tight biogeochemical linkages of ungulate prey with plants and soil microbes. Here, I examined the effects of the recent reintroduction of the gray wolf Canis lupus on ecosystem processes in Yellowstone National Park, where herds of grazing ungulates previously have been shown to stimulate several processes, including soil net nitrogen (N) mineralization. Rates of ungulate grazing intensity and soil net N mineralization were compared before and after wolf reintroduction in grasslands ranging five‐fold in aboveground production. Grazing intensity and grassland net N mineralization declined after wolf reintroduction, a likely partial function of fewer ungulates; wolf predation has been one of several factors implicated in causing the decline in Yellowstone ungulates. In addition, the spatial pattern of grazing and net N mineralization changed after reintroduction. A shift in the spatial patterns of grazer‐associated processes is consistent with a growing body of work indicating that wolves have changed habitat use patterns of ungulates in Yellowstone National Park. These findings suggest widespread wolf effects on ungulate prey, plants, and microbial activity that have spatially reorganized grassland energy and nutrient dynamics in Yellowstone Park.     

Linking alternative food sources to winter habitat selection of herbivores in overbrowsed landscapes

During winter, ungulates in boreal forests must cope with high energetic costs related to locomotion in deep snow and reduced forage abundance and quality. At high density, ungulates face additional constraints, because heavy browsing reduces availability of woody browse, the main source of forage during winter. Under these severe conditions, large herbivores might forage on alternative food sources likely independent of browsing pressure, such as litterfall or windblown trees. We investigated the influence of alternative food sources on winter habitat selection, by studying female white-tailed deer (Odocoileus virginianus) living in 2 landscapes with contrasted browse abundance, recently logged and regenerated landscapes, in a population at high density and on a large island free of predators. We fitted 21 female white-tailed deer with Global Positioning System (GPS) collars and delineated winter home ranges and core areas. We measured snow conditions in different habitat categories and sampled vegetation in the core areas and in the rest of the home ranges to determine how forage abundance, protective cover, and snow conditions influenced habitat selection within the home range. In both landscapes, deer were less likely to use open habitat categories as snow accumulated on the ground. At a finer scale, deer inhabiting the regenerated landscape intensively used areas where balsam fir cover was intermediate with greater balsam fir browse density than in the rest of the home range. In the recently logged landscape, deer were more likely to be found near edges between clear-cuts and balsam fir stands and in areas where windblown balsam fir trees were present; the latter being the most influential variable. Although balsam fir browse was sparse and mainly out of reach in this landscape, deer increased the use of areas where it was present. Our results offer novel insights into the resource selection processes of northern ungulates, as we showed that access to winter forage, such as woody browse and alternative food sources, depends on climatic conditions and stochastic events, such as abundant compacted snow or windthrows. To compensate for these scarce and unpredictable food supplies, deer selected habitat categories, but mostly areas within those habitat categories, where the likelihood of finding browse, litterfall, and windblown trees was greatest. © 2011 The Wildlife Society.     

Contrasting Effects of Wolves and Human Hunters on Elk Behavioral Responses to Predation Risk

ABSTRACT Prey behavioral responses to predation risk in wolf-ungulate-plant systems are of interest to wildlife managers. Using Global Positioning System data collected from telemetry-collared elk (Cervus elaphus) and wolves (Canis lupus), we evaluated elk behavioral responses to spatial and temporal variation in wolf- and human-predation risk on a winter range in the Greater Yellowstone Area, USA. We found elk changed grouping patterns and increased movement rates as predation risk increased and that these behavioral changes were habitat dependent. Elk behavioral responses to wolf- and human-predation risk were similar; however, responses to human-predation risk were stronger than responses to wolf-predation risk. These results suggest that predation risk from wolves or human hunters may result in elk spending more time on private rangelands away from public-land winter ranges, which may exacerbate problems of landowner tolerance of elk on livestock pastures. However, increased movement and changing grouping patterns on winter ranges may also disperse elk grazing impacts and lessen elk impacts on any one area.     

Impacts of an invasive willow (Salix × rubens) on riparian bird assemblages in south‐eastern Australia

We explored how a woody plant invader affected riparian bird assemblages. We surveyed 15 200‐m‐long transects in riparian zones in a much‐changed landscape of eastern Victoria, Australia. Abundance, species‐richness, foraging‐guild richness and composition of birds were compared in transects in three habitat types: (i) riparian zones dominated by the invasive willow Salix × rubens; (ii) riparian zones lined with native woody species; and (iii) riparian zones cleared of almost all woody vegetation. We also measured abundance and richness of arthropods and habitat structure to explore further the effects of food resources and habitat on the avifauna. We observed 67 bird species from 14 foraging guilds. Native riparian transects had more birds, bird species and foraging guilds than willow‐invaded or cleared transects. Habitat complexity increased from cleared to willow‐invaded to native riparian transects, as did abundance of native and woodland‐dependent birds. Native shrub and trees species had more foliage and branch‐associated arthropods than did willows, consistent with a greater abundance and variety of foraging guilds of birds dependent on this resource. Willow spread into cleared areas is unlikely to facilitate greatly native bird abundance and diversity even though habitat complexity is increased. Willow invasion into the native riparian zone, by decreasing food resources and altering habitat, is likely to reduce native bird biodiversity and further disrupt connectivity of the riparian zone.     

Landscape of fear in Europe: wolves affect spatial patterns of ungulate browsing in Białowieża Primeval Forest,Poland

Large carnivores can either directly influence ungulate populations or indirectly affect their behaviour. Knowledge from European systems, in contrast to North American systems, on how this might lead to cascading effects on lower trophic levels is virtually absent. We studied whether wolves Canis lupus via density‐mediated and behaviorally‐mediated effects on their ungulate prey species influence patterns of browsing and tree regeneration inside the Bia?owie?a National Park, Poland. Browsing intensity of tree saplings (height class < 150 cm), irrespective of tree species or forest type, was lower inside a wolf core area (50.5%) where predator presence is highest, than in the remainder of the wolf pack’s home range (58.3%). Additionally, browsing intensity was reduced when the amount of coarse woody debris (CWD), which can act as a ?ungulate escape impediment?, increased (within 5‐m radius) inside the wolf core area. No relationship existed outside the core area. As a result, the proportion of trees growing out of herbivore control increased more strongly with increasing amount of CWD inside compared to outside the wolf core area. This suggests that next to direct effects of wolves on ungulate density caused by a higher predation pressure inside the core area, risk effects are important and are enhanced by habitat characteristics. These results indicate that behaviorally‐mediated effects of predators on prey can become more important than density‐mediated effects in affecting lower trophic levels. This is the first study we are aware of, that shows CWD can create fine‐scale risk effects on ungulates with the potential for cascading effects of large predators on patterns of tree regeneration for a European forest system. This knowledge broadens the discussion on how the impact of large predators on ecosystem functioning depends on the physical landscape, by illustrating these effects for a system which largely contrasts in this respect to the North American systems.     

Winter predation patterns of wolves in Northwestern Wyoming

Wolf (Canis lupus) diets and potential effects on prey have been a prominent subject of interest to wildlife researchers and managers since reintroduction into Yellowstone National Park, Wyoming, USA, in 1995 and 1996. Post-reintroduction, wolves expanded south and recolonized areas in the southern Yellowstone ecosystem. Elk (Cervus elaphus) in this area are supplementally fed during winter (Dec–Mar) at state-managed feedgrounds, resulting in high-density congregations of elk. From December to March 2000–2007, we determined the winter predation patterns of wolves by examining the remains of 289 wolf kills on 3 state-managed feedgrounds and adjacent winter range near Jackson, Wyoming. During winters 2002–2005, we also monitored the movements of radio-collared elk on feedgrounds to describe the response of elk to the presence of wolf kills. Thirty-seven percent (n = 106) of kills were located on elk feedgrounds where elk composition included 49% calves, 42% adult females, 5% adult males, and 5% unknown. Sixty-three percent (n = 183) of kills were located on winter range adjacent to feedgrounds and prey species consisted of 90% elk (38% calves, 35% adult females, 24% adult males, 2% unknown), 9% moose (Alces alces; 13% calves, 69% adult females, 6% adult males, 1% unknown), 1% mule deer (Odocoileus hemionus; 1 fawn, 1 adult female), and 0.5% adult female bison (Bison bison). Mean age of elk killed on feedgrounds was 4.2 years (range = 0–20) and 4.6 years (range = 0–23) on winter range. Calves were selected more than available in most years with female elk killed less than expected. Adult males were killed more than expected in 2005–2007. Eighty-eight percent (n = 198) of the time elk remained on the feedground even when wolves made a kill. Less commonly, elk left the feedground, gathered in larger herds on adjacent feedgrounds absent of wolves, and returned within a few days (6%, n = 13) or left the feedground for another feedground and did not return for the rest of the winter (6%; n = 14). Elk were less likely to leave feedgrounds in the presence of a wolf kill when there were more elk on that feedground. Elk left feedgrounds with greater topography and tree cover (Alkali and Fish Creek) and gathered on the flat, open feedgrounds (Patrol Cabin) more frequently than they left flat, open feedgrounds for feedgrounds with greater topography and tree cover. Our results indicate wolves in our study area primarily preyed on elk and exhibited a strong preference for elk calves. High-density concentrations of elk on feedgrounds will continue to be an attractant for wolves. Although elk leave feedgrounds for reasons other than wolf presence, any displacement of elk from feedgrounds due to wolves will be temporary. State managers have the ability to alter management strategies (e.g., increasing wolf harvest, phasing out elk feeding, increasing the intensity of elk feeding) in an effort to affect predator-prey relationships. © 2019 The Wildlife Society.     

Fine-scale predation risk on elk after wolf reintroduction in Yellowstone National Park, USA

While patterns from trophic cascade studies have largely focused on density-mediated effects of predators on prey, there is increasing recognition that behaviorally mediated indirect effects of predators on prey can, at least in part, explain trophic cascade patterns. To determine if a relationship exists between predation risk perceived by elk (Cervus elaphus) while browsing and elk position within the landscape, we observed a total of 56 female elk during two summers and 29 female elk during one winter. At a fine spatial (0–187 m) and temporal scale (145–300 s), results from our model selection indicated summer vigilance levels were greater for females with calves than for females without calves, with vigilance levels greater for all females at closer escape-impediment distances. Winter results also suggested greater female vigilance levels at closer escape-impediment distances, but further indicated an increase in vigilance levels with closer conifer-edge distances. Placed within the context of other studies, the results were consistent with a behaviorally mediated trophic cascade and provide a potential mechanism to explain the variability in observed woody plant release from browsing in Yellowstone National Park, Wyoming, USA.     

Effects of Snow on Sitka Black-Tailed Deer Browse Availability and Nutritional Carrying Capacity in Southeastern Alaska

ABSTRACT Snow affects the nutritional ecology of northern ungulates during winter through burial of important winter forages. We used nonlinear regression analyses to model snow-burial dynamics of blueberry (Vaccinium spp.) browse biomass, a key winter food item of Sitka black-tailed deer (Odocoileus hemionus sitchensis) in southeastern Alaska, USA. During November 2003—March 2004 we collected data from 546 individually marked twigs located on 100 plants of differing sizes and architectures across a range of snow depths. In general, browse biomass became buried and unavailable to deer at snow depths substantially lower than prewinter twig heights. Plant architecture and plant height were related to the probability of a twig being buried. Probability of twig burial was higher on plants with lateral than on those with erect architectures. Twig height also affected the probability of burial by snow but the relationship was complex. For twigs located on erect plants, probability of burial was greatest for twigs near the bottom and top of the plant due to ground-up burial and bending of flexible apex stems, respectively. We used estimated nonlinear equations to model blueberry browse availability in a simulated upland old-growth habitat patch subject to a range of snow depths. We then compared subsequent estimates of deer winter nutritional carrying capacity for this habitat patch to findings derived using an alternative, simple linear (ground-up) model of winter-browse burial by snow. Comparisons indicated that ground-up models of browse burial overestimated browse availability and nutritional carrying capacity for most snow depths. Our findings demonstrate the importance of applying detailed snow-burial models when characterizing nutritional landscape of northern ungulates during winter.     

Optimal predator management for mountain sheep conservation depends on the strength of mesopredator release

Large predators often suppress ungulate population growth, but they may also suppress the abundance of smaller predators that prey on neonatal ungulates. Antagonistic interactions among predators may therefore need to be integrated into predator–prey models to effectively manage ungulate–predator systems. We present a modeling framework that examines the net impact of interacting predators on the population growth rate of shared prey, using interactions among wolves Canis lupus, coyotes Canis latrans and Dall sheep Ovis dalli dalli as a case study. Wolf control is currently employed on approximately 16 million ha in Alaska to increase the abundance of ungulates for human harvest. We hypothesized that the positive effects of wolf control on Dall sheep population growth could be counteracted by increased levels of predation by coyotes. Coyotes and Dall sheep adult females (ewes) and lambs were radiocollared in the Alaska Range from 1999–2005 to estimate fecundity, age‐specific survival rates, and causes of mortality in an area without wolf control. We used stage‐structured population models to simulate the net effect of wolf control on Dall sheep population growth (λ). Our models accounted for stage‐specific predation rates by wolves and coyotes, compensatory mortality, and the potential release of coyote populations due to wolf control. Wolves were the main predators of ewes, coyotes were the main predators of lambs, and wolves were the main source of mortality for coyotes. Population models predicted that wolf control could increase sheep λ by 4% per year in the absence of mesopredator release. However, if wolf control released coyote populations, our models predicted that sheep λ could decrease by up to 3% per year. These results highlight the importance of integrating antagonistic interactions among predators into predator–prey models, because the net effect of predator management on shared prey can depend critically on the strength of mesopredator release.     

Differing influences of natural and artificial disturbances on riparian cottonwoods from prairie to mountain ecoregions in Alberta, Canada

Aim Ecoregions represent biophysical zones where environmental factors enable the development of particular plant communities. Ecoregions are generally large but abrupt transitions occur in areas with rapid physical change. A particularly abrupt transitional sequence occurs in the Rocky Mountain region of south‐western Alberta where fescue prairie, aspen parkland and mountain ecoregions occur within 15 km. To investigate plant adaptation across ecoregions, our study investigated the influences of a natural disturbance (flooding) and an artificial disturbance (cattle grazing) on reproductive and population processes of black cottonwood (Populus balsamifera subsp. trichocarpa, Torr. & Gray), the dominant riparian tree. Location We studied cottonwoods throughout their elevational range along two free‐flowing, first‐order streams, Yarrow and Drywood creeks. Cottonwood was the only prominent tree in the prairie ecoregion, the dominant riparian tree in the parkland and extended upward through the montane ecoregion where it was a pioneer species for the mixed coniferous–deciduous woodland. Cottonwoods did not occur in the higher elevation sub‐alpine ecoregion. Methods Thirty‐six cross‐sectional sampling transects were located across the three ecoregions with cottonwoods, and in ungrazed and grazed areas of each ecoregion. Rectangular 100 m² tree and 2 m² seedling quadrats were positioned along the transects, and substrate and vegetation were assessed. Historic hydrological data were analysed relative to flood recurrences and seasonal flow patterns. Results Overall, the cottonwoods displayed a sawtooth shaped ‘punctuated progressive age structure’ with many young trees, progressively fewer older trees, and about four pulses of increased recruitment over the past century. This was considered to provide a healthy cottonwood population and recruitment pulses were apparently associated with flood events with appropriate peak timing and magnitude and a gradual post‐flood stage recession. However, analyses of tree, sapling and seedling data indicated that flood‐associated seedling recruitment was less important and clonal processes were more important for cottonwood recruitment in the montane ecoregion, the highest ecoregion with cottonwoods. The correlation between flood events and cottonwood recruitment was strongest in the mid‐elevation parkland ecoregion suggesting greater reliance on flood‐associated seedling recruitment. There was little correlation with flooding and limited recruitment in the fescue prairie ecoregion in recent decades and the disturbed age structure probably results from cattle impacts that have prevented recruitment and produced a decrepit cottonwood forest population. Main conclusions These analyses suggested that a healthy cottonwood population displayed a sawtooth shaped ‘punctuated progressive age structure’ and that cottonwood reproduction processes varied across ecoregions with increased clonality in the highest montane ecoregion. Cattle grazing impacts on reproduction were most severe in the lowest prairie ecoregion that is treeless except for the riparian zone. We conclude that appropriate strategies of instream flow regulation, land‐use policies and practices, and conservation and restoration efforts should be refined according to ecoregion to recognize the differences in cottonwood reproductive and population ecology.     

Environmental tolerance of an invasive riparian tree and its potential for continued spread in the southwestern US

Questions: Exotic plant invasion may be aided by facilitation and broad tolerance of environmental conditions, yet these processes are poorly understood in species‐rich ecosystems such as riparian zones. In the southwestern United States (US) two plant species have invaded riparian zones: tamarisk (Tamarix ramosissima, T. chinensis, and their hybrids) and Russian olive (Elaeagnus angustifolia). We addressed the following questions: (1) is Russian olive able to tolerate drier and shadier conditions than cottonwood and tamarisk? (2) Can tamarisk and cottonwood facilitate Russian olive invasion? Location: Arid riparian zones, southwestern US. Methods: We analyzed riparian tree seedling requirements in a controlled experiment, performed empirical field studies, and analyzed stable oxygen isotopes to determine the water sources used by Russian olive. Results: Russian olive survival was significantly higher in dense shade and low moisture conditions than tamarisk and cottonwood. Field observations indicated Russian olive established where flooding cannot occur, and under dense canopies of tamarisk, cottonwood, and Russian olive. Tamarisk and native riparian plant species seedlings cannot establish in these dry, shaded habitats. Russian olive can rely on upper soil water until 15 years of age, before utilizing groundwater. Conclusions: We demonstrate that even though there is little evidence of facilitation by cottonwood and tamarisk, Russian olive is able to tolerate dense shade and low moisture conditions better than tamarisk and cottonwood. There is great potential for continued spread of Russian olive throughout the southwestern US because large areas of suitable habitat exist that are not yet inhabited by this species.     

Linking anti‐predator behaviour to prey demography reveals limited risk effects of an actively hunting large carnivore

Ecological theory predicts that the diffuse risk cues generated by wide‐ranging, active predators should induce prey behavioural responses but not major, population‐ or community‐level consequences. We evaluated the non‐consumptive effects (NCEs) of an active predator, the grey wolf (Canis lupus), by simultaneously tracking wolves and the behaviour, body fat, and pregnancy of elk (Cervus elaphus), their primary prey in the Greater Yellowstone Ecosystem. When wolves approached within 1 km, elk increased their rates of movement, displacement and vigilance. Even in high‐risk areas, however, these encounters occurred only once every 9 days. Ultimately, despite 20‐fold variation in the frequency of encounters between wolves and individual elk, the risk of predation was not associated with elk body fat or pregnancy. Our findings suggest that the ecological consequences of actively hunting large carnivores, such as the wolf, are more likely transmitted by consumptive effects on prey survival than NCEs on prey behaviour.     

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.     

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.     

Botanical and chemical composition of rumen contents of Sika deer on Mt Goyo,northern Japan

Botanical and chemical compositions of the rumen contents of 58 Sika deer on Mt Goyo, northern Japan, collected from summer of 1988 to spring of 1989, were analyzed.Sasa nipponica, a dwarf bamboo, was important in summer (35.0%) and winter (61.4%), but it decreased to 5.6% and was replaced by browse leaves in fall, and to 28.0% and was replaced by dead leaves, twigs and bark of woody plants in winter. Crude protein was 20–25% lower in the washed fraction than in the gross fraction. It was highest (16.2%) in summer and lowest (8.6%) in winter. High protein content in summer and fall foods was attributed to forb and browse leaves. Seasonal fluctuation in protein content in the foods of these Sika deer was greater than red deer on Rhum, Scotland and smaller than wapiti in Yellowstone National Park, Wyoming, USA. Climatically, winter severity of Mt Goyo is intermediate between the two localities, which seems to explain the seasonal fluctuation of the protein level. Crude fiber wasca 33% in the ‘washed’ fraction, and did not change seasonally.     

Do prey select for vacant hunting domains to minimize a multi‐predator threat?

Many ecosystems contain sympatric predator species that hunt in different places and times. We tested whether this provides vacant hunting domains, places and times where and when predators are least active, that prey use to minimize threats from multiple predators simultaneously. We measured how northern Yellowstone elk (Cervus elaphus) responded to wolves (Canis lupus) and cougars (Puma concolor), and found that elk selected for areas outside the high‐risk domains of both predators consistent with the vacant domain hypothesis. This enabled elk to avoid one predator without necessarily increasing its exposure to the other. Our results demonstrate how the diel cycle can serve as a key axis of the predator hunting domain that prey exploit to manage predation risk from multiple sources. We argue that a multi‐predator, spatiotemporal framework is vital to understand the causes and consequences of prey spatial response to predation risk in environments with more than one predator.     

Faunal isotope records reveal trophic and nutrient dynamics in twentieth century Yellowstone grasslands

Population sizes and movement patterns of ungulate grazers and their predators have fluctuated dramatically over the past few centuries, largely owing to overharvesting, land-use change and historic management. We used δ(13)C and δ(15)N values measured from bone collagen of historic and recent gray wolves and their potential primary prey from Yellowstone National Park to gain insight into the trophic dynamics and nutrient conditions of historic and modern grasslands. The diet of reintroduced wolves closely parallels that of the historic population. We suggest that a significant shift in faunal δ(15)N values over the past century reflects impacts of anthropogenic environmental changes on grassland ecosystems, including grazer-mediated shifts in grassland nitrogen cycle processes.     

Gray wolves as climate change buffers in Yellowstone

Understanding the mechanisms by which climate and predation patterns by top predators co-vary to affect community structure accrues added importance as humans exert growing influence over both climate and regional predator assemblages. In Yellowstone National Park, winter conditions and reintroduced gray wolves (Canis lupus) together determine the availability of winter carrion on which numerous scavenger species depend for survival and reproduction. As climate changes in Yellowstone, therefore, scavenger species may experience a dramatic reshuffling of food resources. As such, we analyzed 55 y of weather data from Yellowstone in order to determine trends in winter conditions. We found that winters are getting shorter, as measured by the number of days with snow on the ground, due to decreased snowfall and increased number of days with temperatures above freezing. To investigate synergistic effects of human and climatic alterations of species interactions, we used an empirically derived model to show that in the absence of wolves, early snow thaw leads to a substantial reduction in late-winter carrion, causing potential food bottlenecks for scavengers. In addition, by narrowing the window of time over which carrion is available and thereby creating a resource pulse, climate change likely favors scavengers that can quickly track food sources over great distances. Wolves, however, largely mitigate late-winter reduction in carrion due to earlier snow thaws. By buffering the effects of climate change on carrion availability, wolves allow scavengers to adapt to a changing environment over a longer time scale more commensurate with natural processes. This study illustrates the importance of restoring and maintaining intact food chains in the face of large-scale environmental perturbations such as climate change.     

Trophic niche overlap and wild ungulate consumption by red fox and wolf in a mountain area in Italy

Red fox (Vulpes vulpes) and wolf (Canis lupus) are two widespread opportunistic predators living in simpatry in many areas. Nonetheless, scarce information are available on their trophic interactions. We investigated food habits of these two carnivores in a mountain area in Italy and assessed the extent of their trophic niche overlap, focusing on the consumption of wild ungulates. Thereby we analyzed the content of 669 red fox scats and 253 wolf scats collected between May 2008 and April 2009. Red foxes resulted to have a more than three times higher niche breadth than wolves. Vegetables, small mammals, wild ungulates, and invertebrates were major items (altogether 92% of volume) of the red fox annual diet. On the contrary wolf annual diet relied on wild ungulates (94% of volume) with wild boar (Sus scrofa) being the main food item. The degree of trophic niche overlap between the two species was found to be low (Pianka's O = 0.356). Diet variation between the warm and the cold seasons was limited in both species, and higher in red fox than in wolf. The two canids appeared to use wild ungulates unevenly being the former more selective for younger preys, smaller in size (newborn piglets and roe deer Capreolus capreolus fawns), whereas the latter exhibited a preference for medium-sized and large ungulates (10–35 kg wild boar and adult roe deer). Even if wild ungulates represent the main shared food category, the different use of age/weight classes by the two predators, together with their possible consumption as carrions by red fox, suggests a very limited trophic competition between wolf and red fox.This study represents a contribution to the knowledge of trophic interaction in predator–prey systems where sympatric carnivores are present.     

The cost of migratory prey: seasonal changes in semi‐domestic reindeer distribution influences breeding success of Eurasian lynx in northern Norway

Migratory prey is a widespread phenomenon that has implications for predator–prey interactions. By creating large temporal variation in resource availability between seasons it becomes challenging for carnivores to secure a regular year‐round supply of food. Some predators may respond by following their migratory prey, however, most predators are sedentary and experience strong seasonal variation in resource availability. Increased predation on alternative prey may dampen such seasonal resource fluctuations, but reduced reproduction rates in predators is a predicted consequence of migratory primary prey behavior that has received little empirical attention. We used data from 23 GPS collared Eurasian lynx Lynx lynx monitored during 2007–2013 in northern Norway, to examine how spatio‐temporal variation in the migratory behavior of semi‐domestic reindeer Rangifer tarandus influences lynx spatial organization and reproductive success using estimates of seasonal home range overlap and breeding success. We found that lynx of both sexes maintained seasonally stable home ranges and exhibited site fidelity across years, independent of whether they had access to reindeer throughout the year or experienced a scarcity of reindeer in winter due to migration. However, lynx without access to reindeer in winter showed a decreased probability of reproducing and a tendency for lowered kitten survival into their first winter, when compared to female lynx with reindeer available year around. This supports the hypothesis that sedentary predators experience demographic costs in systems with migratory primary prey. Changes in the migratory behavior of ungulates, including disrupted migrations, is therefore likely to have bottom–up effects on the population dynamics of sedentary predators as well as the previously documented consequences for ungulate population dynamics.