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1.
Large predators may affect the hunting efficiency of smaller ones directly by decreasing their numbers, or indirectly by altering their behaviour. Either way this may have positive effects on the density of shared prey. Using large outdoor enclosures, we experimentally studied whether the presence of the Tengmalm's owl Aegolius funereus affects the hunting efficiency of the smallest member of the vole-eating predator guild, the least weasel Mustela nivalis, as measured by population responses of coexisting prey species, the field vole Microtus agrestis and the sibling vole M. levis . We compared the density and survival probability of vole populations exposed to no predation, weasel predation or combined predation by a weasel and an owl. The combined predation of both owl and weasel did not result in obvious changes in the density of sibling and field vole populations compared to the control populations without predators, while predation by least weasel alone decreased the densities of sibling voles and induced a similar trend in field vole densities. Survival of field voles was not affected by predator treatment while sibling vole survival was lower in predator treated populations than in control populations. Our results suggest that weasels are intimidated by avian predators, but without changing the effects of predators on competitive situations between the two vole species. Non-lethal effects of intraguild predation therefore will not necessarily change competitive interactions between shared prey species.  相似文献   

2.
Many predator species feed on prey that fluctuates in abundance from year to year. Birds of prey can face large fluctuations in food abundance i.e. small mammals, especially voles. These annual changes in prey abundance strongly affect the reproductive success and mortality of the individual predators and thus can be expected to influence their population dynamics and persistence. The barn owl, for example, shows large fluctuations in breeding success that correlate with the dynamics in voles, their main prey species. Analysis of the impact of fluctuations in vole abundance (their amplitude, peaks and lows, cycle length and regularity) with a simple predator prey model parameterized with literature data indicates population persistence is especially affected by years with low vole abundance. In these years the population can decline to low owl numbers such that the ensuing peak vole years cannot be exploited. This result is independent of the length and regularity of vole fluctuations. The relevance of this result for conservation of the barn owl and other birds of prey that show a numerical response to fluctuating prey species is discussed.  相似文献   

3.
Theoretical models predict that a delayed density-dependent mortality factor with a time lag of ca 9 months is able to drive 3–5-yr population cycles of northern voles. We studied numerical responses of predators in western Finland during 1986–92, in an area with 3-yr population cycles of voles. Abundances of small mammals were monitored in several farmland areas (each 3 km2) by snap-trapping in April, June, August, and October (only in 1986–90), and the abundances of avian, mammalian, and reptilian predators by visual censuses during trapping occasions. The 3-yr cycle studied was a cycle of Microtus voles (field vole M. agrestis and sibling vole M. rossiaemeridionalis ) and their small-sized predators (small mustelids and vole-eating birds of prey). The numerical responses of both migratory avian predators and small mustelids to changes in vole densities were more alike than different. In late summer (August), the time lag in the numerical response of all main predators was short (0–4 months), whereas longer time lags prevailed from spring to early summer. The length of the time lag in spring appeared to be related to the length of the winter, which indicates that strong seasonality may create longer time lags to the numerical response of predators at northern latitudes than at more southern latitudes. Our results suggest that, from spring to early summer, predation by migratory avian predators may act in concordance with mustelid predation to produce the long time lag necessary to drive the 3-yr cycle of voles, whereas almost direct density-dependent predation by all major predators in late summer may dampen spatial variation in prey densities.  相似文献   

4.
Joshua T. Ackerman 《Oikos》2002,99(3):469-480
Coexisting prey species interact indirectly via their shared predators when one prey type influences predation rates of the second prey type. In a temperate system where the predominant shared predator is a generalist, I studied the indirect effects of rodent populations on waterfowl nest success, both within the nesting season among sites and among years. Among six to ten upland fields (14 to 27 ha), mallard ( Anas platyrhynchos ) nest success was positively correlated with rodent abundance in all three years of the study. After removing year effects, mallard nest success remained positively correlated with the relative abundance of rodents. Of the rodent species present, California voles ( Microtus californicus ) were the most important coexisting prey type influencing nest success. Among years, mallard nest success was positively correlated with vole abundance; the asymptotic relationship suggests a threshold response to vole abundance, beyond which predators become satiated and additional voles do little to affect nest success. I tested and rejected three alternative explanations for the observed positive correlation between mallard nest success and rodent abundance that do not involve an indirect effect of coexisting prey populations. The influences of dense nesting cover, nesting density, and predator activity did not explain the observed patterns of nest success. These results suggest that rodent populations buffer predation on waterfowl nests, both within and among years, via the behavioral responses of shared predators to coexisting prey.  相似文献   

5.
In small rodent populations with wide-amplitude fluctuations and low-density bottlenecks, the individuals that survive through the bottlenecks may gain major fitness advantages as they will be the founders of the following population expansion. Most hypotheses assume that there exists a physical or behavioural refuge from increased predation risk, and that the survivors are most likely individuals adapted to use such refuges. A recent hypothesis suggests that survival probability is habitat-dependent so that some otherwise sub-optimal habitats provide a spatial refuge from predation risk by the main predator(s). We used spatially replicated long-term (1981–2004) trapping and tracking data of voles (field vole Microtus agrestis and sibling vole M. rossiaemeridionalis) and their main predators (weasel Mustela nivalis and stoat M. erminea) to test predictions based on this hypothesis. We did not find support for the hypothesis. We did not find marked phase-dependent differences in the habitat-level distribution of Microtus voles. Habitat types with low Microtus vole abundance had, on average, comparable predator activity than the main Microtus vole habitats, indicating that there were no habitat-level refuges from predators. There appeared to be no permanent site-level refuges: the spatial distribution of voles varied from one bottleneck to another. This suggests that survival through bottlenecks is at least partly determined by chance events. We propose that in this kind of systems, where relatively short-lived prey are hunted by nomadic or widely ranging predators, short-term anti-predator responses may increase survival prospects as efficiently as more costly anti-predator adaptations, and there is no apparent need to maintain special adaptations to bottleneck situations that occur at infrequent intervals.Co-ordinating editor: J. Tuomi  相似文献   

6.
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.  相似文献   

7.
The regional synchrony of short-term population fluctuations of small rodents and small game has usually been explained by varying impacts of generalist predators subsisting on both voles and small game (the "alternative prey hypothesis" APH). APH says that densities of predators increase as a response to increasing vole densities and then these predators shift their diet from the main prey to the alternative prey when the main prey decline and vice versa. We studied the diet composition of breeding common buzzards Buteo buteo during 1985-92 in western Finland. Microtus voles were the main prey and water voles, shrews, forest grouse, hares and small birds the most important alternative prey. Our data from the between-year variation in the diet composition of buzzards fulfilled the main predictions of APH. The yearly proportion of main prey (Microtus voles) in the diet was higher in years of high than low vole abundance. The proportion of grouse in the diet of buzzards was negatively related to the abundance of Microtus voles in the field and was nearly independent of grouse abundance in the field. In addition, buzzards mainly took grouse chicks and young hares which is consistent with the prediction of APH. Therefore, we conclude that buzzards are able to shift their diet in the way predicted by the APH and that buzzards, together with other generalist predators, may reduce the breeding success of small game in the decline phase of the vole cycle, and thus substantially contribute to the existence of short-term population cycles of small game.  相似文献   

8.
The consequences of cyclic fluctuations in abundance of prey species on predator continue to improve our understanding of the mechanisms behind population regulation. Among predators, vole‐eating raptors usually respond to changes in prey abundance with no apparent time‐lag and therefore contradict predictions from the predator–prey theory. In such systems, the interplay between demographic traits and population growth rate in relation to prey abundance remains poorly studied, yet it is crucial to characterize the link between ecological processes and population changes. Using a mechanistic approach, we assessed the demographic rates associated to the direct and indirect numerical responses of a specialist raptor (Montagu's harrier) to its cyclic prey (common vole), using long term data from two adjacent study sites in France. First‐year survival rates were weakly affected by vole abundance, probably due to the fact that Montagu's harriers are trans‐Saharan migrants and thus escape the vole collapse occurring in autumn–winter. Recruitment of yearling as well as breeding propensity of experienced adult females were strongly affected by vole abundance and at least partially shaped the trajectory of the breeding population. We argued that the strong density dependent signal detected in predator time series was mostly the phenomenological consequence of the positive direct numerical response of harriers to vole abundance. Accounting for this, we proposed a method to assess density dependence in predator relying on a cyclic prey. Finally, the variation in Montagu's harrier population growth rates was best explained by overwinter growth rates of the prey population and to a lesser extent by previous residual predator density.  相似文献   

9.
Comprehensive analyses of long-term (1977-2003) small-mammal abundance data from western Finland showed that populations of Microtus voles (field voles M. agrestis and sibling voles M. rossiaemeridionalis) voles, bank (Clethrionomys glareolus) and common shrews (Sorex araneus) fluctuated synchronously in 3 year population cycles. Time-series analyses indicated that interspecific synchrony is influenced strongly by density-dependent processes. Synchrony among Microtus and bank voles appeared additionally to be influenced by density-independent processes. To test whether interspecific synchronization through density-dependent processes is caused by predation, we experimentally reduced the densities of the main predators of small mammals in four large agricultural areas, and compared small mammal abundances in these to those in four control areas (2.5-3 km(2)) through a 3 year small-mammal population cycle. Predator reduction increased densities of the main prey species, Microtus voles, in all phases of the population cycle, while bank voles, the most important alternative prey of predators, responded positively only in the low and the increase phase. Manipulation also increased the autumn densities of water voles (Arvicola terrestris) in the increase phase of the cycle. No treatment effects were detected for common shrews or mice. Our results are in accordance with the alternative prey hypothesis, by which predators successively reduce the densities of both main and alternative prey species after the peak phase of small-mammal population cycles, thus inducing a synchronous low phase.  相似文献   

10.
Predation has been invoked as a factor synchronizing the population oscillations of sympatric prey species, either because predators kill prey unselectively (the Shared Predation Hypothesis; hereafter SPH), or because predators switch to alternative prey after a density decline in their main prey (the Alternative Prey Hypothesis; APH). A basic assumption of the APH is that the impact of predators on alternative prey depends more on the density of main prey than on the predator/alternative prey ratio. Both SPH and APH assume that the impact of predators on alternative prey is at least periodically strong enough to depress prey populations. To examine these assumptions, we utilized data from replicated field experiments in large areas where we reduced the breeding densities of avian predators during three years and the numbers of least weasels (Mustela nivalis) in two years when vole populations declined. In addition, we reduced the breeding densities of avian predators in two years when vole populations were high. The reduction of least weasels increased the abundance of their alternative prey, small birds breeding on the ground, but did not affect the abundance of common shrews (Sorex araneus). In years when vole populations declined, the reduction of avian predators increased the abundance of their alternative prey, common shrews and small birds. Therefore, vole‐eating predators do at least periodically depress the abundance of their alternative prey. At high vole densities, the reduction of avian predators did not increase the abundance of common shrews, although the ratio of avian predators to alternative prey was similar to years when vole populations declined, which supported APH. In contrast, the abundance of small birds increased after the reduction of avian predators also at high vole densities, which supported SPH. The manipulations had no obvious effect on the number of game birds, which are only occasionally killed by these small‐sized predators. We conclude that in communities where most predators are small or specialize on a single prey type, the synchronizing impact of predation is restricted to a few similar‐sized species.  相似文献   

11.
Interspecific competition is assumed to generate negative effects on coexisting species, possibly including slower population growth and lower survival. The field vole ( Microtus agrestis ) and the sibling vole ( M. rossiaemeridionalis ) are sympatric close relatives which compete for similar resources. Previous non-experimental studies suggest that the smaller sibling vole is a superior competitor, yet more vulnerable to predation than the larger field vole. We studied the effects of coexistence on population densities, reproductive parameters, and survival in these two species by means of experimentation in large, predator-free outdoor enclosures. While populations of both species reached higher densities in the absence of the other, field voles appeared to suffer more from interspecific competition than sibling voles. The proportion of young individuals in the population was higher in the sibling vole than in the field vole at the end of the experiment. The presence of a coexisting species reduced the survival of field voles. Sibling voles, on the other hand, appeared to suffer more from intraspecific competition than interspecific competition. On a population level, the sibling vole seems to be a superior competitor in the absence of predators due to better survival and possibly a higher reproductive capacity. However, predation probably has a profound influence on the interspecific dynamics of these two species indicating that in natural surroundings apparent competition (i.e. competition via shared predators) is stronger than direct competition.  相似文献   

12.
Interspecific competition is usually understood as different species competing directly with each other for limited resources. However, predators can alter such competitive interactions substantially. Predation can promote the coexistence of species in a situation where it would otherwise be impossible, for example if a tradeoff between the competitive abilities and predation resistance of the prey species exists. The field vole Microtus agrestis and the sibling vole M. rossiaemeridionalis are sympatric grassland species, which compete for the same resources. At the population level sibling voles are suggested to be superior competitors to field voles, yet more vulnerable to predation. We tested the effects of predation on the two species in 0.5 ha outdoor enclosures by exposing vole populations to radio-collared freely-hunting least weasels Mustela nivalis nivalis for three weeks. Lethal and non-lethal impacts of predation limited population densities of both species during and after the experimental period, but the effect was more pronounced in sibling voles in which population densities decreased markedly during the treatment period and even after that. Field vole population densities remained stable under weasel predation, while densities increased in controls. Survival in both species was lower in treatment populations compared to controls, but the effect tended to be more pronounced in sibling voles and in females of both species. The average mass of adults in both species declined in the treatment populations. These results suggest that predation by least weasels can limit vole populations locally, even during favourable summer conditions, and have extended negative effects on the dynamics of vole populations. In addition, predation alleviated interspecific competition between the vole species and is, therefore, a potential factor enabling the coexistence of them.  相似文献   

13.
The cyclic population dynamics of vole and predator communities is a key phenomenon in northern ecosystems, and it appears to be influenced by climate change. Reports of collapsing rodent cycles have attributed the changes to warmer winters, which weaken the interaction between voles and their specialist subnivean predators. Using population data collected throughout Finland during 1986–2011, we analyse the spatio-temporal variation in the interactions between populations of voles and specialist, generalist and avian predators, and investigate by simulations the roles of the different predators in the vole cycle. We test the hypothesis that vole population cyclicity is dependent on predator–prey interactions during winter. Our results support the importance of the small mustelids for the vole cycle. However, weakening specialist predation during winters, or an increase in generalist predation, was not associated with the loss of cyclicity. Strengthening of delayed density dependence coincided with strengthening small mustelid influence on the summer population growth rates of voles. In conclusion, a strong impact of small mustelids during summers appears highly influential to vole population dynamics, and deteriorating winter conditions are not a viable explanation for collapsing small mammal population cycles.  相似文献   

14.
We studied the effects of removal of breeding nomadic avian predators (the kestrel, Falco tinnunculus and Tengmalm's owl, Aegolius funereus) on small mammals (voles of the genera Microtus and Clethrionomys and the common shrew, Sorex araneus) during 1989–1992 in western Finland to find out if these predators have a regulating or limiting impact on their prey populations. We removed potential breeding sites of raptors from five manipulation areas (c. 3 km2 each), whereas control areas had nest-boxes in addition to natural cavities and stick-nests. Densities of small mammals were monitored by snap-trapping in April, June, and August, and densities of mammalian predators (the least weasel, Mustela nivalis nivalis, the stoat, M. erminea and the red fox, Vulpes vulpes) by snow tracking in early spring and late autumn. The yearly mean number of raptor breeding territories was 0.2–1.0 in reduction areas and 3.0–8.2 in control areas. Breeding raptors alone did not regulate prey populations in the long term, but probably caused short-term changes in the population dynamics of both the main prey, the sibling vole (Microtus rossiaemeridionalis) and an alternative prey (the common shrew). The densities of an alternative prey, the bank vole (Clethrionomys glareolus) decreased in raptor reduction areas, most likely due to increased least weasel predation pressure in the absence of breeding avian predators.  相似文献   

15.
The alternative prey hypothesis predicts that predators respond both functionally and numerically (with a time lag) to fluctuations in the main prey abundance, which affects the survival of alternative prey. This pattern was found in northern Europe in the community formed by voles (Microtidae), red foxes (Vulpes vulpes) and roe deer (Capreolus capreolus). We studied the same predator—prey community in a temperate latitude where, according to the predation hypothesis, only the functional response of predators to changes in main prey availability should occur. In the years 1997–2007, in western Poland, we estimated the index of common vole (Microtus arvalis) abundance (burrow counts), the density of foxes (spotlight counts), the young production in foxes (young/adult ratio), the index of fox predation on fawns (prey remains near dens) as well as the reproduction index (fawn/female ratio) and density of roe deer (total counts). The vole abundance fluctuated considerably, the young production in foxes did not correlate with the main prey availability, but the density of foxes showed direct numerical response. The index of fox predation on fawns decreased with the vole abundance and negatively affected the fawn/female ratio in roe deer. Thus, the relationships between voles and foxes were not fully consistent with the predation hypothesis. The direct numerical response of foxes should tend to stabilize this predator—prey community. It is suggested, however, that responses showed by vole-eating predators in temperate latitudes may sometimes affect their alternative prey, including animals with unfavourable conservation status.  相似文献   

16.
VIDAR SELÅS 《Ibis》2006,148(4):678-686
According to the alternative prey hypothesis, autumn populations of ground-nesting game birds fluctuate in synchrony with vole numbers because generalist predators that mainly eat voles switch to alternative prey, such as eggs and chicks, when vole numbers decline. In hunting statistics from Nord-Trøndelag, central Norway, 1901–24, annual fluctuations in the number of Willow Grouse Lagopus lagopus and Western Capercaillie Tetrao urogallus , but not of Woodcock Scolopax rusticola , were positively related to vole numbers in the current year. Both Woodcock and grouse indices were related to hunting indices of Goshawk Accipiter gentilis and to weather variables assumed to influence the birds' survival or reproduction, suggesting that the indices actually reflected local population levels. Synchronous vole and grouse fluctuations are consistent with the alternative prey hypothesis (although predator densities were low in the early 1900s), but the asynchronous Woodcock fluctuations refute the hypothesis. Rather, because the Woodcock does not feed on plants utilized by voles and grouse, I suggest that food quality is the ultimate factor for the synchrony in vole and grouse numbers in Norway.  相似文献   

17.
There is a pressing need to understand how changing climate interacts with land‐use change to affect predator–prey interactions in fragmented landscapes. This is particularly true in boreal ecosystems facing fast climate change and intensification in forestry practices. Here, we investigated the relative influence of autumn climate and habitat quality on the food‐storing behaviour of a generalist predator, the pygmy owl, using a unique data set of 15 850 prey items recorded in western Finland over 12 years. Our results highlighted strong effects of autumn climate (number of days with rainfall and with temperature <0 °C) on food‐store composition. Increasing frequency of days with precipitation in autumn triggered a decrease in (i) total prey biomass stored, (ii) the number of bank voles (main prey) stored, and (iii) the scaled mass index of pygmy owls. Increasing proportions of old spruce forests strengthened the functional response of owls to variations in vole abundance and were more prone to switch from main prey to alternative prey (passerine birds) depending on local climate conditions. High‐quality habitat may allow pygmy owls to buffer negative effects of inclement weather and cyclic variation in vole abundance. Additionally, our results evidenced sex‐specific trends in body condition, as the scaled mass index of smaller males increased while the scaled mass index of larger females decreased over the study period, probably due to sex‐specific foraging strategies and energy requirements. Long‐term temporal stability in local vole abundance refutes the hypothesis of climate‐driven change in vole abundance and suggests that rainier autumns could reduce the vulnerability of small mammals to predation by pygmy owls. As small rodents are key prey species for many predators in northern ecosystems, our findings raise concern about the impact of global change on boreal food webs through changes in main prey vulnerability.  相似文献   

18.
Mechanisms generating the well-known 3-5 year cyclic fluctuations in densities of northern small rodents (voles and lemmings) have remained an ecological puzzle for decades. The hypothesis that these fluctuations are caused by delayed density-dependent impacts of predators was tested by replicated field experimentation in western Finland. We reduced densities of all main mammalian and avian predators through a 3 year vole cycle and compared vole abundances between four reduction and four control areas (each 2.5-3 km(2)). The reduction of predator densities increased the autumn density of voles fourfold in the low phase, accelerated the increase twofold, increased the autumn density of voles twofold in the peak phase, and retarded the initiation of decline of the vole cycle. Extrapolating these experimental results to their expected long-term dynamic effects through a demographic model produces changes from regular multiannual cycles to annual fluctuations with declining densities of specialist predators. This supports the findings of the field experiment and is in agreement with the predation hypothesis. We conclude that predators may indeed generate the cyclic population fluctuations of voles observed in northern Europe.  相似文献   

19.
Summary We studied responses of stoats and least weasels to fluctuating vole abundances during seven winters in western Finland. Density indices of mustelids were derived from snow-tracking, diet composition from scat samples, and vole abundances from snap-trapping. Predation rate was estimated by the ratio of voles to mustelids and by the vole kill rate by predators (density of predator x percentage of voles in the diet). We tested the following four predictions of the hypothesis that small mustelids cause the low phase of the microtine cycle. (1) The densities of predators should lag well behind the prey abundances, as time lags tend to have destabilizing effects. The densities of stoats fluctuated in accordance with the vole abundances, whereas the spring densities of least weasels tracked the vole abundances with a half-year lag and the autumn densities with a 1-year lag. (2) Predators should not shift to alternative prey with declining vole densities. The yearly proportion of Microtus voles (the staple prey) in the diet of stoats varied widely (range 16–82%) and was positively correlated with the winter abundance of these voles. In contrast, the same proportion in the food of least weasels was independent of the vole abundance. (3) The ratio of voles to small mustelids should be smallest in poor vole years and largest in good ones. This was also observed. (4) Vole densities from autumn to spring should decrease more in those winters when vole kill rates are high than when they are low. The data on least weasels agreed with this prediction. Our results from least weasels were consistent with the predictions of the hypothesis, but stoats behaved like semi-generalist predators. Accordingly, declines and lows in the microtine cycle may be due to least weasel predation, but other extrinsic factors may also contribute to crashes.  相似文献   

20.
Competing species benefit from eavesdropping on each other's signals by learning about shared resources or predators. But conspicuous signals are also open to exploitation by eavesdropping predators and should also pose a threat to other sympatric prey species. In western Finland, sibling voles Microtus rossiameridionalis and field voles M. agrestis compete for food and space, and both species rely upon scent marks for intraspecific communication. Both vole species are prey to a range of terrestrial scent hunting predators such as least weasels, however, the competitively superior sibling voles are taken preferentially. We tested in large out‐door enclosures whether field voles eavesdrop on the signals of its competitor, and whether they behave as though this eavesdropping carries a risk of predation. We presented field voles with scent marks from unknown conspecifics and sibling voles and measured their visitation, activity and scent marking behaviours at these scents under high (weasel present) and low (weasel absent) predation risk. Field voles readily visited both field and sibling vole scents under both high and low predation risk; however their activity at sibling vole scent marks declined significantly under increased predation risk. In contrast, predation risk did not affect field voles’ activity at conspecific scents. Thus, field voles were compelled to maintain eavesdropping on heterospecific scents under an increased risk of predation, however they compensated for this additional risk by reducing their activity at these risky scents. Scent marking rates declined significantly under high predation risk. Our results therefore reveal a hidden complexity in the use of social signals within multi‐species assemblages that is clearly sensitive to the potential for increased predation risk. The predation risks of interspecific eavesdropping demonstrated here represents a significant generalisation of the concept of associational susceptibility.  相似文献   

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