Can predation cause the 10-year hare cycle?

Summary We relate causes of mortality of snowshoe hares to density of hares over an 8-year period that included a peak in numbers. We then use simulation modeling to examine whether these density-dependent relationships could produce changes in hare density similar to those observed in our study are in Yukon, Canada.Predation during winter was the largest source of mortality for snowshoe hares at Kluane, Yukon during 1978–84. There was a one-year lag in the response of winter predation mortality rate to hare density. There was a two-year lag in the response of winter mortality not caused by predators to hare density.A simple simulation model with density-dependent predation produced 8–11 year cycles only within a narrow range of parameters that are inconsistent with data from the Kluane region. However, a simulation model that predicted winter mortality rates using a delayed density-dependent numerical response and a Type II functional response by predators, produced 8–11 year cycles within the range of parameter values measured in our study. Yet another simulation model that predicted both summer and winter mortality rates using a delayed density-dependent numerical response and a Type II functional response by predators, did not produce 8–11 year cycles within the range of parameter values measured in our study. Lack of data on juvenile mortality may be one reason for this result.     

Long-term changes in the feeding pattern of red foxes Vulpes vulpes and their predation on brown hares Lepus europaeus in western Poland

The aim of this study was to estimate long-term changes in the winter feeding pattern of red foxes Vulpes vulpes and in their predation on brown hares Lepus europaeus in relation to the decreasing abundance of hares in western Poland in 1965/1966–2006/2007. The frequencies of occurrence in the stomachs of culled foxes (N?=?726) were used as indices of prey capture rates. The average autumn density of brown hares in the study area decreased from 48 individuals/km² at the turn of the 1960s and 1970s to seven individuals/km² in 1999–2006. Hares and small rodents were the main food classes of foxes in western Poland at the turn of the 1960s and 1970s; however, the occurrence of hares in the fox diet subsequently decreased, and they were replaced by livestock carrion. The relationship between the occurrence frequency of hares in the fox diet and the hare density was best described by sigmoid equation. It indicates that the red fox showed a type III functional response to long-term changes in hare abundance. When predation rate index was estimated on the basis of functional response, the potential fox predation was density-dependent at low to intermediate hare densities (<25 individuals/km²). This finding suggests that the increase in the number of low-density hare populations may require intensive management measures, e.g. simultaneous use of fox control and habitat improvement.     

Avian top predator and the landscape of fear: responses of mammalian mesopredators to risk imposed by the golden eagle

Top predators may induce extensive cascading effects on lower trophic levels, for example, through intraguild predation (IGP). The impacts of both mammalian and avian top predators on species of the same class have been extensively studied, but the effects of the latter upon mammalian mesopredators are not yet as well known. We examined the impact of the predation risk imposed by a large avian predator, the golden eagle (Aquila chrysaetos, L.), on its potential mammalian mesopredator prey, the red fox (Vulpes vulpes, L.), and the pine marten (Martes martes, L.). The study combined 23 years of countrywide data from nesting records of eagles and wildlife track counts of mesopredators in Finland, northern Europe. The predation risk of the golden eagle was modeled as a function of territory density, density of fledglings produced, and distance to nearest active eagle territory, with the expectation that a high predation risk would reduce the abundances of smaller sized pine martens in particular. Red foxes appeared not to suffer from eagle predation, being in fact most numerous close to eagle nests and in areas with more eagle territories. This is likely due to similar prey preferences of the two predators and the larger size of foxes enabling them to escape eagle predation risk. Somewhat contrary to our prediction, the abundance of pine martens increased from low to intermediate territory density and at close proximity to eagle nests, possibly because of similar habitat preferences of martens and eagles. We found a slightly decreasing trend of marten abundance at high territory density, which could indicate that the response in marten populations is dependent on eagle density. However, more research is needed to better establish whether mesopredators are intimidated or predated by golden eagles, and whether such effects could in turn cascade to lower trophic levels, benefitting herbivorous species.     

Role of predation in short-term population fluctuations of some birds and mammals in Fennoscandia

Summary We tested the hypothesis that synchronous fluctuations in small game species in boreal Fennoscandia are caused by varying predation pressure. The main prey of predators are the cyclically superabundant voles. Small game species (alternative prey) are rare compared to voles. The following 4 predictions were checked: (1) Predators should shift their diet from main prey to alternative prey as main prey decline. — This was confirmed using data on red fox (Vulpes vulpes L.) diet.; (2) The mortality rate of alternative prey should be inversely correlated to the abundance of main prey. — This was true for mountain hare (Lepus timidus L.) mortality rates and the rate of nest predation on black grouse (Tetrao tetrix L.).; (3) The total consumption of prey by all the predators should at least equal the critical losses in alternative prey during a decline year. — A tentative estimate of predator consumption amounted to 10 times the losses in grouse and hare.; and (4) The absence of synchrony between the species in the boreonemoral region should be associated with a more diverse diet of predators. — This was the case for red fox diets throughout Sweden. Although all 4 predictions were confirmed, we could not necessarily exclude other hypotheses involving changes in quality or quantity of plant food.     

Predation on rodents in Białowieza primeval forest, Poland

Autumn and winter predation on bank vole Clethrtonomys glareolus and yellow-necked mouse Apodemus flavicollis was studied in 1985/86-88/89 in an 11 2 km² area of the Bialowieza National Park Rodents regularly increased in numbers from spring to autumn and decreased throughout winter Out of 23 species of predators, the most common were tawny owl Strix aluco (43-57 adult ind 10 km⁻²) weasel Mustela ntvahs (17-27 ind), buzzard Buteo buteo (12-16) and pine marten Martes martes (5-8) Voles and mice were the staple food for two specialists the stoat Mustela erminea and the weasel, and two generahsts the tawny owl and the pine marten The generalists exploited different alternative prey when rodents were scarce tawny owl -amphibians, marten - small mammals, and the red fox Vulpes vulpes - ungulate carcasses and hares The depth of snow and abundance of voles were two major factors shaping the contribution of voles to tawny owl and marten diets No such relationships were found for mice and generalist predators The predation by 8 species (tawny owl, buzzard, marten, weasel, stoat, polecat Mustela putortus fox, and raccoon dog Nyctereutes procyonoides) from 1 October to 15 April in 1986/87, 87/88, and 88/89 was estimated to be on average 28-35 voles and 14-17 mice ha Estimates suggested that three species were responsible for 86-95% of the total predation impact tawny owl (56-71%), weasel (11-21%) and marten (10-15%) The predation impact was similar to the annual decrease in rodent numbers from autumn till spring voles 35 (SD 111) lnds ha^-1in autumn and 8 (SD 2 6) lnds ha^-1in spring, and mice 24 (SD 16 3) mds ha⁻¹ in autumn and 3 (SD 2 5) in spring Predation, therefore, was regarded as the main agent of rodent mortality throughout autumn and winter     

Owl predation on snowshoe hares: consequences of antipredator behaviour

We show evidence of differential predation on snowshoe hares (Lepus americanus) by great horned owls (Bubo virginianus) and ask whether predation mortality is related to antipredator behaviour in prey. We predicted higher predation on (1) young and inexperienced hares, (2) hares in open habitats lacking cover to protect from owl predation, and (3) hares in above average condition assuming that rich food patches are under highest risk of predation. Information on killed hares was obtained at nest sites of owls and by monitoring hares using radio-telemetry. The availability of age classes within the hare population was established from live-trapping and field data on reproduction and survival. Great horned owls preferred juvenile over adult hares. Juveniles were more vulnerable to owl predation before rather than after dispersal, suggesting that displacement or increased mobility were not causes for this increased mortality. Owls killed ratio-collared hares more often in open than in closed forest types, and they avoided or had less hunting success in habitats with dense shrub cover. Also, owls took hares in above average condition, although it is unclear whether samples from early spring are representative for other seasons. In conclusion, these results are consistent with the hypothesis that variation in antipredator behaviours of snowshoe hares leads to differential predation by great horned owls.     

Implications of shared predation for space use in two sympatric leporids

Spatial variation in habitat riskiness has a major influence on the predator–prey space race. However, the outcome of this race can be modulated if prey shares enemies with fellow prey (i.e., another prey species). Sharing of natural enemies may result in apparent competition, and its implications for prey space use remain poorly studied. Our objective was to test how prey species spend time among habitats that differ in riskiness, and how shared predation modulates the space use by prey species. We studied a one‐predator, two‐prey system in a coastal dune landscape in the Netherlands with the European hare (Lepus europaeus) and European rabbit (Oryctolagus cuniculus) as sympatric prey species and red fox (Vulpes vulpes) as their main predator. The fine‐scale space use by each species was quantified using camera traps. We quantified residence time as an index of space use. Hares and rabbits spent time differently among habitats that differ in riskiness. Space use by predators and habitat riskiness affected space use by hares more strongly than space use by rabbits. Residence time of hare was shorter in habitats in which the predator was efficient in searching or capturing prey species. However, hares spent more time in edge habitat when foxes were present, even though foxes are considered ambush predators. Shared predation affected the predator–prey space race for hares positively, and more strongly than the predator–prey space race for rabbits, which were not affected. Shared predation reversed the predator–prey space race between foxes and hares, whereas shared predation possibly also released a negative association and promoted a positive association between our two sympatric prey species. Habitat riskiness, species presence, and prey species’ escape mode and foraging mode (i.e., central‐place vs. noncentral‐place forager) affected the prey space race under shared predation.     

Cycles in voles and small game in relation to variations in plant production indices in Northern Sweden

Population dynamics for voles (Cricetidae), Tengmalm's owl (Aegolius funereus (L.)), red fox (Vulpes vulpes (L.)) willow grouse (Lagopus lagopus (L.)), black grouse (Lyrurus tetrix (L.)), capercaillie (Tetrao urogallus L.), hazel hen (Tetrastes bonasia (L.)), mountain hare (Lepus timidus L.) and tularemia (Francisella tularensis (McCoy & Chapin)) and game bird recruitment were studied by index methods in northern Sweden. In addition contemporary temperature records and spruce (Picea abies (L.) Karst.) and pine (Pinus silvestris L.) cone crops (as indices for plant production) and the occurrence of forest damage, caused by voles eating bark, were studied.During 1970–80 two synchronous 4-year cycles were observed for voles, predators (Tengmalm's owl and red fox) and their alternative prey species (grouse and mountain hare). In grouse the change of numbers was correlated with that of recruitment. Autumn vole numbers peaked about a year before the other species and extensive forest damage occurred at winter peak densities of voles. These population fluctuations are consistent with a predator-prey model for their regulation. In short the model suggests that vole-food plant interactions trigger the cycle of voles, that voles generate the cycle of predators and that these in turn synchronize alternative prey populations to the others at vole declines.For voles, grouse and red fox the amplitude was higher in the first cycle compared to the second one whilst the opposite was true for the mountain hare. Although temperature and cone crops showed large interannual variations they still implied that herbivore food conditions were better during the former cycle. Hence, the reduction of the amplitude of the vole cycle may be explained by inter-cyclic differences in plant food conditions, implying food shortage (as indicated by bark-eating) at different population levels. The similar decrease of grouse and red fox populations may also be explained by deteriorated food conditions and/or for the fox by an outbreak of sarcoptic mange (Sarcoptes scabiae var. vulpes). The increased amplitude of the mountain hare cycle was part of a long-term rise in numbers after a tularemia epidemic in 1967. This is interpreted as a recovery, probably towards the generally higher pre-epidemic population level.     

The fluctuating world of a tundra predator guild: bottom‐up constraints overrule top‐down species interactions in winter

Global warming is predicted to change ecosystem functioning and structure in Arctic ecosystems by strengthening top‐down species interactions, i.e. predation pressure on small herbivores and interference between predators. Yet, previous research is biased towards the summer season. Due to greater abiotic constraints, Arctic ecosystem characteristics might be more pronounced in winter. Here we test the hypothesis that top‐down species interactions prevail over bottom‐up effects in Scandinavian mountain tundra (Northern Sweden) where effects of climate warming have been observed and top‐down interactions are expected to strengthen. But we test this ‘a priori’ hypothesis in winter and throughout the 3–4 yr rodent cycle, which imposes additional pulsed resource constraints. We used snowtracking data recorded in 12 winters (2004–2015) to analyse the spatial patterns of a tundra predator guild (arctic fox Vulpes lagopus, red fox Vulpes vulpes, wolverine Gulo gulo) and small prey (ptarmigan, Lagopus spp). The a priori top‐down hypothesis was then tested through structural equation modelling, for each phase of the rodent cycle. There was weak support for this hypothesis, with top‐down effects only discerned on arctic fox (weakly, by wolverine) and ptarmigan (by arctic fox) at intermediate and high rodent availability respectively. Overall, bottom‐up constraints appeared more influential on the winter community structure. Cold specialist predators (arctic fox and wolverine) showed variable landscape associations, while the boreal predator (red fox) appeared strongly dependent on productive habitats and ptarmigan abundance. Thus, we suggest that the unpredictability of food resources determines the winter ecology of the cold specialist predators, while the boreal predator relies on resource‐rich habitats. The constraints imposed by winters and temporary resource lows should therefore counteract productivity‐driven ecosystem change and have a stabilising effect on community structure. Hence, the interplay between summer and winter conditions should determine the rate of Arctic ecosystem change in the context of global warming.     

Functional and numerical responses of four lemming predators in high arctic Greenland

The high‐arctic tundra ecosystem has the world's simplest vertebrate predator–prey community, with only four predators preying upon one rodent species, the collared lemming (Dicrostonyx groenlandicus). We document the functional and numerical responses of all the four predators in NE Greenland. Using these data, we assess the impact of predation on the dynamics of the collared lemming with a 4 yr cycle and >100‐fold difference between maximum and minimum densities. All predator species feed mostly (>90%) on lemmings when lemming density is >1 ha^?1, but the shapes of the predators’ responses vary greatly. The snowy owl (Nyctea scandiaca) is present and breeds only when lemming densities at snowmelt are >2 ha^?1, giving rise to a step‐like numerical response. The long‐tailed skua (Stercorarius longicaudus) has a type III functional response and shifts from alternate food (mainly berries and insects) to lemmings with increasing lemming density. The skua surpasses all the other predators in summer by its total response. The type III functional response of the Arctic fox (Alopex lagopus) starts to increase at much lower lemming densities than the responses of the avian predators, but it has only a weak numerical response. Finally, the stoat (Mustela erminea) is the most specialized predator and the only one with a clearly delayed numerical response. According to their specific functional and numerical responses, each predator plays a key role at some point of the lemming cycle, but only the stoat has the potential to drive the lemming cycle. Stoat predation is greatly reduced in the winter preceding the lemming peak, and it reaches a maximum in the winter preceding the lowest lemming summer density. Stoat predation appears to maintain low lemming densities for at least two successive years. Our study provides empirical support for the specialist predator hypothesis about small mammal population cycles.     

Predation Risk Drives Habitat‐Specific Sex Ratio in a Monomorphic Species,the Brown Hare (Lepus europaeus)

In dimorphic species, sexual habitat segregation is generally explained by the differences in nutritional needs or by a trade‐off between fulfilling food requirements and avoiding predation. However, it remains unclear whether predation risk is strong enough to drive the differences in habitat use between sexes as predicted by the predation sensitivity hypothesis. Here we test in a monomorphic species, the brown hare (Lepus europaeus), the prediction that abundance of the gender more sensitive to predation is higher in safer habitat. We used data on 1645 individually marked hares in western Poland during autumn–winter seasons of 1966/1967–1978/1979 to estimate sex‐specific annual survival rates. We analyzed the stomach contents of 134 foxes shot in 1965/1966–1994/1995 to evaluate fox predation on hares. Finally, we employed data on 26 790 hares live‐trapped in 1965/1966–1994/1995 to analyze hare sex ratio across habitats. We found that male annual survival rate was lower than that of females and that the predation risk by foxes on hares was lower in agricultural than forest habitat. Our finding, that males were more often trapped by nets in agricultural than the forest habitat, provides indirect evidence for the predation sensitivity hypothesis. We conclude that predation risk can be a driving force for habitat‐specific sex ratio in a monomorphic species such as the brown hare.     

The effect of experimental removal of red foxesVulpes vulpes on spring density of brown haresLepus europaeus in western Poland

Red foxesVulpes vulpes (Linnaeus, 1758) were experimentally removed in two nearby areas located in western Poland to verify the hypothesis about the limiting impact of their predation on the low-density population of brown haresLepus europaeus (Pallas, 1778) (4.4-10.6 ind./km² in late autumn). In 1996/1997–2001/2002 foxes were culled (mainly in autumn and winter) in the reduction area (32 km²), whereas in the control area (34 km²) intensive culling was carried out only in 2000/2001–2001/2002. Indices of fox and hare spring densities were estimated using spotlight counts, as mean numbers of individuals observed per 10 km of the counting route. Annual changes in the fox density indices were negatively correlated with the bag of foxes, and annual changes in the hare density indices were negatively related to the annual changes in fox density indices. The fox density indices were significantly lower in the reduction area than in the control one only in 2000–2001 (2.8 times, on average), and in the same years, the hare population responded with higher density (1.7 times, on average). The hare responses took place without time delay, which suggests that the changes in fox abundance affected the situation of hares primarily in the autumn-winter season.     

Variation in the diet composition of a generalist predator, the red fox, in relation to season and density of main prey

Diet composition of a generalist predator, the red fox (Vulpes vulpes) in relation to season (winter or summer) and abundance of multi-annually cyclic voles was studied in western Finland from 1983 to 1995. The proportion of scats (PS; a total of 58 scats) including each food category was calculated for each prey group. Microtus voles (the field vole M. agrestis and the sibling vole M. rossiaemeridionalis) were the main prey group of foxes (PS = 0.55) and they frequently occurred in the scats both in the winter and summer (PSs 0.50 and 0.62, respectively). There was a positive correlation between the PSs of Microtus voles in the winter diet of foxes and the density indices of these voles in the previous autumn. Other microtine rodents (the bank vole Clethrionomys glareolus, the water vole Arvicola terrestris and the muskrat Ondatra zibethicus) were consumed more in winter than in summer. The unusually high small mustelid predation by red foxes (PS = approx. 0.10) in our study area gives qualitative support for the hypothesis on the limiting impact of mammalian predators on least weasel and stoat populations. None of the important prey groups was preyed upon more at low than at high densities of main prey (Microtus voles). This is consistent with the notion that red foxes are generalist predators that tend to opportunistically subsist on many prey groups. Among these prey groups, particularly hares and birds (including grouse), were frequently used as food by foxes.     

Seasonal Effects of Habitat on Sources and Rates of Snowshoe Hare Predation in Alaskan Boreal Forests

Survival and predation of snowshoe hares (Lepus americanus) has been widely studied, yet there has been little quantification of the changes in vulnerability of hares to specific predators that may result from seasonal changes in vegetation and cover. We investigated survival and causes of mortalities of snowshoe hares during the late increase, peak, and decline of a population in interior Alaska. From June 2008 to May 2012, we radio-tagged 288 adult and older juvenile hares in early successional and black spruce (Picea mariana) forests and, using known-fate methods in program MARK, evaluated 85 survival models that included variables for sex, age, and body condition of hares, as well as trapping site, month, season, year, snowfall, snow depth, and air temperature. We compared the models using Akaike’s information criterion with correction for small sample size. Model results indicated that month, capture site, and body condition were the most important variables in explaining survival rates. Survival was highest in July, and more generally during summer, when alternative prey was available to predators of hares. Low survival rates coincided with molting periods, breeding activity in the spring, and the introduction of juveniles to the sample population in the fall. We identified predation as the cause of mortality in 86% of hare deaths. When the source of predation could be determined, hares were killed more often by goshawks (Accipiter gentilis) than other predators in early successional forest (30%), and more often by lynx (Lynx canadensis) than other predators in black spruce forest (31%). Great horned owls (Bubo virginianus) and coyotes (Canis latrans) represented smaller proportions of hare predation, and non-predatory causes were a minor source (3%) of mortality. Because hares rely on vegetative cover for concealment from predators, we measured cover in predation sites and habitats that the hares occupied and concluded that habitat type had a greater influence on the sources of predation than the amount of cover in any given location within a habitat. Our observations illustrate the vulnerability of hares to predators in even the densest coniferous habitat available in the boreal forest, and indicate strong seasonal changes in the rates and sources of predation.     

From process to pattern: how fluctuating predation risk impacts the stress axis of snowshoe hares during the 10-year cycle

Predation is a central organizing process affecting populations and communities. Traditionally, ecologists have focused on the direct effects of predation—the killing of prey. However, predators also have significant sublethal effects on prey populations. We investigated how fluctuating predation risk affected the stress physiology of a cyclic population of snowshoe hares (Lepus americanus) in the Yukon, finding that they are extremely sensitive to the fluctuating risk of predation. In years of high predator numbers, hares had greater plasma cortisol levels at capture, greater fecal cortisol metabolite levels, a greater plasma cortisol response to a hormone challenge, a greater ability to mobilize energy and poorer body condition. These indices of stress had the same pattern within years, during the winter and over the breeding season when the hare:lynx ratio was lowest and the food availability the worst. Previously we have shown that predator-induced maternal stress lowers reproduction and compromises offspring’s stress axis. We propose that predator-induced changes in hare stress physiology affect their demography through negative impacts on reproduction and that the low phase of cyclic populations may be the result of predator-induced maternal stress reducing the fitness of progeny. The hare population cycle has far reaching ramifications on predators, alternate prey, and vegetation. Thus, predation is the predominant organizing process for much of the North American boreal forest community, with its indirect signature—stress in hares—producing a pattern of hormonal changes that provides a sensitive reflection of fluctuating predator pressure that may have long-term demographic consequences.     

Red fox trap success is correlated with piping plover chick survival

Predation management is an important component of managing species of conservation concern. The piping plover (Charadrius melodus; plover), a disturbance-dependent and conservation-reliant shorebird that nests on sandy beaches and barrier islands on the Atlantic Coast, was listed under the United States Endangered Species Act in 1986, with habitat loss and predation stated as key causes of its decline. We evaluated the relationship between a suite of predators and plover chick survival from 2015–2018. We used a camera grid to establish indices of the abundance of 3 known chick predators: red fox (Vulpes vulpes), raccoon (Procyon lotor), and domestic cat. We used camera detections in a survival model to assess potential relationships between predator species detection and plover chick survival. Plover chick survival was negatively related with red fox detection, but not with detection of the other 2 predators. In addition to the correlation with red fox detections, chick survival was negatively related to high plover nesting density. Our results suggest that red foxes were predators of piping plover chicks during our study, likely augmented by other density-dependent sources of mortality. Targeted predator management could aid in conservation of piping plovers in this system as a short-term solution, but long-term recovery plans must also address habitat limitation.     

Guardian or threat: does golden eagle predation risk have cascading effects on forest grouse?

Previous studies on intraguild predation have mainly focused on within-class assemblages, even though avian top predators may also influence mammalian mesopredator prey. By using nation-wide long-term data from Finland, northern Europe, we examined the impacts of golden eagles (Aquila chrysaetos) together with red foxes (Vulpes vulpes) and pine martens (Martes martes) on forest-dwelling herbivores, black grouse (Tetrao tetrix) and hazel grouse (Tetrastes bonasia). We hypothesized that eagles may alleviate the overall predation pressure on grouse by imposing intraguild predation risk on mesopredators. The predation impact of eagle was modelled using eagle density estimates and distance to eagle nest. Wildlife triangle counts were used as predation impact proxies of mammalian mesopredators and as measures of response in grouse. Our results show that eagle density correlated negatively with black grouse abundance indices while being positively associated with the proportion of juveniles in both grouse species, irrespective of the abundance of mesopredators. Yet, foxes and martens alone had a negative effect on the abundance indices and the proportion of young in the two grouse species. This suggests that the possible cascading effects of eagles are not mediated by decreased mesopredator numbers, but instead by fear effects. Alternatively, they may be mediated by other species than fox or marten studied here. In conclusion, we found support for the hypothesis that eagles provide protection for juvenile black and hazel grouse, whereas they are a threat for adult grouse. This important information helps us to better understand the role of avian top predators in terrestrial ecosystems.     

Role of prey and intraspecific density dependence on the population growth of an avian top predator

Exploring predator–prey systems in diverse ecosystems increases our knowledge about ecological processes. Predator population growth may be positive when conspecific density is low but predators also need areas with prey availability, associated with competition, which increases the risk of suffering losses but stabilises populations. We studied relationships between European rabbits Oryctolagus cuniculus (prey) and adult eagle owls Bubo bubo (predators) in south-western Europe. We assessed models explaining the predator population growth and stability. We estimated the abundance of rabbits and adult eagle owls during three years in eight localities of central-southern Spain. We explored models including rabbit and adult eagle owl abundance, accounting for yearly variations and including the locality as a random variable. We found that population growth of adult eagle owls was positive in situations with low conspecific abundance and tended to be negative but approaching equilibrium in situations of higher conspecific abundance. Population growth was also positively related to previous summer rabbit density when taking into account eagle owl conspecific abundance, possibly indicating that rabbits may support recruitment. Furthermore, abundance stability of adult eagle owls was positively related to previous winter–spring rabbit density, which could suggest predator population stabilisation through quick territory occupation in high-quality areas. These results exemplify the trade-off between prey availability and abundance of adult predators related to population growth and abundance stability in the eagle owl–rabbit system in south-western Europe. Despite rabbits have greatly declined during the last decades and eagle owls locally specialise on them, eagle owls currently have a favourable conservation status. As eagle owls are the only nocturnal raptor with such dependence on rabbits, this could point out that predators may overcome prey decreases in areas with favourable climate and prey in the absence of superior competitors with similar foraging mode.     

The recent expansion of the brown hare (<Emphasis Type="Italic">Lepus europaeus</Emphasis>) in Sweden with possible implications to the mountain hare (<Emphasis Type="Italic">L. timidus</Emphasis>)

The brown hare (Lepus europaeus) expanded its Swedish distribution since the 1980s northwards and locally to new areas within its former range. Of 115 brown hare populations within the former range reported in a hunter enquiry, those established after 1980 were situated higher above the sea level than older ones and higher than neighbouring (<50 km) older populations. Reports on increased use of forest habitats by brown hares were equally frequent among recent and older populations, suggesting a process promoted solely by less harsh winters. Supposed hare hybrids were more often reported from hunting grounds with recent brown hare establishment, i.e. where the species expands in time and in space. In a 27-year dataset on brown hare observations, the recent increased use of forest habitats was supported in that maximum distances to agricultural land for brown hare sightings were higher in mild winters, whereas the proportions of the annual observations made during winter were lower. In 40-year bag records from two Swedish counties, the dynamics of the mountain hare (Lepus timidus) responded positively to snow parameters, whereas brown hares responded negatively. We suggest that the state of mountain hare populations primarily depends on winter conditions and predation pressure, whereas possible effects of hybridization are unclear. If winter conditions remain as in the last 15 years, mountain hare numbers are not likely to increase in southern Sweden, whereas the brown hare may expand even further. In either case, hybrids will occur in sympatric areas in frequencies probably related to the density of the respective true species.     

Predator-Mediated Indirect Effects of Snowshoe Hares on Dall's Sheep in Alaska

ABSTRACT Indirect interactions among species can strongly influence population dynamics and community structure but are often overlooked in management of large mammals. We estimated survival of Dall's sheep (Ovis dalli) in the central Alaska Range, USA, during years of differing snowshoe hare (Lepus americanus) abundance to test whether indirect interactions with a cyclic hare population affect Dall's sheep either negatively, by subsidizing predators (apparent competition), or positively, by diverting predation (apparent commensalism). Annual survival of adult female sheep was consistently high (0.85 for all yr and age classes combined). In contrast, annual estimates of lamb survival ranged from 0.15 to 0.63. The main predators of lambs were coyotes (Canis latrans) and golden eagles (Aquila chrysaetos), which rely on hares as their primary food and prey on lambs secondarily. Coyotes and eagles killed 78% of 65 radiocollared lambs for which cause of death was known. Lamb survival was negatively related to hare abundance during the previous year, and lamb survival rates more than doubled when hare abundance declined, supporting the hypothesis of predator-mediated apparent competition between hares and sheep. However, stage-specific predation and delays in predator responses to changes in hare numbers led to a positive relationship between abundance of adult Dall's sheep and hares. Lacking reliable estimates of survival, a manager might erroneously conclude that hares benefit sheep. Thus, support for different indirect effects can be obtained from different types of data, which demonstrates the need to determine the mechanisms that create indirect interactions. Long-term survey data suggest that predation by coyotes is limiting this sheep population below levels typical when coyotes were rare or absent. Understanding the nature of indirect interactions is necessary to effectively manage complex predator–prey communities.