The three-dimensional arrangement of vegetation has an essential influence on habitat quality and... more The three-dimensional arrangement of vegetation has an essential influence on habitat quality and, therefore, the presence and abundance of animal species at local scales (Tews et al., 2004). The complexity and diversity of vegetation determine the diversity and behavior of higher organisms by influencing the availability and diversity of resources and niches (Hekkala, Tarvainen,
Due to their reclusive nature, information on the population structure of many bat species is lac... more Due to their reclusive nature, information on the population structure of many bat species is lacking or scarce. The pattern of small scale population genetic structure could reveal the degree of gene flow among colonies, and the evolutionary consequences of short-distance dispersal. In this study, we used nine microsatellite loci to assess the small-scale genetic population structure of Daubenton's bats in the Archipelago Sea comparing it to samples from sites elsewhere in Finland and Europe. The Archipelago Sea is a highly variable environment with possible dispersal barriers. Our results indicate a low level of population genetic structuring among the populations sampled. We found significant isolation by distance in both sexes, indicating a gradual increase of population differentiation across a large geographic scale. In Finland alone, isolationby-distance was also found, with high levels of gene flow among local populations. Isolation-by-distance was stronger in females, suggesting that males disperse longer distances.
In bacteria associated with humans, antimicrobial resistance is common, both in clinical isolates... more In bacteria associated with humans, antimicrobial resistance is common, both in clinical isolates and in the less-studied commensal flora, and it is thought that commensal and environmental bacteria might be a hidden reservoir of resistance. Gilliver et al. have reported that resistance is also prevalent in faecal bacteria from wild rodents living in northwest England. Here we test the faeces of moose, deer and vole in Finland and find an almost complete absence of resistance in enterobacteria. Resistance is thus not a universal property of enterobacterial populations, but may be a result of the human use of antibiotics.
In small mammal populations with multiannual oscillations in density, the occurrence of large ind... more In small mammal populations with multiannual oscillations in density, the occurrence of large individuals in the peak phase (the "Chitty effect") is a typical feature, but mechanisms behind this phenomenon have remained unclear. We analysed long-term data sets collected in western Finland between 1984 and 1992 to: (1) find out how the body size and body condition of voles (Microtus agrestis, M. rossiaemeridionalis, Clethrionomys glareolus) and shrews (Sorex araneus) was associated with the 3-year population cycle of voles, and (2) relate the quality (body condition) of the individuals to changes in the biotic environment in order to detect how the different hypotheses about the mechanisms behind the Chitty effect can explain the observed variation. In the 3-year cycle studied, the mean body size and quality were strongly related to density oscillations in voles but not in sympatric shrews. Voles were lean in the decline phase but very stocky in the summer of the peak phase. This pattern appeared to be mainly caused by changes in body condition or body shape rather than mere size (body length). The quality of voles appeared to be delayed density dependent, especially in autumn when the dominant time lag was 12 months. Previous vole density was strongly related to changes in the environment (activity of specialist predators, production of hay until early summer). We suggest that the previous density of voles mainly affects the quality of voles indirectly through changes in the biotic environment, and that the proximate cause behind the Chitty effect is the combined effect of changes in predation pressure and availability of food.
We studied responses of stoats and least weasels to fluctuating vole abundances during seven wint... more We studied responses of stoats and least weasels to fluctuating vole abundances during seven winters in western Finland. Density indices ofmustelids 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 • 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.
Proceedings of the Royal Society of London. Series B: Biological Sciences, 2002
Mechanisms generating the well-known 3-5 year cyclic fluctuations in densities of northern small ... more 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.
Proceedings of the Royal Society B: Biological Sciences, 2005
Comprehensive analyses of long–term (1977–2003) small–mammal abundance data from western Finland ... more Comprehensive analyses of long–term (1977–2003) small–mammal abundance data from western Finland showed that populations ofMicrotusvoles (field volesM.agrestisand sibling volesM.rossiaemeridionalis), bank voles (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 amongMicrotusand 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 km2) through a 3 year small–mammal population cycle. Predator reduction increased densities of the main prey species,Microtusvoles, in all phas...
We studied the indirect effects of vertebrate predator exclusion on plant communities in boreal g... more We studied the indirect effects of vertebrate predator exclusion on plant communities in boreal grassland in western Finland to find out whether the removal of the top trophic level would result in a trophic cascade. Predators were excluded from 1996 to 2000 by eight predator-proof fences (each 0.5 ha) constructed on old fields. Despite a major increase in vole densities, the expected trophic cascade attenuated rapidly so that the indirect effects of predator exclusion were restricted to a few plant species. The cause for the rapid attenuation of the trophic cascade appeared to be strong seasonality, as peak densities of voles were attained at the end of the growing season of vegetation, and vole populations declined before the next growing season so that the herbivory pressure during the growing season remained low or moderate. Accordingly, most plants escaped the heaviest grazing pressure either in time (plants completed their reproduction and withered before winter) or in space (living parts hidden under frozen ground and ice). However, heavy winter herbivory reduced the biomass of available vegetation and killed woody species (willows) at vole peaks, which implies that predator exclusion may have a strong effect on secondary succession. During summer, voles reduced the coverage of only a few preferred food plants (Elymus repens, Phleum pratense, Vicia cracca). Voles also maintained annual and biennial species in the community by creating gaps in the closed vegetation. We conclude that abiotic factors (harsh winter conditions) limited peak numbers of herbivores below a threshold density where herbivores could have caused a community-level decline in the biomass of herbaceous plants during summer.
Three mechanisms have been proposed to induce spatial synchrony in fluctuations of small mammal p... more Three mechanisms have been proposed to induce spatial synchrony in fluctuations of small mammal populations: climate-related environmental effects, predation and dispersal. We conducted a field experiment in western Finland to evaluate the relative roles of these mechanisms in inducing spatial synchrony among cyclic populations of field voles Microtus agrestis. The study was conducted during the increase and peak phases of a vole population cycle on four agricultural field sites situated 1.5 Á/7.0 km apart. Each field contained two 0.5-ha fenced enclosures and one 1-ha unfenced control area. One enclosure per field allowed access by small mustelid predators and the other by avian predators; all enclosures prevented the dispersal of voles. The unfenced control areas allowed access by all predators as well as dispersal by voles. Enclosed vole populations were in a treatment-wise asynchronous phase before the predator access treatments were applied. The growth rates of all enclosed populations were tightly synchronized during the course of the experiment. Conversely, synchrony both among the unfenced populations and between the fenced and unfenced populations was practically non-existent. During winter, in the increase phase of the cycle, vole populations in all treatments declined to low densities due to a seasonal effect of winter food depletion. During summer, in the peak year of the vole cycle, all populations fluctuated in synchrony. At this time, both small mustelids and birds of prey appeared to be abundant enough to induce synchrony. Dispersal was identified as a potential contributor to synchronization, but the magnitude of its effects could not be reliably discerned. Our results indicate that no single mechanism can account for the observed patterns of spatial synchrony among cyclic northern vole populations. Rather, spatial synchronization is induced by different mechanisms, namely seasonality and predation, acting successively during different seasons and phases of the vole cycle.
Norrdahl, K. and Korpimäki, E. 2002. Changes in population structure and reproduction during a 3-... more Norrdahl, K. and Korpimäki, E. 2002. Changes in population structure and reproduction during a 3-yr population cycle of voles.-Oikos 96: 331-345. Cyclic changes in population growth rate are caused by changes in survival and/or reproductive rate. To find out whether cyclic changes in reproduction are an important part of the mechanism causing cyclic fluctuations in small mammal populations, we studied changes in the population structure and reproduction of field voles (Microtus agrestis), sibling voles (M. rossiaemeridionalis), bank voles (Clethrionomys glareolus), and common shrews (Sorex araneus) in western Finland during 1984-1992, in an area with 3-yr vole cycles. We also modelled the population growth of voles using parameter values from this study. The animals studied were collected by snap trapping in April, May, June, August, September, and, during 1986-1990, also in October. We found several phase-related differences in the population structure (age structure, sex ratio, proportion of mature individuals) and reproduction (litter size, length of the breeding season) of voles. In non-cyclic common shrews, the only significant phase-related difference was a lower proportion of overwintered individuals in the increase phase. According to the analyses and the vole model, phase-related changes in litter size had only a minor impact on population growth rate. The same was true for winter breeding in the increase phase. The length and intensity of the summer breeding season had an effect on yearly population growth but this impact was relatively weak compared to the effect of cyclic changes in survival. The population increase rates of Microtus were delayed dependent on density (8-12-month time lag). Our results indicate that cyclic changes in reproduction are not an important part of the mechanism driving cyclic fluctuations in vole populations. Low survival of young individuals appeared to play an important role in the shift from the peak to the decline phase in late summer and early autumn.
We studied whether the presence of breeding kestrels (Falco tinnunculus) affected nest predation ... more We studied whether the presence of breeding kestrels (Falco tinnunculus) affected nest predation and breeding habitat selection of curlews (Numenius arquata) on an open flat farmland area in western Finland. We searched for nests of curlews from an area of 6 km(2) during 1985-1993. For each nest found, we recorded the fate of the nest, and the distance to the nearest kestrel nest and to the nearest perch. We measured the impact of breeding kestrels on nest predation by constructing artificial curlew nests in the vicinity of ten kestrel nests in 1993. Curlew nests were closer to kestrel nests than expected from random distribution, eventhough kestrels fed on average 5.5% of curlew chick production. Predation risk by kestrels was lower than predation risk by corvids and other generalist predators, which predated 9% of curlew nests surviving farming practices and an unknown proportion of chicks. Artificial nest experiment showed that nest predation was lower close to kestrel nests than further away suggesting that the breeding association of curlews and kestrels was a behavioural adaptation against nest predation. Thus, the presence of a predator may sometimes be beneficial to prey, and prey animals have behavioural adaptations to these situations.
Reproductive output and the growth of captive voles were quanti®ed under high and low avian preda... more Reproductive output and the growth of captive voles were quanti®ed under high and low avian predation risk in a semi-natural experiment. Voles were exposed to Eurasian kestrels (Falco tinnunculus), the main avian predator of vole species studied (Clethrionomys glareolus, Microtus agrestis and M. rossiaemeridionalis). Vole pairs were housed in cages settled under nest-boxes occupied by breeding kestrels or in control cages settled under empty nest-boxes for 2 weeks. The experiment was conducted in midsummer when kestrels had half-grown nestlings, because in that time hunting adults and begging nestlings produce noise and scats which may indicate signi®cant predation threat to voles housed underneath the nest-boxes. The risk of kestrel predation did not have any obvious impact on pregnancy rates, mean litter sizes, or growth rates of kestrel-exposed voles compared with control voles studied. These results indicate that the risk of avian predation does not depress the reproductive investment of voles. Key words Avian predation risk á Indirect eects of predation á Reproductive investment á Breeding suppression á Voles Oecologia (1998) 115:149±153
The abundances of potential avian prey species may be lower in the vicinity of the nests of preda... more The abundances of potential avian prey species may be lower in the vicinity of the nests of predatory birds than farther away. We predicted that, if this density depression is due to predation, the observed density pattern should develop gradually during the breeding ...
1. Graham & Lambin (2002) have reported on a weasel-reduction experiment, concluding that the imp... more 1. Graham & Lambin (2002) have reported on a weasel-reduction experiment, concluding that the impact of weasel predation on field vole survival was neither sufficient nor necessary to initiate and drive the cyclic decline of field vole populations in Kielder Forest, northern England. They also stated that their findings contradict conclusively the specialist predator hypothesis put forward to explain population cycles of voles in North Europe. 2. Straightforward inferences from Kielder Forest to the northern boreal zone are misleading, because the population cycles of voles in Kielder Forest differ essentially from North European vole cycles. The low amplitude of the vole cycles in Kielder Forest, their restricted spatial synchrony in comparison to northern Europe and the virtual lack of interspecific synchrony in Kielder Forest suggest that there are essential differences between the mechanisms responsible for the two types of cyclic fluctuations of voles. 3. The weasel-reduction experiment may provide a misleading picture on the role of predators, even in the Kielder Forest cycle. The experimental reduction of weasels alone may not stop the population decline of voles, because competing larger predators are expected to increase their hunting in the weasel-reduction areas. The small spatial scale of the experiment, which produced only slight, short-term differences in weasel densities between reduction and control areas, also suggests that other predators could have compensated easily for the weasels that were removed. 4. We propose a new version of the predation hypothesis to explain low-amplitude population cycles of voles in temperate Europe, including the Kielder Forest. The interaction between generalist predators and vole populations might account for these cycles because generalists can have a functional response that is destabilizing in the neighbourhood of the equilibrium point. As most generalists are orders of magnitude larger than weasels, and thus need much more food for survival, generalist-driven cycles should be characterized by high prey minima, as observed in Kielder Forest.
The three-dimensional arrangement of vegetation has an essential influence on habitat quality and... more The three-dimensional arrangement of vegetation has an essential influence on habitat quality and, therefore, the presence and abundance of animal species at local scales (Tews et al., 2004). The complexity and diversity of vegetation determine the diversity and behavior of higher organisms by influencing the availability and diversity of resources and niches (Hekkala, Tarvainen,
Due to their reclusive nature, information on the population structure of many bat species is lac... more Due to their reclusive nature, information on the population structure of many bat species is lacking or scarce. The pattern of small scale population genetic structure could reveal the degree of gene flow among colonies, and the evolutionary consequences of short-distance dispersal. In this study, we used nine microsatellite loci to assess the small-scale genetic population structure of Daubenton's bats in the Archipelago Sea comparing it to samples from sites elsewhere in Finland and Europe. The Archipelago Sea is a highly variable environment with possible dispersal barriers. Our results indicate a low level of population genetic structuring among the populations sampled. We found significant isolation by distance in both sexes, indicating a gradual increase of population differentiation across a large geographic scale. In Finland alone, isolationby-distance was also found, with high levels of gene flow among local populations. Isolation-by-distance was stronger in females, suggesting that males disperse longer distances.
In bacteria associated with humans, antimicrobial resistance is common, both in clinical isolates... more In bacteria associated with humans, antimicrobial resistance is common, both in clinical isolates and in the less-studied commensal flora, and it is thought that commensal and environmental bacteria might be a hidden reservoir of resistance. Gilliver et al. have reported that resistance is also prevalent in faecal bacteria from wild rodents living in northwest England. Here we test the faeces of moose, deer and vole in Finland and find an almost complete absence of resistance in enterobacteria. Resistance is thus not a universal property of enterobacterial populations, but may be a result of the human use of antibiotics.
In small mammal populations with multiannual oscillations in density, the occurrence of large ind... more In small mammal populations with multiannual oscillations in density, the occurrence of large individuals in the peak phase (the "Chitty effect") is a typical feature, but mechanisms behind this phenomenon have remained unclear. We analysed long-term data sets collected in western Finland between 1984 and 1992 to: (1) find out how the body size and body condition of voles (Microtus agrestis, M. rossiaemeridionalis, Clethrionomys glareolus) and shrews (Sorex araneus) was associated with the 3-year population cycle of voles, and (2) relate the quality (body condition) of the individuals to changes in the biotic environment in order to detect how the different hypotheses about the mechanisms behind the Chitty effect can explain the observed variation. In the 3-year cycle studied, the mean body size and quality were strongly related to density oscillations in voles but not in sympatric shrews. Voles were lean in the decline phase but very stocky in the summer of the peak phase. This pattern appeared to be mainly caused by changes in body condition or body shape rather than mere size (body length). The quality of voles appeared to be delayed density dependent, especially in autumn when the dominant time lag was 12 months. Previous vole density was strongly related to changes in the environment (activity of specialist predators, production of hay until early summer). We suggest that the previous density of voles mainly affects the quality of voles indirectly through changes in the biotic environment, and that the proximate cause behind the Chitty effect is the combined effect of changes in predation pressure and availability of food.
We studied responses of stoats and least weasels to fluctuating vole abundances during seven wint... more We studied responses of stoats and least weasels to fluctuating vole abundances during seven winters in western Finland. Density indices ofmustelids 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 • 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.
Proceedings of the Royal Society of London. Series B: Biological Sciences, 2002
Mechanisms generating the well-known 3-5 year cyclic fluctuations in densities of northern small ... more 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.
Proceedings of the Royal Society B: Biological Sciences, 2005
Comprehensive analyses of long–term (1977–2003) small–mammal abundance data from western Finland ... more Comprehensive analyses of long–term (1977–2003) small–mammal abundance data from western Finland showed that populations ofMicrotusvoles (field volesM.agrestisand sibling volesM.rossiaemeridionalis), bank voles (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 amongMicrotusand 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 km2) through a 3 year small–mammal population cycle. Predator reduction increased densities of the main prey species,Microtusvoles, in all phas...
We studied the indirect effects of vertebrate predator exclusion on plant communities in boreal g... more We studied the indirect effects of vertebrate predator exclusion on plant communities in boreal grassland in western Finland to find out whether the removal of the top trophic level would result in a trophic cascade. Predators were excluded from 1996 to 2000 by eight predator-proof fences (each 0.5 ha) constructed on old fields. Despite a major increase in vole densities, the expected trophic cascade attenuated rapidly so that the indirect effects of predator exclusion were restricted to a few plant species. The cause for the rapid attenuation of the trophic cascade appeared to be strong seasonality, as peak densities of voles were attained at the end of the growing season of vegetation, and vole populations declined before the next growing season so that the herbivory pressure during the growing season remained low or moderate. Accordingly, most plants escaped the heaviest grazing pressure either in time (plants completed their reproduction and withered before winter) or in space (living parts hidden under frozen ground and ice). However, heavy winter herbivory reduced the biomass of available vegetation and killed woody species (willows) at vole peaks, which implies that predator exclusion may have a strong effect on secondary succession. During summer, voles reduced the coverage of only a few preferred food plants (Elymus repens, Phleum pratense, Vicia cracca). Voles also maintained annual and biennial species in the community by creating gaps in the closed vegetation. We conclude that abiotic factors (harsh winter conditions) limited peak numbers of herbivores below a threshold density where herbivores could have caused a community-level decline in the biomass of herbaceous plants during summer.
Three mechanisms have been proposed to induce spatial synchrony in fluctuations of small mammal p... more Three mechanisms have been proposed to induce spatial synchrony in fluctuations of small mammal populations: climate-related environmental effects, predation and dispersal. We conducted a field experiment in western Finland to evaluate the relative roles of these mechanisms in inducing spatial synchrony among cyclic populations of field voles Microtus agrestis. The study was conducted during the increase and peak phases of a vole population cycle on four agricultural field sites situated 1.5 Á/7.0 km apart. Each field contained two 0.5-ha fenced enclosures and one 1-ha unfenced control area. One enclosure per field allowed access by small mustelid predators and the other by avian predators; all enclosures prevented the dispersal of voles. The unfenced control areas allowed access by all predators as well as dispersal by voles. Enclosed vole populations were in a treatment-wise asynchronous phase before the predator access treatments were applied. The growth rates of all enclosed populations were tightly synchronized during the course of the experiment. Conversely, synchrony both among the unfenced populations and between the fenced and unfenced populations was practically non-existent. During winter, in the increase phase of the cycle, vole populations in all treatments declined to low densities due to a seasonal effect of winter food depletion. During summer, in the peak year of the vole cycle, all populations fluctuated in synchrony. At this time, both small mustelids and birds of prey appeared to be abundant enough to induce synchrony. Dispersal was identified as a potential contributor to synchronization, but the magnitude of its effects could not be reliably discerned. Our results indicate that no single mechanism can account for the observed patterns of spatial synchrony among cyclic northern vole populations. Rather, spatial synchronization is induced by different mechanisms, namely seasonality and predation, acting successively during different seasons and phases of the vole cycle.
Norrdahl, K. and Korpimäki, E. 2002. Changes in population structure and reproduction during a 3-... more Norrdahl, K. and Korpimäki, E. 2002. Changes in population structure and reproduction during a 3-yr population cycle of voles.-Oikos 96: 331-345. Cyclic changes in population growth rate are caused by changes in survival and/or reproductive rate. To find out whether cyclic changes in reproduction are an important part of the mechanism causing cyclic fluctuations in small mammal populations, we studied changes in the population structure and reproduction of field voles (Microtus agrestis), sibling voles (M. rossiaemeridionalis), bank voles (Clethrionomys glareolus), and common shrews (Sorex araneus) in western Finland during 1984-1992, in an area with 3-yr vole cycles. We also modelled the population growth of voles using parameter values from this study. The animals studied were collected by snap trapping in April, May, June, August, September, and, during 1986-1990, also in October. We found several phase-related differences in the population structure (age structure, sex ratio, proportion of mature individuals) and reproduction (litter size, length of the breeding season) of voles. In non-cyclic common shrews, the only significant phase-related difference was a lower proportion of overwintered individuals in the increase phase. According to the analyses and the vole model, phase-related changes in litter size had only a minor impact on population growth rate. The same was true for winter breeding in the increase phase. The length and intensity of the summer breeding season had an effect on yearly population growth but this impact was relatively weak compared to the effect of cyclic changes in survival. The population increase rates of Microtus were delayed dependent on density (8-12-month time lag). Our results indicate that cyclic changes in reproduction are not an important part of the mechanism driving cyclic fluctuations in vole populations. Low survival of young individuals appeared to play an important role in the shift from the peak to the decline phase in late summer and early autumn.
We studied whether the presence of breeding kestrels (Falco tinnunculus) affected nest predation ... more We studied whether the presence of breeding kestrels (Falco tinnunculus) affected nest predation and breeding habitat selection of curlews (Numenius arquata) on an open flat farmland area in western Finland. We searched for nests of curlews from an area of 6 km(2) during 1985-1993. For each nest found, we recorded the fate of the nest, and the distance to the nearest kestrel nest and to the nearest perch. We measured the impact of breeding kestrels on nest predation by constructing artificial curlew nests in the vicinity of ten kestrel nests in 1993. Curlew nests were closer to kestrel nests than expected from random distribution, eventhough kestrels fed on average 5.5% of curlew chick production. Predation risk by kestrels was lower than predation risk by corvids and other generalist predators, which predated 9% of curlew nests surviving farming practices and an unknown proportion of chicks. Artificial nest experiment showed that nest predation was lower close to kestrel nests than further away suggesting that the breeding association of curlews and kestrels was a behavioural adaptation against nest predation. Thus, the presence of a predator may sometimes be beneficial to prey, and prey animals have behavioural adaptations to these situations.
Reproductive output and the growth of captive voles were quanti®ed under high and low avian preda... more Reproductive output and the growth of captive voles were quanti®ed under high and low avian predation risk in a semi-natural experiment. Voles were exposed to Eurasian kestrels (Falco tinnunculus), the main avian predator of vole species studied (Clethrionomys glareolus, Microtus agrestis and M. rossiaemeridionalis). Vole pairs were housed in cages settled under nest-boxes occupied by breeding kestrels or in control cages settled under empty nest-boxes for 2 weeks. The experiment was conducted in midsummer when kestrels had half-grown nestlings, because in that time hunting adults and begging nestlings produce noise and scats which may indicate signi®cant predation threat to voles housed underneath the nest-boxes. The risk of kestrel predation did not have any obvious impact on pregnancy rates, mean litter sizes, or growth rates of kestrel-exposed voles compared with control voles studied. These results indicate that the risk of avian predation does not depress the reproductive investment of voles. Key words Avian predation risk á Indirect eects of predation á Reproductive investment á Breeding suppression á Voles Oecologia (1998) 115:149±153
The abundances of potential avian prey species may be lower in the vicinity of the nests of preda... more The abundances of potential avian prey species may be lower in the vicinity of the nests of predatory birds than farther away. We predicted that, if this density depression is due to predation, the observed density pattern should develop gradually during the breeding ...
1. Graham & Lambin (2002) have reported on a weasel-reduction experiment, concluding that the imp... more 1. Graham & Lambin (2002) have reported on a weasel-reduction experiment, concluding that the impact of weasel predation on field vole survival was neither sufficient nor necessary to initiate and drive the cyclic decline of field vole populations in Kielder Forest, northern England. They also stated that their findings contradict conclusively the specialist predator hypothesis put forward to explain population cycles of voles in North Europe. 2. Straightforward inferences from Kielder Forest to the northern boreal zone are misleading, because the population cycles of voles in Kielder Forest differ essentially from North European vole cycles. The low amplitude of the vole cycles in Kielder Forest, their restricted spatial synchrony in comparison to northern Europe and the virtual lack of interspecific synchrony in Kielder Forest suggest that there are essential differences between the mechanisms responsible for the two types of cyclic fluctuations of voles. 3. The weasel-reduction experiment may provide a misleading picture on the role of predators, even in the Kielder Forest cycle. The experimental reduction of weasels alone may not stop the population decline of voles, because competing larger predators are expected to increase their hunting in the weasel-reduction areas. The small spatial scale of the experiment, which produced only slight, short-term differences in weasel densities between reduction and control areas, also suggests that other predators could have compensated easily for the weasels that were removed. 4. We propose a new version of the predation hypothesis to explain low-amplitude population cycles of voles in temperate Europe, including the Kielder Forest. The interaction between generalist predators and vole populations might account for these cycles because generalists can have a functional response that is destabilizing in the neighbourhood of the equilibrium point. As most generalists are orders of magnitude larger than weasels, and thus need much more food for survival, generalist-driven cycles should be characterized by high prey minima, as observed in Kielder Forest.
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