Summary1. In northern Scandinavia there are indications of a long-term decline in the abundance of the three dominant vole species, Clethrionomys glareolus , Clethrionomys rufocanus and Microtus agrestis , since the 1970s. One explanation proposes that intensified clear-cutting has created even-aged and homogeneous forest stands with poor overall conditions for survival and reproduction of the voles. 2. We investigated the relationship between forest age and structural habitat factors and its implications for the species richness and abundance of small mammals. In particular, we assessed the population dynamics of C. glareolus , a forest-dwelling species with rather general habitat requirements . 3. Extensive snap-trapping of small mammals was conducted during 1998-2000 on 24 study sites in boreal forests in northern Sweden. Trapping was carried out along transects running from immature forests of six age classes (0 -50 years) into adjacent reference sites (> 100 years). At each trapping station we recorded 14 habitat variables that were reduced to three principal components (PCs). The PCs were related to late successional traits, such as forest age and cover of tree layers (PC1), cover of tall vegetation in the field layer (PC2) and structural heterogeneity in the forest floor (PC3). 4. The species richness of small mammals, as well as the total abundance of C. glareolus , was positively influenced by tall vegetation (PC2) and structural heterogeneity (PC3) but not by late successional traits (PC1). The youngest forests had higher scores for both PC2 and PC3 compared with older forests. 5. The youngest forests also had the highest species richness and total abundance of C. glareolus . This was associated with a generally higher rate of change in numbers of C. glareolus during summer in the youngest forests compared with adjacent reference sites. In contrast, survival during winter was lower in the youngest forests. We found this result to be consistent with a source-sink scenario where young individuals, primarily born in old forest stands in early summer, migrate into younger forests to breed, but where the probabilities for winter survival are poor. 6. Our study demonstrates that both the species richness of small mammals and the population dynamics of C. glareolus are influenced to a great extent by structural habitat factors that are altered by common forest management practices in northern Sweden. In order to conserve species richness of small mammals and to minimize population fluctuations of C. glareolus in northern Scandinavia, we outline forest management practices that will provide heterogeneous environments, such as leaving logging residues on site after forest harvesting.

Cyclic populations of bank voles (Clethrionomys glareolus), grey-sided voles (Clethrionomys rufocanus), and field voles (Microtus agrestis) made up > 90% of the diet of Tengmalm's owl (Aegolius funereus) in northern Sweden during the breeding seasons of 1980–1986. These voles also made up > 90% of snap-trapped small mammals in the study area. Comparisons of the species composition in the diet during laying with that of animals trapped indicated that bank voles were usually taken approximately in proportion to their relative abundance, whereas field voles were frequently taken more and grey-sided voles less than expected from their relative abundance. However, bank voles predominated in the diet during laying early in the season (snow depth > 40 cm). Later in the season (snow depth ≤ 40 cm) the porportion of field voles and grey-sided voles increased during laying, suggesting increased availability of these species as the snow melted. Breeding density (percentage of nest boxes with ≥ 1 egg) showed a positive correlation whereas laying date showed a negative correlation with food supply in autumn. In contrast, clutch size, number of fledglings per successful nest, and an index of the annual production of fledglings showed positive correlations with food supply in spring. Clutch size declined with season but at different levels in different years, and laying date per se only explained 11% of clutch size variation among years. In contrast, the "year effect" (related to food supply) explained 29% of the variation in clutch size. Clutches were even larger at later laying dates (when, however, the food supply was better) in 1984 than in 1985. Nest survival and survival per egg (until fledging) in successful nests varied over the years, but did not covary with the cyclic food supply. However, the lowest nest survival was found in a year when voles declined dramatically over winter. Egg size did not show any variation among years.

In northern Sweden breeding males of Tengmalm's owls (Aegolius funereus (L.)) were site tenacious during and between the peaks of the vole (staple food) cycles, but females only during the peaks. Most of these adults shifted nest boxes between successive years. They selected nest boxes randomly in a radius of 3 km. Juveniles, in contrast to site tenacious adults, dispersed outside their natal area. The females moved longer than the males prior to their first breeding. Five adult females were found to be nomadic. One of these nomadic females previously bred site tenaciously as long as food was abundant. Juveniles and adult males were not found to be nomadic. Emigration of adult females and juveniles occurred most frequently when vole populations declined. The breeding population increased sharply and received immigrants suggesting that nomadism may be essential in the population dynamics. Site tenacity and nomadism are discussed in terms of costbenefit to males and females, respectively. Emphasis is on the main functional roles of males (feeding femle and young) and females (incubation).

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

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