Numerical response of small mustelids to vole abundance: delayed or not?

One of the most studied problems in population ecology has been to understand the relative roles of top–down and bottom–up forces in regulating animal populations. This has also been a key issue in studies of vole population dyna mics. Vole populations exhibit a wide variation of dynamics, from seasonal fluctuations to multiannual variations or cyclicity. One of the hypotheses to explain cyclic population dynamics is predation by the specialist predators. A common counterargument against the predation hypothesis has been the lack of conclusive observations of the time delay in the predators’ numerical response. We studied the interaction between voles and their specialist small mustelid predators, the stoat Mustela erminea and the least weasel Mustela n. nivalis, by modelling their interaction to data sets that cover large areas of Finland. Vole abundance was monitored with biannual trappings and their predators with snow‐tracking. Results show a high dependence of the predators on the voles, and this connection is generally tighter in weasels than in stoats. Weasel abundance is affected most strongly by the vole abundance in previous spring, 8.5– 10 months earlier, while in stoats the effect of autumn abundance of voles, 2.5–6 months earlier, was the strongest. These results, together with the observation that the weasels’ effects on voles are stronger after a time lag of 6–9.5 than 2–4.5 months, indicate the existence of a time lag in weasels’ numerical response. A time lag in the predators’ numerical response is a necessary condition for the predators to drive population cycles in its prey, and therefore our results support the specialist predation hypothesis.     

Seasonal changes in the numerical responses of predators to cyclic vole populations

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.     

Vole population cycles in northern and southern Europe: is there a need for different explanations for single pattern?

1. Students of population cycles in small rodents in Fennoscandia have accumulated support for the predation hypothesis, which states that the gradient in cycle length and amplitude running from southern to northern Fennoscandia reflects the relative influence of specialist and generalist predators on vole dynamics, itself modulated by the presence of snow cover. The hypothesized role of snow cover is to isolate linked specialist predators, primarily the least weasel, Mustela n. nivalis L. and their prey, primarily field voles Microtus agrestis L., from the stabilizing influence of generalist predators. 2. The predation hypothesis does not readily account for the high amplitude and regular 3-year cycles of common voles documented in agricultural areas of western, central and eastern Europe. Such cycles are rarely mentioned in the literature pertaining to Fennoscandian cycles. 3. We consider new data on population cycles and demographic patterns of common voles Microtus arvalis Pallas in south-west France. We show that the patterns are wholly consistent with five of six patterns that characterize rodent cycles in Fennoscandia and that are satisfactorily explained by the predation hypothesis. They include the: (a) existence of cycle; (b) the occurrence of long-term changes in relative abundance and type of dynamics; (c) geographical synchrony over large areas; (d) interspecific synchrony; and (e) voles are large in the increase and peak phase and small in decline and low phase, namely. There is a striking similarity between the patterns shown by common vole populations in south-west France and those from Fennoscandian cyclic rodent populations, although the former are not consistent with a geographical extension of the latitudinal gradient south of Fennoscandia. 4. It is possible that the dominant interaction leading to multiannual rodent oscillations is different in different regions. We argue, however, that advocates of the predation hypothesis should embrace the challenge of developing a widely applicable explanation to population cycles, including justifying any limits to its applicability on ecological and not geographical grounds.     

Responses of stoats and least weasels to fluctuating food abundances: is the low phase of the vole cycle due to mustelid predation?

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.     

Species‐specific limitation of vole population growth by least weasel predation: facilitation of coexistence?

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.     

Disease dynamics in cyclic populations of field voles (Microtus agrestis): cowpox virus and vole tuberculosis (Mycobacterium microti)

The possible role of pathogens in rodent population cycles has been largely neglected since Elton's 'epidemic hypothesis' of 1931. To revisit this question, 12 adjacent, cyclic but out-of-phase populations of field voles (Microtus agrestis) in North East England were studied and the initial results are presented here. The prevalences of antibodies to cowpox virus and of clinical signs of Mycobacterium microti infection (vole tuberculosis) showed delayed (not direct) density dependence (with a lag of three to six months). This did not result from changes in population structure, even though there were such changes associated with the different phases of the cycle. The prevalences rose as vole numbers rose, and peaked as numbers declined. The apparent lag in the numerical response of infection prevalence to changes in host abundance is consistent with the hypothesis that diseases, singly or in combination, play a hitherto underestimated role in the dynamics of cyclic populations.     

The impact of short-term predator removal on vole dynamics in an arctic-alpine landscape

During 1991–95, mammalian predators (weasel, Mustela nivalis , stoat, M. erminea , mink, M. vison , and red fox, Vulpes vulpes ) were excluded in late summers from a 2 ha piece of a north Norwegian mountain slope. The exclosure extended from an outpost of luxuriant sub-arctic birch forest to a typical arctic–alpine habitat complex, including productive willow scrublands, tundra heaths, dry ridges and snow-beds. The exclosure thus encompassed the entire range of habitat conditions encountered in a typical north Fennoscandian mountain and tundra landscapes. During 1991–95, the exclosure was predator-proof from late July to late September. In wintertime and in early summer, the exclosure was accessible to mammalian predators. Vole dynamics in the short-term exclosure were compared to dynamics in five reference areas with similar habitat conditions.
In 1991, when vole densities were rising in the area, neither collective vole densities nor densities of individual vole species differed significantly between the exclosure and the replicated controls. Spring densities of voles were never significantly different between the exclosure and the controls. With respect to autumnal densities of voles, however, the exclosure was a statistical outlier in the peak year 1992 and throughout the gradual decline phase of 1993–95. In the peak year, the difference in collective vole densities was modest (30%), but increased to two-fold during the first two decline years and was almost four-fold in the crash year of 1995. The strongest response was displayed by field voles ( Microtus agrestis ), hypothesized to be the pivotal prey species of weasels, especially by females and young individuals, i.e. by the functional categories especially sensitive to mammalian predation. These results are consistent with the hypothesis that predation plays a pivotal role for the regulation of herbivorous mammals in relatively productive arctic–alpine habitats.     

Multiple predators induce risk reduction in coexisting vole species

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.     

Dynamic effects of predators on cyclic voles: field experimentation and model extrapolation

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.     

An empirically based model for latitudinal gradient in vole population dynamics

Vole dynamics in northern Europe exhibit a well-defined geographical gradient, with oscillatory populations being confined to high latitudes. It has been proposed that oscillations in northern vole populations are driven by their interaction with specialist predators (weasels), while the more southern rodent populations are relatively stable because of regulation by generalist predators. We tested this generalist/specialist predation hypothesis by constructing an empirically based model for vole population dynamics, estimating its parameters, and making predictions about the quantitative pattern of the latitudinal shift in vole dynamics. Our results indicated that the model accurately predicted the latitudinal shift in the amplitude and periodicity of population fluctuations. Moreover, the model predicted that vole dynamics should shift from stable to chaotic as latitude is increased, a result in agreement with nonlinear time-series analysis of the data. The striking success of the model at predicting the shifts in amplitude and stability along the geographical gradient in northern Europe provides strong support for the key role of specialist and generalist predators in vole population dynamics.     

Habitat requirements of weasels <Emphasis Type="Italic">Mustela nivalis</Emphasis> constrain their impact on prey populations in complex ecosystems of the temperate zone

Differences in habitat use by prey and predator may lead to a shift of occupied niches and affect dynamics of their populations. The weasel Mustela nivalis specializes in hunting rodents, therefore habitat preferences of this predator may have important consequences for the population dynamics of its prey. We investigated habitat selection by weasels in the Bia?owie?a Forest in different seasons at the landscape and local scales, and evaluated possible consequences for the population dynamics of their prey. At the landscape scale, weasels preferred open habitats (both dry and wet) and avoided forest. In open areas they selected habitats with higher prey abundance, except during the low-density phase of the vole cycle, when the distribution of these predators was more uniform. Also in winter, the distribution of weasels at the landscape scale was proportional to available resources. In summer, within open dry and wet habitats, weasels preferred areas characterised by dense vegetation, but avoided poor plant cover. In winter, weasels used wet open areas proportionally to availability of habitats when hunting, but in contrast to summer, they rested only in habitats characterized by a lower water level, which offered better thermal conditions. At the local scale, the abundance of voles was a less important factor affecting the distribution of these predators. Although we were not able to provide direct evidence for the existence of refuges for voles, our results show that they may be located within habitat patches, where availability of dense plant cover and physiological constraints limit the activity of weasels. Our results indicate that in complex ecosystems of the temperate zone, characterized by a mosaic pattern of vegetation types and habitat specific dynamics of rodents, impact of weasels on prey populations might be limited.     

Spatio-temporal patterns of habitat use in voles and shrews modified by density, season and predators

1.?Although the intrinsic habitat preferences of a species can be considered to be fixed, the realized habitat use depends on the prevailing abiotic and biotic conditions. Often the core habitats are occupied by dense and stable populations, while marginal habitats become occupied only at times of high density. In a community of interacting species, habitat uses of different species become inter-related, for example an increased density of a strong competitor forcing a weaker competitor to use more marginal habitats. 2.?We studied the spatio-temporal distribution patterns of three common small mammal species, the bank vole Myodes glareolus; the field vole Microtus agrestis; and the common shrew Sorex araneus, in a 4-year trapping study carried out on six large islands, each containing a mixture of three main habitat types (forest, field and clear-cut). We experimentally released least weasels (Mustela n. nivalis) to some of the islands to see how the focal species respond to increased predation pressure. 3.?Both vole species were largely restricted to their core habitats (bank voles to forests and field voles to fields) at times of low population density. With increasing density, the relative habitat use of both species increased in the clear-cut areas. The common shrew was a generalist in its habitat use at all population densities. 4.?The release of the weasels changed the habitat use of all study species. 5.?The vole species showed a stronger aggregated pattern than the common shrew, especially at low population density. The vole aggregations remained in the same localities between seasons, except in the case of bank voles after the weasels were released. 6.?Bank voles and field voles avoided each other at high density. 7.?We conclude that intrinsically differential habitat requirements and flexibility to modify habitat use facilitate the coexistence of the two competing vole species in mosaic landscapes consisting of boreal forests and open habitats.     

Large-scale spatial and temporal patterns in population dynamics of the stoat, Mustela erminea, and the least weasel, M. nivalis, in Finland

The stoat and the least weasel are specialist predators of small rodents, and therefore their numbers are likely to depend on the availability of voles. These small predators are ecologically alike, but they differ somewhat in their diet. The stoat is larger in size than the least weasel and therefore capable of using a wider variety of prey species, while the least weasel is more restricted to small mammals. Voles in northern Fennoscandia exhibit cyclic dynamics of 3–5 years with large-scale spatial synchrony and geographical trends in cycle length and amplitude. We predicted that the cyclic dynamics of voles are reflected in the dynamics of their predators with slight differences between the stoat and the least weasel. In this study we use snow-tracking data to characterize the dynamics of small mustelids. The data were collected from different parts of Finland using permanent triangle-shaped census routes of 12 km in 1989 to 2003. Population fluctuations of small mustelids were generally multiannually periodic and in synchrony over large areas, but we did not find any clear geographical gradient in the attribute of small mustelid dynamics comparable to those observed in vole population fluctuations. Instead, we found a similar decreasing temporal trend in the abundances of both species as has been recently reported for voles.     

Predation rate, prey preference and predator switching: experiments on voles and weasels

We studied the predation rate and prey selection of the least weasel ( Mustela nivalis nivalis ) on its two most common prey species in boreal environments, the bank vole ( Clethrionomys glareolus ) and the field vole ( Microtus agrestis ), in large outdoor enclosures. We also studied the response of weasels to odours of the two species in the laboratory. The enclosure experiment was conducted using constant vole densities (16 voles/ha) but with varying relative abundance of the two species. Weasels showed higher predation rates on bank voles, and males had higher predation rate than females. Females killed disproportionately more of the more abundant prey species, but they preferred bank voles to field voles when both were equally available. Overall, the predation rate also increased with increasing abundance of bank voles. Therefore our results are in agreement with earlier laboratory results showing preference for bank voles, even if no intrinsic preference for odours of either species was observed in our laboratory study. We suggest that the least weasel hunts according to prey availability, prey aggregation and suitability of hunting habitat, and that this causes the observed dependence of least weasels on field voles and emphasises the role of the field vole in the vole-weasel interaction in cyclic vole populations. Furthermore, our results suggest that predation by weasels may facilitate the coexistence of the two vole species via predator switching, and that it may cause the observed synchrony in dynamics between vole species.     

Predator–vole interactions in northern Europe: the role of small mustelids revised

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.     

Population changes of different predators during a water vole cycle in a central European mountainous habitat

Population changes in long-eared owls Asio otus , polecats Mustela putorius , red foxes Vulpes vulpes , stone martens Martes foina and badgers Meles meles were monitored during a water vole Arvicola terrestris scherman cycle in western Switzerland. Long-eared owls confirmed their status of highly mobile specialist predators in responding strongly and without time lag to water vole population changes. Even though polecats are considered generalists, they exhibited also a strong response to water vole fluctuations. Their numbers tracked water vole densities with a 1-yr time lag. Marked population changes were also recorded in red foxes and stone martens, but these changes were not related to water vole densities. Lastly, badgers did not show any significant population changes during the water vole cycle. We discuss the possible reasons for these differences and conclude that multi-factorial approach is clearly required to understand population processes in predator-prey systems.     

Patterns in grouse and Woodcock Scolopax rusticola hunting yields from central Norway 1901–24 do not support the alternative prey hypothesis for grouse cycles

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.     

Do predators limit the abundance of alternative prey? Experiments with vole‐eating avian and mammalian predators

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.     

Predation on two vole species by a shared predator: antipredatory response and prey preference

In prey communities with shared predators, variation in prey vulnerability is a key factor in shaping community dynamics. Conversely, the hunting efficiency of a predator depends on the prey community structure, preferences of the predator and antipredatory behavioural traits of the prey. We studied experimentally, under seminatural field conditions, the preferences of a predator and the antipredatory responses of prey in a system consisting of two Myodes species of voles, the grey-sided vole (M. rufocanus Sund.) and the bank vole (M. glareolus Schreb.), and their specialist predator, the least weasel (Mustela nivalis nivalis L.). To quantify the preference of the weasels, we developed a new modelling framework that can be used for unbalanced data. The two vole species were hypothesised to have different habitat-dependent vulnerabilities. We created two habitats, open and forest, to provide different escape possibilities for the voles. We found a weak general preference of the weasels for the grey-sided voles over the bank voles, and a somewhat stronger preference specifically in open habitats. The weasels clearly preferred male grey-sided voles over females, whereas in bank voles, there was no difference. The activity of voles changed over time, so that voles increased their movements immediately after weasel introduction, but later adjusted their movements to times of lowered predation risk. Females that were more active had an elevated mortality risk, whereas in the case of males, the result was the opposite. We conclude that, in vulnerability to predation, the species- or habitat-specific characteristics of these prey species are playing a minor role compared to sex-specific characteristics.     

Population dynamics in a cyclic environment: consequences of cyclic food abundance on tawny owl reproduction and survival

1. Understanding which factors regulate population dynamics may help us to understand how a population would respond to environmental change, and why some populations are declining.
2. In southern Finland, vole abundance shows a three-phased cycle of low, increase and decrease phases, but these have been fading out in recent years. During five such cycles (1981–1995), all tawny owls Strix aluco were censused in a 250-km² study area, and their reproduction and survival were monitored.
3. Males and females showed similar dynamics, but experienced breeders recruited more offspring and had higher survival than first breeders. Offspring recruitment, but not survival of breeding individuals varied in accordance with vole abundance.
4. The population's numerical response to prey abundance was primarily due to first-breeding individuals entering the population in the increase phase when immigration was the highest. First-breeding birds were younger, but experienced breeders were older in more favourable vole years.
5. A stage-specific matrix population model integrating survival and fecundity showed that, despite obvious variation in fecundity between vole cycle phases, this variation had limited importance for overall tawny owl population dynamics, but that the survival of experienced breeders during the low phase is most important for population growth.
6. Model and data agreed that the vole cycle drives the dynamics of this avian predator by limiting the recruitment of new breeders during the low phase. Population dynamics hence differ not only from the classic example of the species in a more temperate region in the UK where the number of territories is stable across years, but also from the dynamics of other avian vole predators in Fennoscandia where the recurring crash in vole abundance drastically lowers adult survival thereby creating vacancies.