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Apex predators play key roles in food webs and their recovery can trigger trophic cascades in some ecosystems. Intra‐guild competition can reduce the abundances of smaller predators and perceived predation risk can alter their foraging behaviour thereby limiting seed dispersal by frugivorous carnivores. However, little is known about how plant–frugivore mutualisms could be disturbed in the presence of larger predators.We evaluated the top‐down effect of the regional superpredator, the Iberian lynx Lynx pardinus, on the number of visits and fruits consumed by medium‐sized frugivorous carnivores, as well as the foraging behaviour of identified individuals, by examining the consumption likelihood and the foraging time.We carried out a field experiment in which we placed Iberian pear Pyrus bourgaeana fruits beneath fruiting trees and monitored pear removal by frugivorous carnivores, both inside and outside lynx ranges. Using camera traps, we recorded the presence of the red fox Vulpes vulpes, the Eurasian badger Meles meles and the stone marten Martes foina, as well as the number of fruits they consumed and their time spent foraging.Red fox was the most frequent fruit consumer carnivore. We found there were fewer visits and less fruit consumed by foxes inside lynx ranges, but lynx presence did not seem to affect badgers. We did not observe any stone marten visits inside lynx territories. The foraging behaviour of red foxes was also altered inside lynx ranges whereby foxes were less efficient, consuming less fruit per unit of time and having shorter visits. Local availability of fruit resources, forest coverage and individual personality also were important variables to understand visitation and foraging in a landscape of fear.Our results show a potential trophic cascade from apex predators to primary producers. The presence of lynx can reduce frugivorous carnivore numbers and induce shifts in their feeding behaviour that may modify the seed dispersal patterns with likely consequences for the demography of many fleshy‐fruited plant species. We conclude that knowledge of the ecological interactions making up trophic webs is an asset to design effective conservation strategies, particularly in rewilding programs. [ABSTRACT FROM AUTHOR]

Scavenging by vertebrates can have important impacts on food web stability and persistence, and can alter the distribution of nutrients throughout the landscape. However, scavenging communities have been understudied in most regions around the globe, and we lack understanding of the biotic drivers of vertebrate scavenging dynamics.In this paper, we examined how changes in prey density and carrion biomass caused by population cycles of a primary prey species, the snowshoe hare Lepus americanus, influence scavenging communities in the northern boreal forest. We further examined the impact of habitat and temperature on scavenging dynamics.We monitored the persistence time, time until first scavenger, and number of species scavenging experimentally‐placed hare carcasses over four consecutive years in the southwestern Yukon. We simultaneously monitored hare density and carrion biomass to examine their influence relative to temperature, habitat, and seasonal effects. For the primary scavengers, we developed species‐specific scavenging models to determine variation on the effects of these factors across species, and determine which species may be driving temporal patterns in the entire community.We found that the efficiency of the scavenging community was affected by hare density, with carcass persistence decreasing when snowshoe hare densities declined, mainly due to increased scavenging rates by Canada lynx Lynx canadensis. However, prey density did not influence the number of species scavenging a given carcass, suggesting prey abundance affects carrion recycling but not necessarily the number of connections in the food web. In addition, scavenging rates increased in warmer temperatures, and there were strong seasonal effects on the richness of the vertebrate scavenging community.Our results demonstrate that vertebrate scavenging communities are sensitive to changes in species' demography and environmental change, and that future assessments of food web dynamics should consider links established through scavenging. [ABSTRACT FROM AUTHOR]

Juvenile survival to first breeding is a key life‐history stage for all taxa. Survival through this period can be particularly challenging when it coincides with harsh environmental conditions such as a winter climate or food scarcity, leading to highly variable cohort survival. However, the small size and dispersive nature of juveniles generally make studying their survival more difficult.In territorial species, a key life‐history event is the acquisition of a territory. A territory is expected to enhance survival, but how it does so is not often identified. We tested how the timing of territory acquisition influenced the winter survival of juvenile North American red squirrels Tamiasciurus hudsonicus, hereafter red squirrels, and how the timing of this event mediated the sources of mortality. We hypothesized that securing a territory prior to when food resources become available would reduce juvenile susceptibility to predation and climatic factors overwinter.Using 27 years of data on the survival of individually marked juvenile red squirrels, we tested how the timing of territory acquisition influenced survival, whether the population density of red squirrel predators and mean temperature overwinter were related to individual survival probability, and if territory ownership mediated these effects.Juvenile red squirrel survival was lower in the years of high predator abundance and in colder winters. Autumn territory owners were less susceptible to lynx Lynx canadensis and possibly mustelid Mustela and Martes spp., predation. Autumn territory owners had lower survival in colder winters, but surprisingly non‐owners had higher survival in cold winters.Our results show how the timing of a life‐history event like territory acquisition can directly affect survival and also mediate the effects of biotic and abiotic factors later in life. This engenders a better understanding of the fitness consequences of the timing of key life‐history events. [ABSTRACT FROM AUTHOR]

Restricting movements to familiar areas should increase individual fitness as it provides animals with information about the spatial distribution of resources and predation risk. While the benefits of familiarity for locating resources have been reported previously, the potential value of familiarity for predation avoidance has been accorded less attention. It has been suggested that familiarity should be beneficial for anti‐predator behaviour when direct cues of predation risk are unclear and do not allow prey to identify well‐defined spatial refuges. However, to our knowledge, this hypothesis has yet to be tested.Here, we assessed how site familiarity, measured as the intensity of use of a given location, is associated with the probability of roe deer Capreolus capreolus being killed by two predators with contrasting hunting tactics, the Eurasian lynx Lynx lynx and human hunters. While risk of human hunting was confined to open habitats, risk of lynx predation was more diffuse, with no clear refuge areas.We estimated cause‐specific mortality rates in a competing risk framework for 212 GPS‐collared roe deer in two ecologically distinct areas of Central Europe to test the hypothesis that the daily risk of being killed by lynx or hunters should be lower in areas of high familiarity.We found strong evidence that site familiarity reduces the risk of being predated by lynx, whereas the evidence that the risk of being hunted is linked to site familiarity was weak.We suggest that local knowledge about small‐scale differences in predation risk and information about efficient escape routes affect an individual's ability to avoid or escape an attack by an ambush predator. Our study emphasizes the role of site familiarity in determining the susceptibility of prey to predation. Further research will be required to understand better how a cognitive map of individual spatial information is beneficial for avoiding predation in the arms race that drives the predator–prey shell game. [ABSTRACT FROM AUTHOR]

Humans, as super predators, can have strong effects on wildlife behaviour, including profound modifications of diel activity patterns. Subsequent to the return of large carnivores to human‐modified ecosystems, many prey species have adjusted their spatial behaviour to the contrasting landscapes of fear generated by both their natural predators and anthropogenic pressures. The effects of predation risk on temporal shifts in diel activity of prey, however, remain largely unexplored in human‐dominated landscapes.We investigated the influence of the density of lynx Lynx lynx, a nocturnal predator, on the diel activity patterns of their main prey, the roe deer Capreolus capreolus, across a gradient of human disturbance and hunting at the European scale.Based on 11 million activity records from 431 individually GPS‐monitored roe deer in 12 populations within the EURODEER network (http://eurodeer.org), we investigated how lynx predation risk in combination with both lethal and non‐lethal human activities affected the diurnality of deer.We demonstrated marked plasticity in roe deer diel activity patterns in response to spatio‐temporal variations in risk, mostly due to human activities. In particular, roe deer decreased their level of diurnality by a factor of 1.37 when the background level of general human disturbance was high. Hunting exacerbated this effect, as during the hunting season deer switched most of their activity to night‐time and, to a lesser extent, to dawn, although this pattern varied noticeably in relation to lynx density. Indeed, in the presence of lynx, their main natural predator, roe deer were relatively more diurnal. Overall, our results revealed a strong influence of human activities and the presence of lynx on diel shifts in roe deer activity.In the context of the recovery of large carnivores across Europe, we provide important insights about the effects of predators on the behavioural responses of their prey in human‐dominated ecosystems. Modifications in the temporal partitioning of ungulate activity as a response to human activities may facilitate human–wildlife coexistence, but likely also have knock‐on effects for predator–prey interactions, with cascading effects on ecosystem functioning. [ABSTRACT FROM AUTHOR]

Apex predators may buffer bottom-up driven ecosystem change, as top-down suppression may dampen herbivore and mesopredator responses to increased resource availability. However, theory suggests that for this buffering capacity to be realized, the equilibrium abundance of apex predators must increase. This raises the question: will apex predators maintain herbivore/mesopredator limitation, if bottom-up change relaxes resource constraints?, Here, we explore changes in mesopredator (red fox Vulpes vulpes) abundance over 220 years in response to eradication and recovery of an apex predator (Eurasian lynx Lynx lynx), and changes in land use and climate which are linked to resource availability., A three-step approach was used. First, recent data from Finland and Sweden were modelled to estimate linear effects of lynx density, land use and winter temperature on fox density. Second, lynx density, land use and winter temperature was estimated in a 22 650 km2 focal area in boreal and boreo-nemoral Sweden in the years 1830, 1920, 2010 and 2050. Third, the models and estimates were used to project historic and future fox densities in the focal area., Projected fox density was lowest in 1830 when lynx density was high, winters cold and the proportion of cropland low. Fox density peaked in 1920 due to lynx eradication, a mesopredator release boosted by favourable bottom-up changes - milder winters and cropland expansion. By 2010, lynx recolonization had reduced fox density, but it remained higher than in 1830, partly due to the bottom-up changes. Comparing 1830 to 2010, the contribution of top-down limitation decreased, while environment enrichment relaxed bottom-up limitation. Future scenarios indicated that by 2050, lynx density would have to increase by 79% to compensate for a projected climate-driven increase in fox density., We highlight that although top-down limitation in theory can buffer bottom-up change, this requires compensatory changes in apex predator abundance. Hence apex predator recolonization/recovery to historical levels would not be sufficient to compensate for widespread changes in climate and land use, which have relaxed the resource constraints for many herbivores and mesopredators. Variation in bottom-up conditions may also contribute to context dependence in apex predator effects. [ABSTRACT FROM AUTHOR]

Lack proposed that the average clutch size of altricial species should be determined by the average maximum number of young the parents can raise such that all females in a given population should share a common optimal clutch size. Support for this model remains equivocal and recent studies have suggested that intra-population variation in clutch size is adaptive because each female has its own optimal clutch size associated with its intrinsic ability to raise offspring., Although Lack litter size and condition-dependent litter size are presented as two competing models, both are based on the concept of individual optimization. We propose a unified optimal litter size model (called 'adaptive litter size') and identify a set of conditions under which a common vs. a state-dependent optimal litter size should be observed., We test whether females of Eurasian lynx ( Lynx lynx) have a common optimal litter size, or whether they adjust their litter size according to their state. We used a detailed individual-based data set collected from contrasting populations of Eurasian lynx in Scandinavia., Observed reproductive patterns in female lynx provide strong support for the existence of a common optimal litter size. Litter size did not vary according to female body mass or reproductive category, or among contrasted populations and years. A litter size of 2 was associated with a higher fitness than both smaller and larger litters, and thus corresponded to the 'adaptive litter size' for female lynx., We suggest that the reproductive pattern of female lynx might correspond to a risk avoidance tactic common to all individuals, which has evolved in response to strong environmental constraints generated by a highly unpredictable food supply during lactation. [ABSTRACT FROM AUTHOR]

1. Predation plays a major role in shaping the structure and dynamics of ecological communities, and the functional response of a predator is of crucial importance to the dynamics of any predator–prey system by linking the trophic levels. For large mammals, there is a dearth of field studies documenting functional responses, and observations at low prey density are particularly scarce. Furthermore, there is a lack of understanding about how variables such as season, social status and climate modulate the functional response curves. 2. We analysed kill rate data collected over a 10-year period based on radio-marked lynx ( Lynx lynx) mainly preying on roe deer ( Capreolus capreolus) along a steep prey density gradient in south-eastern Norway. 3. The asymptotic kill rate was reached at a very low prey density for both solitary individuals and family groups (i.e. females with their dependent kittens), indicative of an efficient predator. This highlights the importance of understanding the interplay between predator and prey at low prey densities. 4. A purely prey-dependent functional response was a poor descriptor of the data, as the curve was strongly modulated by season and differences between lynx of different social status. In addition, there was a clear effect of abiotic climatic factors (indexed by the North Atlantic Oscillation) on observed kill rates in the more snow-rich portion of our study area. 5. Our analysis suggests that simple functional response curves might be poor descriptors of predator consumption rates in complex natural system, and that auxiliary factors are likely to induce complexity into any predator–prey systems that would not be captured by simple deterministic approaches. [ABSTRACT FROM AUTHOR]

1. Environmental conditions shape population growth through their impact on demographic parameters. While knowledge has accumulated concerning the effects of population density and climatic conditions, a topical question now concerns how predation and harvest influence demographic parameters and population growth (λ). 2. We performed a comparative demographic analysis based on projection matrix models for female roe deer. Population-specific matrices were parameterized based on longitudinal data from five intensively monitored populations in Norway and France, spanning a large variability in environmental characteristics such as densities of large predators, hunter harvest and seasonality. 3. As expected for a large iteroparous vertebrate, temporal variation was invariably higher in recruitment than in adult survival, and the elasticity of adult survival was consistently higher than that of recruitment. However, the relative difference in elasticity of λ to recruitment and adult survival varied strongly across populations, and was closely correlated with adult survival. 4. Different traits accounted for most of the variance in λ in different ecological settings. Adult survival generally contributed more in populations with low mean adult survival and low mean growth rate during the study period. Hunters and predators (Eurasian lynx and red foxes) occurred in two of our study populations and contributed substantially to the variance in λ, accounting for a total of 35% and 70% in the two populations respectively. 5. Across populations, we did not find any evidence that roe deer increased their reproductive output when faced with harsh conditions, resulting in some populations having negative growth rates. 6. Generation time, a measure of the speed of the life-history cycle, increased from less than 4 years in the most productive population (‘roe deer heaven’) to more than 6 years in declining populations facing predation from lynx, red fox and hunters (‘roe deer hell’), and was tightly and inversely correlated with λ. Such a deceleration of the life cycle in declining populations might be a general feature in large herbivores. 7. Our results shows that the plethora of environmental conditions faced by populations of large herbivores also induce high intraspecific variation in their ranking along the ‘fast–slow’ continuum of life-history tactics. [ABSTRACT FROM AUTHOR]

Summary We used path analysis to investigate the causal relationships between Iberian lynx and Egyptian mongoose track numbers, and to estimate the direct effect of the former on the latter in the Doñana area (2750 km2 ), south-western Spain. Relative abundance of rabbits, shrub cover and protection level were also considered in the path analysis. An observational study consisting of a repeated track survey with a split-plot design was also carried out between 1990 and 1996 to independently test the results obtained by path analysis. Maximum likelihood ratio tests of the goodness-of-fit of the model to the data and other indices used all indicated that the proposed theoretical model depicting the relationships among variables was adequate. Egyptian mongooses suffered a significant direct negative effect from lynx, and rabbits and protection level positively, and significantly affected lynx track abundance; rabbits were also significantly affected by shrub cover (all these standardized partial correlation coefficients ranged from 0.34 to 0.41). Shrub cover did not directly affect either lynx or mongoose tracks. Relatively important indirect effects were found between shrubs and lynx (0.12), and between rabbits and mongooses (-0.14), and protection and mongooses (-0.17). The two latter non-intuitive indirect effects were through the lynx path, as a consequence of the negative direct influence of lynx on mongooses. The repeated track censuses confirmed that lynx presence significantly affects number of mongoose tracks. In areas without lynx, mean number of mongoose tracks were 3.6 times higher than in areas with lynx. It remains an interesting question which of several proposed hypotheses explains co-existence of both species in south-western Spain. [ABSTRACT FROM AUTHOR]

Perceived predation risk and the resulting antipredator behaviour varies across space, time and predator identity. Communities with multiple predators that interact and differ in their use of space, time of activity and hunting mode create a complex landscape for prey to avoid predation. Anthropogenic presence and disturbance have the potential to shift interactions among predators and prey and the where and when encounters occur. We examined how white-tailed deer Odocoileus virginianus fawn spatiotemporal antipredator behaviour differed along an anthropogenic disturbance gradient that had black bears Ursus americanus, coyotes Canis latrans, bobcats Lynx rufus and humans present. We quantified (a) spatial co-occurrence in species distributions, (b) temporal overlap across the diel cycle and (c) spatiotemporal associations between humans, bears, coyotes, bobcats, adult male deer and fawns. We also examined how deer vigilance behaviour changed across the anthropogenic disturbance gradient and survey duration. Anthropogenic disturbance influenced spatiotemporal co-occurrence across multiple scales, often increasing spatiotemporal overlap among species. In general, species' spatial co-occurrence was neutral or positive in anthropogenically disturbed environments. Bears and fawns, coyotes and adult male deer, and bobcats and fawns all had higher temporal overlap in the agriculture-development matrix sites. In addition, factors that influenced deer vigilance (e.g. distance to forest edge and predator relative abundance) in the agriculture-development matrix sites did not in the forest matrix site. By taking into account the different antipredator behaviours that can be detected and the different scales these behaviours might occur, we were able to gain a more comprehensive picture of how humans reduce available niche space for wildlife, creating the neutral and positive spatiotemporal associations between species that studies have been seeing in more disturbed areas., (© 2021 British Ecological Society. This article has been contributed to by US Government employees and their work is in the public domain in the USA.)

1. The Mesopredator Release Hypothesis (MRH) suggests that top predator suppression of mesopredators is a key ecosystem function with cascading impacts on herbivore prey, but it remains to be shown that this top-down cascade impacts the large-scale structure of ecosystems. 2. The Exploitation Ecosystems Hypothesis (EEH) predicts that regional ecosystem structures are determined by top-down exploitation and bottom-up productivity. In contrast to MRH, EEH assumes that interference among predators has a negligible impact on the structure of ecosystems with three trophic levels. 3. We use the recolonization of a top predator in a three-level boreal ecosystem as a natural experiment to test if large-scale biomass distributions and population trends support MRH. Inspired by EEH, we also test if top-down interference and bottom-up productivity impact regional ecosystem structures. 4. We use data from the Finnish Wildlife Triangle Scheme which has monitored top predator (lynx, Lynx lynx), mesopredator (red fox, Vulpes vulpes) and prey (mountain hare, Lepus timidus) abundance for 17 years in a 200 000 km(2) study area which covers a distinct productivity gradient. 5. Fox biomass was lower than expected from productivity where lynx biomass was high, whilst hare biomass was lower than expected from productivity where fox biomass was high. Hence, where interference controlled fox abundance, lynx had an indirect positive impact on hare abundance as predicted by MRH. The rates of change indicated that lynx expansion gradually suppressed fox biomass. 6. Lynx status caused shifts between ecosystem structures. In the 'interference ecosystem', lynx and hare biomass increased with productivity whilst fox biomass did not. In the 'mesopredator release ecosystem', fox biomass increased with productivity but hare biomass did not. Thus, biomass controlled top-down did not respond to changes in productivity. This fulfils a critical prediction of EEH. 7. We conclude that the cascade involving top predators, mesopredators and their prey can determine large-scale biomass distribution patterns and regional ecosystem structures. Hence, interference within trophic levels has to be taken into account to understand how terrestrial ecosystem structures are shaped.

Top predators are increasingly recognized as important regulators of ecosystem structure. Elmhagen et al. in this issue show how a recolonizing population of lynx in Finland is in the process of imposing control of the abundance of a mesopredator, the red fox, and relaxing predation pressure on a prey species. Their study shows how ecological restoration programs could use the power of top predators to limit mesopredator populations and control total predation pressure on prey species.

1. Prey responses to high predation risk can be morphological or behavioural and ultimately come at the cost of survival, growth, body condition, or reproduction. These sub-lethal predator effects have been shown to be mediated by physiological stress. We tested the hypothesis that elevated glucocorticoid concentrations directly cause a decline in reproduction in individual free-ranging female snowshoe hares, Lepus americanus. We measured the cortisol concentration from each dam (using a faecal analysis enzyme immunoassay) and her reproductive output (litter size, offspring birth mass, offspring right hind foot (RHF) length) 30 h after birth. 2. In a natural monitoring study, we monitored hares during the first and second litter from the population peak (2006) to the second year of the decline (2008). We found that faecal cortisol metabolite (FCM) concentration in dams decreased 52% from the first to the second litter. From the first to the second litter, litter size increased 122%, offspring body mass increased 130%, and offspring RHF length increased 112%. Dam FCM concentrations were inversely related to litter size (r(2) = 0.19), to offspring birth mass (r(2) = 0.32), and to offspring RHF length (r(2) = 0.64). 3. In an experimental manipulation, we assigned wild-caught, pregnant hares to a control and a stressed group and held them in pens. Hares in the stressed group were exposed to a dog 1-2 min every other day before parturition to simulate high predation risk. At parturition, unsuccessful-stressed dams (those that failed to give birth to live young) and stressed dams had 837% and 214%, respectively, higher FCM concentrations than control dams. Of those females that gave birth, litter size was similar between control and stressed dams. However, offspring from stressed dams were 37% lighter and 16% smaller than offspring from control dams. Increasing FCM concentration in dams caused the decline of offspring body mass (r(2) = 0.57) and RHF (r(2) = 0.52). 4. This is the first study in a free-ranging population of mammals to show that elevated, predator-induced, glucocorticoid concentrations in individual dams caused a decline in their reproductive output measured both by number and quality of offspring. Thus, we provide evidence that any stressor, not just predation, which increases glucocorticoid concentrations will result in a decrease in reproductive output.