Changes in individual quality during a 3-year population cycle of voles.

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.