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Population ecologists have traditionally focused on the patterns and causes of population variation in the temporal domain for which a substantial body of practical analytic techniques have been developed. More recently, numerous studies have documented how populations may fluctuate synchronously over large spatial areas; analyses of such spatially extended time-series have started to provide additional clues regarding the causes of these population fluctuations and explanations for their synchronous occurrence. Here, we report on the development of a phase-based method for identifying coupling between temporally coincident but spatially distributed cyclic time-series, which we apply to the numbers of muskrat and mink recorded at 81 locations across Canada. The analysis reveals remarkable parallel clines in the strength of coupling between proximate populations of both species--declining from west to east--together with a corresponding increase in observed synchrony between these populations the further east they are located.

A new geographical gradient in the dynamics of small rodents is demonstrated by analysing 29 time series of density indices of the common vole (Microtus arvalis) from Poland, the Czech Republic and the Slovak Republic. This gradient extends from more stable northerly populations in coastal Poland to more variable and cyclic populations in the southernmost parts of the Slovak Republic, and is hence a reversal of the Fennoscandian gradient. All studied variables (such as mean density, cycle amplitude, density variability and the coefficients in a second-order autoregressive model) exhibit consistent latitudinal variation. Possible underlying factors are discussed. In particular, we suggest that seasonality may be a key element in explaining the observed new gradient.

Life-history traits relating to growth and reproduction vary greatly among species and populations and among individuals within populations. In vole populations, body size and age at maturation may vary considerably among locations and among years within the same location. Individuals in increasing populations are typically larger and start reproduction earlier in the spring than those in declining populations. The cause of such life-history variation within populations has been subject of much discussion. Much of the controversy concerns whether the memory of past conditions, leading to delayed effects on life-history traits, resides in the environment (for example, predators, pathogens or food) or intrinsically within populations or individuals (age distribution, physiological state, genetic or maternal effects). Here we report from an extensive field transplant experiment in which voles were moved before the breeding season between sites that differed in average overwintering body mass. Transplanted voles did not retain the characteristics of their source population, and we demonstrate an over-riding role of the immediate environment in shaping life-history traits of small rodents.

In a continuous habitat, restricted dispersal and local genetic drift are likely to create a pattern of increasing genetic differentiation with distance. Here, we describe the genetic structure of Siberian lemming (Lemmus sibiricus) populations in a continuous tundra habitat on the western coast of the Taimyr Peninsula, in order to determine the spatial scale at which genetic differentiation and isolation by distance occur. Sampling was carried out at three different geographical scales: (1) a continuous 11 km transect; (2) localities 10-30 km apart; and (3) two localities at 300 and 600 km from the main study area. Two types of genetic markers were used: partial sequences of the mitochondrial DNA control region and four microsatellite loci. On this basis the study populations were genetically quite homogeneous within patches extending over 8 km or more. Contrary to theoretical predictions, no pattern of isolation by distance among patches could be identified. This observation was interpreted as representing populations in migration-drift disequilibrium after a recent major mixing event. The lack of concordance between mtDNA haplotype phylogeny and the geographical distribution of haplotypes supported this interpretation. Spatial autocorrelation among individual genotypes on a local scale was weak and observed only in females, indicating a considerable amount of mostly male-mediated gene flow. Average gene flow per generation was estimated to be in the range of several hundred metres.

In order to evaluate the biogeographical hypothesis that the Norwegian lemming (Lemmus lemmus) survived the last glacial period in some Scandinavian refugia, we examined variation in the nucleotide sequence of the mitochondrial control region (402 base pairs (bp)) and the cytochrome b (cyt b) region (633 bp) in Norwegian and Siberian (Lemmus sibiricus) lemmings. The phylogenetic distinction and cyt b divergence estimate of 1.8% between the Norwegian and Siberian lemmings suggest that their separation pre-dated the last glaciation and imply that the Norwegian lemming is probably a relic of the Pleistocene populations from Western Europe. The star-like control region phylogeny and low mitochondrial DNA diversity in the Norwegian lemming indicate a reduction in its historical effective size followed by population expansion. The average estimate of post-bottleneck time (19-21 kyr) is close to the last glacial maximum (18-22 kyr BP). Taking these findings and the fossil records into consideration, it seems likely that, after colonization of Scandinavia in the Late Pleistocene, the Norwegian lemming suffered a reduction in its population effective size and survived the last glacial maximum in some local Scandinavian refugia, as suggested by early biogeographical work.

The pattern and scale of the genetic structure of populations provides valuable information for the understanding of the spatial ecology of populations, including the spatial aspects of density fluctuations. In the present paper, the genetic structure of periodically fluctuating lemmings (Dicrostonyx groenlandicus) in the Canadian Arctic was analysed using mitochondrial DNA (mtDNA) control region sequences and four nuclear microsatellite loci. Low genetic variability was found in mtDNA, while microsatellite loci were highly variable in all localities, including localities on isolated small islands. For both genetic markers the genetic differentiation was clear among geographical regions but weaker among localities within regions. Such a pattern implies gene flow within regions. Based on theoretical calculations and population census data from a snap-trapping survey, we argue that the observed genetic variability on small islands and the low level of differentiation among these islands cannot be explained without invoking long distance dispersal of lemmings over the sea ice. Such dispersal is unlikely to occur only during population density peaks.

Variation in the nucleotide sequence of the mitochondrial control region (250 bp) and the cytochrome b region (870 bp) was examined in collared lemmings (Dicrostonyx groenlandicus) from 19 localities in northern Alaska and the Canadian Arctic. The division of D. groenlandicus in two phylogeographical groups with limited divergence across the Mackenzie River is consistent with the separation of this species in more than one refugial area located to the northwest of the Laurentide ice sheet during the last glaciation. Populations of D.groenlandicus from formerly glaciated areas are no less variable than those in nonglaciated areas. Instead, the low intrapopulation and intraregional diversity estimates in D. groenlandicus are probably a result of regional bottleneck events due to range contractions during Holocene warming events. These results are consistent with findings previously reported on collared lemmings (D. torquatus) from the Eurasian Arctic.

A 3- to 5-year cycle of vole abundances is a characteristic phenomenon in the ecology of northern regions, and their explanation stands as a central theoretical challenge in population ecology. Although many species of voles usually coexist and are in severe competition for food and breeding space, the role of interspecific competition in vole cycles has never been evaluated statistically. After studying community effects on the population dynamics of the gray-sided vole (Clethrionomys rufocanus) in the subarctic birch forest at Kilpisjärvi, Finland, we report statistical results showing that both interspecific and intraspecific effects are important in the direct year-to-year density dependence. However, interspecific effects are not detectable in the 2-year delayed density dependence that is crucial for generating the characteristic cycles. Furthermore, we show that most of the competition takes place during the winter. The results are evaluated against two models of community dynamics. One assumes that the delayed effects are caused by an interaction with a specialist predator, and the other assumes that they are caused by overgrazing food plants. These statistical results show that vole cycles may be generated by a species-specific trophic interaction. The results also suggest that the gray-sided vole may be the focal species in the birch-forest community, as field voles may be in the taiga and as lemmings may be on the tundra.

We analysed two 26-year long (1970-1995) time-series on annual population growth rates of Norwegian lemmings (Lemmus lemmus) from Finse, south Norway, using a threshold autoregressive (TAR) approach. We demonstrate that the population dynamics is both phase- and density-dependent. The phase-dependence accounts for the observed nonlinearity. We used the deduced stochastic model structure as a basis for evaluating the dynamic properties of this system. The dynamics is characterized either by limit cycles or chaos (the latter with a strong semi-periodic component). Stochasticity is seen to play an important role in the determination of the periodicity. The ecological implications of these statistical and mathematical results are discussed.

Current ecological information on periodically fluctuating microtine populations are demonstrated to support a hypothesis involving both predation and intrinsic self-regulation as necessary and sufficient factors for explaining the "microtine density cycle'. The structure of the cyclic time series is largely two dimensional with strong delayed density dependence. Together with recent field studies on rodent demography, our modelling suggests that trophic interaction is a likely candidate to generate the dimensionality observed for northern microtine rodent dynamics. It is shown that the trophic interaction must be fairly strong. This suggests that specialist predation is the most likely one among the classes of trophic interactions. We also argue that some - but not too strong - self-regulation must occur to generate the structure of the available time series on northern European microtines.

A total of 31 years of abundance data from 90 populations of the grey-sided vole (Clethrionomys rufocanus) in northern Hokkaido (Japan) were analysed with respect to population dynamic characteristics. Both non-periodic and multi-annually periodic fluctuations occur among the studied populations. The length of the period varies from 2 to 5 years. The dynamics appear approximately linear on a logarithmic scale, and a log-linear stochastic difference model with one time-lag is found to recreate the periodograms of the time series. The deterministic (log-linear) component of these models determines the periodicity while stochasticity sustains the cycles. The dynamics of the Hokkaidian vole populations are found to vary clinally from the western coast eastwards and towards the interior of the study area. This gradient corresponds superficially to the latitudinal gradient seen in microtine populations in Fennoscandia. However, under close scrutiny, these gradients differ greatly: the Hokkaidian gradient is caused by a cline in delayed density dependence. Statistical delayed density dependence is more negative towards the east and the interior.