Population limitation of the northern red-backed vole in the boreal forests of northern Canada

1. Across the vast boreal forests of North America, no population cycles in Clethrionomys species occur. In Eurasia, by contrast, some Clethrionomys populations of the same species undergo regular 3-5-year cycles. We examined the effects of nutrients, food, competitors, predators and climate on population limitation in the northern red-backed vole (Clethrionomys rutilus Pallas) in the south-western Yukon to determine why this difference occurs. 2. From 1986 to 1996 we added food, reduced large mammal predators and excluded snowshoe hares (Lepus americanus Erxleben) from large plots and found that none of these manipulations affected red-backed vole abundance. Adding nutrients as nitrogen, phosphorus and potassium (NPK) fertilizer had a slight negative effect, probably acting through a reduction in dwarf shrub productivity caused by competition from grasses. 3. We monitored weasel populations directly through trapping and indirectly through snow tracking. Predation by these vole specialists was irrelevant as a limiting factor most of the time because voles in this area do not reach the densities needed to sustain weasel populations. Other boreal forest mammal and bird predators did not focus on red-backed voles. However, when red-backed vole populations increased in the forest and Microtus voles also increased in the meadows, weasel populations increased and may have temporarily depressed red-backed voles in winter. 4. We monitored one major potential food, white spruce seeds, but seed fall was not related to population changes in red-backed voles, even after mast years. 5. We assessed the impact of weather variables, and the average depth of the snow pack during winter (October-March) was correlated directly with vole demography, having both direct effects in that year and delayed effects in the following year. 6. Our long-term trapping data (1973-96) indicate that Clethrionomys populations fluctuated, with peaks following hare peaks by 2-3 years. 7. We propose that the key variable limiting these vole populations is overwinter survival, and this is a function of overwinter food from berries produced during the previous summer by dwarf shrubs. These shrubs may be stimulated by abundant moisture from winter snows or by periodic fertilization from large quantities of pellets produced at snowshoe hare peaks.     

Specialist and generalist natural enemies as an explanation for geographical gradients in population cycles of northern herbivores

Within Fennoscandia, two well-studied groups of herbivores exhibit clear geographical gradients in their population dynamics. Populations of a forest lepidopteran ( Epirrita autumnata , the autumnal moth) and voles of the genera Microtus and Clethrionomys show pronounced multi-annual cycles in the north but become more stable towards the south. Here we review empirical and theoretical studies on these species, mainly regarding the biological mechanisms that are assumed to generate the pattern of population dynamics in both systems. We conclude that the specialist/generalist predation hypothesis offers a common explanation for the population cycles and their geographical gradients irrespective of whether a herbivorous insect or small mammals are concerned. According to this hypothesis, originally developed for the Fennoscandian voles, but now applied also to E. autumnata , population cycles are generated by specialist natural enemies (predators for the voles and parasitoids for E. autumnata ). Furthermore, the dynamic shift from cycles to stability is assumed to be caused by an increase in the density and diversity of generalist natural enemies from north to south in Fennoscandia.     

Bot fly parasitism of the red-backed vole: host survival,infection risk,and population growth

Parasites can play an important role in the dynamics of host populations, but empirical evidence remains sparse. We investigated the role of bot fly (Cuterebra spp.) parasitism in red-backed voles (Myodes gapperi) by first assessing the impacts of the parasite on the probability of vole survival under stressful conditions as well as on the reproductive activity of females. We then identified the main factors driving both the individual risk of infection and the abundance of bot flies inside red-backed voles. Finally, we evaluated the impacts of bot fly prevalence on the growth rate of vole populations between mid-July and mid-August. Thirty-six populations of red-backed voles were sampled in the boreal forest of Québec, Canada. The presence and the abundance of parasites in voles, two host life history traits (sex and body condition), three indices of habitat complexity (tree basal area, sapling basal area, coarse woody debris volume), and vole abundance were considered in models evaluating the effects of bot flies on host populations. We found that the probability of survival of red-backed voles in live traps decreased with bot fly infection. Both the individual risk of infection and the abundance of bot flies in red-backed voles were driven mainly by vole abundance rather than by the two host life history traits or the three variables of habitat complexity. Parasitism had population consequences: bot fly prevalence was linked to a decrease in short-term growth rate of vole populations over the summer. We found that bot flies have the potential to reduce survival of red-backed voles, an effect that may apply to large portions of populations. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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.     

Spatio‐temporal covariation in abundance between the cyclic common vole Microtus arvalis and other small mammal prey species

Populations of the common vole Microtus arvalis in mid‐western France show cyclic dynamics with a three‐year period. Studies of cyclic vole populations in Fennoscandia have often found inter‐specific synchrony between the voles and other small mammals which share the voles' predators. Although predators are central to the favoured mechanism to explain Fennoscandian vole cycles and the spatial variation of small mammal populations, their role in vole cycles elsewhere, including France, is less clear. Establishing whether alternative prey species in France cycle in parallel with voles as they do in Fennoscandia is thus an important step towards understanding the generality of predators' influence on cyclic vole populations. We applied spatial and temporal autocorrelation and cross‐correlation methods to French populations of M. arvalis and two sympatric non‐cyclic small mammal species, Apodemus sylvaticus and Crocidura russula. Patterns of time‐lagged cross‐correlation between the abundance of M. arvalis and the other two species suggested synchrony in their dynamics beyond that expected of stochastic environmental variation, and indicated a weak three‐year cycle in A. sylvaticus and C. russula that was in phase with that of M. arvalis. We interpret the synchrony between these species as the effect of shared predators and environmental stochasticity. Abundance within species showed weak spatial autocorrelation in June at scales consistent with dispersal being the mechanism responsible, but a more general lack of spatial structure within and between species was consistent with the strong spatial synchrony at regional scales often found in fluctuations of small mammal abundance.     

Does risk of predation by mammalian predators affect the spacing behaviour of rodents? Two large-scale experiments

Predator-prey interactions between small mammals and their avian and mammalian predators have attracted much attention. However, large-scale field experiments examining small-mammal antipredatory responses under the risk of predation by mammals are rare. As recently pointed out, the scale of experiments may cause misleading results in studies of decision-making under predation risk. We studied the effect of small mustelid predators on the spacing behaviour of the gray-tailed vole (Microtus canicaudus) and the bank vole (Clethrionomys glareolus) in two separate field enclosure experiments. The experiments were conducted during the breeding season in North America and northern Europe, where small mustelids have been suggested to be important mammalian predators of voles. As in most of the earlier laboratory studies, predation risk was simulated using fresh mustelid faeces and urine. This made it possible to compare the results from experiments at different spatial scales. We did not find any effect of increased predation risk on spacing behaviour (mean and/or maximum distance moved and home range size) or trappability in either vole species. Simulated predation risk did not affect the breeding of females in gray-tailed voles, as has previously been shown in bank voles. The results disagree with most of the studies conducted in laboratory conditions with small mammals. We discuss whether this discrepancy could be caused by differences in the scale of the experiments. Received: 12 April 1999 / Accepted: 7 October 1999     

Genetic similarity among Eurasian subspecies of boreal owls Aegolius funereus

Boreal owls Aegolius funereus (referred to as Tengmalm's owls in Europe) breed in boreal forests throughout the Holarctic region and in high-elevation subalpine forests further south. They are currently classified as seven subspecies; six found throughout Eurasia, and one in North America. The geographic distribution of boreal owls in North America and Eurasia is similar, as are their patterns of dispersal and irruption. Because a recent genetic study of boreal owls in North America found very little genetic differentiation among widely disparate locations, we expected that boreal owls in Eurasia similarly would have very little genetic differentiation. Using seven microsatellite markers, we analyzed genetic samples from 275 boreal owls in North America, 36 in Norway, and five in eastern Russia. We found no detectable genetic differentiation between Norwegian and Russian owls, but notable differentiation between North American and Eurasian owls. Low intra-continental genetic differentiation likely results from high rates of long-distance dispersal among subpopulations of boreal owls. In light of these results, we recommend further genetic sampling of boreal owls throughout Eurasia in order to determine whether six separate subspecies here are warranted.     

Gradients in population fluctuations of Tengmalm's owl Aegolius funereus in Europe

Summary I studied the importance of geographical location, snow cover and food to the fluctuations in 30 breeding populations of Tengmalm's owl (Aegolius funereu) in Europe. Cyclicity indices were positively related both to latitude and longitude, but within Fennoscandia they were better correlated with snow cover. Population fluctuations increased northward, while food niche breadth and degree of site tenacity decreased northwards.Microtine fluctuations become more pronounced northwards and are more synchronized, while number of alternative prey increases southwards. These factors promote instability in North European and stability in central European owl populations. Furthermore, snow conditions were more important within Fennoscandia, since this small owl cannot hunt voles protected by deep snow. Environmental predictability and diversity of available food for Tengmalm's owl increase southwards in Europe. Thus, the owl is a resident generalist predator of small mammals and birds in central Europe and adopts a partial migration strategy (males being resident and females nomadic) in South and West Finland, changing its habits to nomadic microtine specialist in areas with pronounced vole cycles (in northern Fennoscandia). These changes fit well with the recent suggestion that gradients in density variations of small rodents are related to sustainable numbers of generalist predators.     

Is the heart of Fennoscandian rodent cycle still beating? A 14‐year study of small mammals and Tengmalm's owls in northern Norway

Analyses of spatial and temporal patterns of the small mammal cycle in Fennoscandia have led to two main conclusions: a south-north geographical gradient in the strength of density dependence and period length, and a change in temporal dynamics with less clear periodicity and lower amplitude of fluctuations in recent years. Fourteen years (1985–1998) of small mammals trapping and Tengmalm's owls monitoring data provided clear evidence for 3-yr cycles of two vole species in inner Troms, north Norway, at the same latitude and only 80 km from Kilpisjärvi, north Finland, where 4–5-yr cycles have been the norm until the end of 1980s. The response of Tengmalm's owls to variation in small rodents abundance was not delayed, as was observed in central Finland. Common shrews appeared to have more stable dynamics. The features of the cycle, and a reassessment of the previously described patterns, show that the dynamics may be different over short distances. Furthermore, snow cover and habitat fragmentation should explicitly be taken into account in any understanding of the Fennoscandian gradient, and surrogate variables such as latitude should be avoided. We stress the need for a more extensive coverage of spatial and temporal patterns in cyclicity with respect to assumed ecological causes.     

Qualitative analysis of a predator-prey system with limit cycles

Fairly regular multiannual microtine rodent cycles are observed in boreal Fennoscandia. In the southern parts of Fennoscandia these multiannual cycles are not observed. It has been proposed that these cycles may be stabilized by generalist predation in the south.We show that if the half-saturation of the generalist predators is high compared to the number of small rodents the cycles are likely to be stabilized by generalist predation as observed. We give examples showing that if the half-saturation of the generalist predators is low compared to the number of small rodents, then multiple equilibria and multiple limit cycles may occur as the generalist predator density increases.     

SHORT-TERM DEMOGRAPHIC RESPONSE OF THE RED-BACKED VOLE TO SPRUCE BEETLE INFESTATIONS IN ALASKA

Abstract: How small mammals are affected by habitat changes caused by forest insect epidemics is largely unknown. Our objective was to determine the influence of spruce beetle (Dendroctonus rufipennis) epidemics on the dynamics of northern red-backed vole (Clethrionomys rutilus) populations approximately 10 years post-infestation. We conducted a mark-recapture study on northern red-backed voles for 2 field seasons in the Copper River Basin, Alaska, USA, where recent beetle infestations were widespread. Using the robust sampling design, we produced estimates of vole abundance, survival, and recruitment in 3 locations that varied in their degree of beetle-induced spruce mortality. Vole abundance inversely related to the level of spruce mortality. Vole recruitment showed a larger contribution from both immigration and in situ reproduction in the low infestation site than in the medium and heavy infestation sites. No differences in vole survival were detectable across the 3 locations with varied beetle-induced spruce mortality levels. Measured vole food resources and protective vegetative cover did not vary greatly across infestation levels. Abundance and recruitment parameters indicate a negative change induced by spruce beetle infestations. However, the effect of beetles was not large enough to cause the variation in vole survival. Spruce mortality levels may need to be over 50% before greatly influencing the habitat and the demographics of northern red-backed voles.     

Increased autumn rainfall disrupts predator–prey interactions in fragmented boreal forests

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.     

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.     

Diet variation of common buzzards in Finland supports the alternative prey hypothesis

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.     

Population fluctuations of the wood lemming<Emphasis Type="Italic">Myopus schisticolor</Emphasis> in eastern and western Finland

Olavi Eskelinen, Pertti Sulkava and Risto Sulkava From 1982 to 2003 we studied fluctuations in populations of the wood lemmingMyopus schisticolor (Liljeborg, 1844) in the Heinävesi (eastern Finland) and Keuruu regions (western Finland) by counting field signs and dead animals in standardized field surveys. We compared the population fluctuations of lemmings to those of other voles, owls and small mustelids in these regions. The lemming population in Heinävesi fluctuated regularly in 3-year cycles and in synchrony with the field vole population. Populations of owls also fluctuated in synchrony with wood lemming and field vole populations. In the Keuruu region, oscillations in the wood lemming population were irregular, and neither lemming and vole populations nor lemming and owl populations were correlated. Although direct mechanistic evidence is lacking, specialist predators such as weasels and owls probably cause the cyclicity in the field vole and wood lemming populations in the Heinävesi area. On the other hand, scarcity of high-quality habitats, unfavourable winter weather conditions and generalist predators may prevent the development of cyclicity in the wood lemming population of Keuruu.     

Small mammal responses to moose supplementary winter feeding

Supplementary feeding of wild large herbivores is a widespread practice in North America and Europe. The presence of feeding stations may have ecological consequences through changes to animal distributions, patterns of herbivory and a net nutrient input into the ecosystem. In Fennoscandia, supplementary feeding of moose in winter (Alces alces) is increasing. Although it has been shown to affect bird communities, its effects on small mammal communities were unknown. Here, we studied the effects of moose supplementary feeding stations on plants and on abundance, reproduction, and biomass of small mammals in years with low and high vole abundance. We sampled small mammals with snap traps and conducted surveys of the field layer vegetation, at varying distances from moose supplemental feeding stations. Due to the vegetation changes induced by feeding stations, abundance of common shrews (Sorex araneus) and Microtus voles were positively affected by long-term moose winter feeding, while bank voles (Myodes glareolus) were not affected. Moose feeding stations did not affect reproduction, individual body mass, or the total biomass of small mammals. Moose winter-feeding stations have impacts on nontarget species, providing islands of preferred grass and forb habitat for Microtus spp. and common shrews, allowing them to penetrate into a matrix of less preferred forest habitat.     

Gene conversion in the mitochondrial genome on interspecific hybridization in voles of the <Emphasis Type="Italic">Clethrionomys</Emphasis> genus

The phenomenon of interspecific hybridization accompanied by transfer of the mitochondrial genome from the northern red-backed vole (Clethrionomys rutilus) to the bank vole (Cl. glareolus) in northeastern Europe is well known already for 25 years. However, the possibility of recombination between homologous segments of maternal and paternal mtDNAs of the voles during fertilization was not previously studied. Analysis of data on variability of nucleotide sequences of the mitochondrial gene for cytochrome b in populations of red-backed and bank voles in the area of their sympatry has shown that as a result of interspecific hybridization, the mitochondrial gene pool of bank voles contains not only mtDNA haplotypes of red-backed vole females, but also mtDNA haplotypes of bank voles bearing short nucleotide tracts of red-backed vole mtDNA. This finding supports the hypothesis that an incomplete elimination of red-backed vole paternal mtDNA during the interspecific hybridization between bank vole females and red-backed vole males leads to the gene conversion of bank vole maternal mtDNA tracts by homologous ones of mtDNA of red-backed vole males.     

Mycophagy by small mammals in the coniferous forests of North America: nutritional value of sporocarps of Rhizopogon vinicolor, a common hypogeous fungus

We evaluated the nutritional value of sporocarps of Rhizopogon vinicolor, a common hypogeous fungus in the coniferous forests of North America, for two small mammal species: the Californian red-backed vole (Clethrionomys californicus) and the northern flying squirrel (Glaucomys sabrinus). Although the nitrogen concentration of sporocarps was high, much of it was in non-protein form or associated with cell walls, suggesting that it may be of low nutritional value or protected from mammalian digestive enzymes. Sporocarps also had high concentrations of cell wall constituents, indicating low availability of digestible energy. When fed a diet of this fungus alone in a controlled feeding experiment both mammal species lost a small amount of body mass. Digestibilities of dry matter, nitrogen, cell wall constituents and energy from sporocarps by both species were lower than the digestibilities of other food types by other similarly sized small mammals. Red-backed voles digested the various components of sporocarps at least as well as the flying squirrels, even though they were almost six-fold smaller in body mass. This observation supports the notion that red-backed voles, like other microtine rodents, have morphological and physiological adaptations of the digestive system that are postulated to permit greater digestion of fibrous diets than predicted on the basis of body size. Despite this, our results re-affirm previous conclusions that hypogeous fungi are only of moderate nutritional value for most small, hindgut-fermenting mammals. Future studies should focus on the importance of mixed-species of fungi in the diet of small mammalian mycophagists. Accepted: 4 December 1998     

Habitat composition as a determinant of reproductive success of Tengmalm's owls under fluctuating food conditions

The effect of landscape composition on the breeding success of vole-eating Tengmalm's owl ( Aegolius funereus ) was studied in western Finland at five different spatial scales (250–4000 m) around the nests during two consecutive three-year population cycles of voles. Landscape composition had strongest effects on owl breeding in the decrease phase of vole cycles. Significant variation in owl breeding occurred along the productivity gradient from farmland predominated areas to barren hinterland. Owls tended to produce earlier clutches on territories predominated by agricultural areas in increasing vole years. A similar trend was observed in the decreasing phase of the vole cycle; owls breeding on barren hinterland seemed to delay breeding compared to owls breeding near agricultural areas. Surprisingly, nestling survival and fledgling production in the decreasing phase declined steeply with increasing proportion of farmland. Clutch size was not significantly related to landscape composition. The number of fledglings decreased with increases in clear-cut and sapling areas in the decrease phase. During the declining years of vole abundance nestling survival increased from western farmland areas towards the eastern outlying district. These results indicate sudden summer decline of vole populations on farmland predominated habitats. This is probably due to that the number of vole-eating predators, and hence their impact on vole populations is apparently higher in farmland areas than on forested hinterland. This finding gives support for the 'spill-over' hypothesis, which states that predators and their exploitation tends to 'spill over' from luxuriant habitats to the barren habitats.