Seasonality, density dependence, and spatial population synchrony.

Studies of spatial population synchrony constitute a central approach for understanding the drivers of ecological dynamics. Recently, identifying the ecological impacts of climate change has emerged as a new important focus in population synchrony studies. However, while it is well known that climatic seasonality and sequential density dependence influences local population dynamics, the role of season-specific density dependence in shaping large-scale population synchrony has not received attention. Here, we present a widely applicable analytical protocol that allows us to account for both season and geographic context-specific density dependence to better elucidate the relative roles of deterministic and stochastic sources of population synchrony, including the renowned Moran effect. We exemplify our protocol by analyzing time series of seasonal (spring and fall) abundance estimates of cyclic rodent populations, revealing that season-specific density dependence is a major component of population synchrony. By accounting for deterministic sources of synchrony (in particular season-specific density dependence), we are able to identify stochastic components. These stochastic components include mild winter weather events, which are expected to increase in frequency under climate warming in boreal and Arctic ecosystems. Interestingly, these weather effects act both directly and delayed on the vole populations, thus enhancing the Moran effect. Our study demonstrates how different drivers of population synchrony, presently altered by climate warming, can be disentangled based on seasonally sampled population time-series data and adequate population models.

Climate variability and density-dependent population dynamics: Lessons from a simple High Arctic ecosystem.

Ecologists are still puzzled by the diverse population dynamics of herbivorous small mammals that range from high-amplitude, multiannual cycles to stable dynamics. Theory predicts that this diversity results from combinations of climatic seasonality, weather stochasticity, and density-dependent food web interactions. The almost ubiquitous 3- to 5-y cycles in boreal and arctic climates may theoretically result from bottom-up (plant-herbivore) and top-down (predator-prey) interactions. Assessing, empirically, the roles of such interactions and how they are influenced by environmental stochasticity has been hampered by food web complexity. Here, we take advantage of a uniquely simple High Arctic food web, which allowed us to analyze the dynamics of a graminivorous vole population not subjected to top-down regulation. This population exhibited high-amplitude, noncyclic fluctuations-partly driven by weather stochasticity. However, the predominant driver of the dynamics was overcompensatory density dependence in winter that caused the population to frequently crash. Model simulations showed that the seasonal pattern of density dependence would yield regular 2-y cycles in the absence of stochasticity. While such short cycles have not yet been observed in mammals, they are theoretically plausible if graminivorous vole populations are deterministically bottom-up regulated. When incorporating weather stochasticity in the model simulations, cyclicity became disrupted and the amplitude was increased-akin to the observed dynamics. Our findings contrast with the 3- to 5-y population cycles that are typical of graminivorous small mammals in more complex food webs, suggesting that top-down regulation is normally an important component of such dynamics.

Predator co-occurrence in alpine and Arctic tundra in relation to fluctuating prey.

Large carnivores influence ecosystem dynamics in multiple ways, for example, by suppressing meso-carnivores and providing carrions for smaller scavengers. Loss of large carnivores is suggested to cause meso-carnivore increase and expansion. Moreover, competition between meso-carnivores may be modified by the presence of larger carnivores. In tundra ecosystems, the smallest meso-carnivore, the Arctic fox, has experienced regional declines, whereas its larger and competitively superior congener, the red fox, has increased, potentially due to changes in the abundance of apex predators. We explored if variation in the occurrence of wolverine and golden eagle impacted the occurrence and co-occurrence of the Arctic fox and red fox in relation to varying abundances of small rodents within the Scandinavian tundra. We applied multi-species occupancy models to an extensive wildlife camera dataset from 2011-2020 covering 98 sites. Daily detection/non-detection of each species per camera trap site and study period (late winter; March-May) was stacked across years, and species occupancy was related to small rodent abundance while accounting for time of the year and status of simulated carcass. The Arctic fox was more likely to co-occur with the red fox when the wolverine was present and less likely to co-occur with the red fox when golden eagles were present and the wolverine was absent. Red foxes increased in occupancy when co-occurring with the larger predators. The Arctic fox responded more strongly to small rodent abundance than the red fox and co-occurred more often with the other species at carcasses when rodent abundance was low. Our findings suggest that the interspecific interactions within this tundra predator guild appear to be surprisingly intricate, driven by facets of fear of predation, interspecific mediation and facilitation, and food resource dynamics. These dynamics of intraguild interactions may dictate where and when conservation actions targeted towards the Arctic fox should be implemented.

Iterative model predictions for wildlife populations impacted by rapid climate change.

To improve understanding and management of the consequences of current rapid environmental change, ecologists advocate using long-term monitoring data series to generate iterative near-term predictions of ecosystem responses. This approach allows scientific evidence to increase rapidly and management strategies to be tailored simultaneously. Iterative near-term forecasting may therefore be particularly useful for adaptive monitoring of ecosystems subjected to rapid climate change. Here, we show how to implement near-term forecasting in the case of a harvested population of rock ptarmigan in high-arctic Svalbard, a region subjected to the largest and most rapid climate change on Earth. We fitted state-space models to ptarmigan counts from point transect distance sampling during 2005-2019 and developed two types of predictions: (1) explanatory predictions to quantify the effect of potential drivers of ptarmigan population dynamics, and (2) anticipatory predictions to assess the ability of candidate models of increasing complexity to forecast next-year population density. Based on the explanatory predictions, we found that a recent increasing trend in the Svalbard rock ptarmigan population can be attributed to major changes in winter climate. Currently, a strong positive effect of increasing average winter temperature on ptarmigan population growth outweighs the negative impacts of other manifestations of climate change such as rain-on-snow events. Moreover, the ptarmigan population may compensate for current harvest levels. Based on the anticipatory predictions, the near-term forecasting ability of the models improved nonlinearly with the length of the time series, but yielded good forecasts even based on a short time series. The inclusion of ecological predictors improved forecasts of sharp changes in next-year population density, demonstrating the value of ecosystem-based monitoring. Overall, our study illustrates the power of integrating near-term forecasting in monitoring systems to aid understanding and management of wildlife populations exposed to rapid climate change. We provide recommendations for how to improve this approach.

End-user involvement to improve predictions and management of populations with complex dynamics and multiple drivers.

Sustainable management of wildlife populations can be aided by building models that both identify current drivers of natural dynamics and provide near-term predictions of future states. We employed a Strategic Foresight Protocol (SFP) involving stakeholders to decide the purpose and structure of a dynamic state-space model for the population dynamics of the Willow Ptarmigan, a popular game species in Norway. Based on local knowledge of stakeholders, it was decided that the model should include food web interactions and climatic drivers to provide explanatory predictions. Modeling confirmed observations from stakeholders that climate change impacts Ptarmigan populations negatively through intensified outbreaks of insect defoliators and later onset of winter. Stakeholders also decided that the model should provide anticipatory predictions. The ability to forecast population density ahead of the harvest season was valued by the stakeholders as it provides the management extra time to consider appropriate harvest regulations and communicate with hunters prior to the hunting season. Overall, exploring potential drivers and predicting short-term future states, facilitate collaborative learning and refined data collection, monitoring designs, and management priorities. Our experience from adapting a SFP to a management target with inherently complex dynamics and drivers of environmental change, is that an open, flexible, and iterative process, rather than a rigid step-wise protocol, facilitates rapid learning, trust, and legitimacy.

Dietary variation in Icelandic arctic fox (Vulpes lagopus) over a period of 30 years assessed through stable isotopes.

Identifying resources driving long-term trends in predators is important to understand ecosystem changes and to manage populations in the context of conservation or control. The arctic fox population in Iceland has increased steadily over a period of 30 years, an increase that has been attributed to an overall increase in food abundance. We hypothesized that increasing populations of geese or seabirds were driving this growth. We analyzed stable isotopes in a long-term series of collagen samples to determine the role of these different resources. The isotopic signatures of arctic foxes differed consistently between coastal and inland habitats. While δ15N displayed a non-linear change over time with a slight increase in the first part of the period followed by a decline in both habitats, δ13C was stable. Stable isotope mixing models suggested that marine resources and rock ptarmigan were the most important dietary sources, with marine resources dominating in coastal habitats and rock ptarmigan being more important inland. Our results suggest that seabirds may have been driving the arctic fox population increase. The rapidly increasing populations of breeding geese seem to have played a minor role in arctic fox population growth, as rock ptarmigan was the most important terrestrial resource despite a considerable decrease in their abundance during recent decades. This study shows that a long-term population trend in a generalist predator may have occurred without a pronounced change in main dietary resources, despite ongoing structural changes in the food web, where one species of herbivorous birds increased and another decreased.

Contrasting changes in space use induced by climate change in two Arctic marine mammal species.

Global warming is inducing major environmental changes in the Arctic. These changes will differentially affect species owing to differences in climate sensitivity and behavioural plasticity. Arctic endemic marine mammals are expected to be impacted significantly by ongoing changes in their key habitats owing to their long life cycles and dependence on ice. Herein, unique biotelemetry datasets for ringed seals (RS; Pusa hispida) and white whales (WW; Delphinapterus leucas) from Svalbard, Norway, spanning two decades (1995-2016) are used to investigate how these species have responded to reduced sea-ice cover and increased Atlantic water influxes. Tidal glacier fronts were traditionally important foraging areas for both species. Following a period with dramatic environmental change, RS now spend significantly more time near tidal glaciers, where Arctic prey presumably still concentrate. Conversely, WW spend significantly less time near tidal glacier fronts and display spatial patterns that suggest that they are foraging on Atlantic fishes that are new to the region. Differences in levels of dietary specialization and overall behavioural plasticity are likely reasons for similar environmental pressures affecting these species differently. Climate change adjustments through behavioural plasticity will be vital for species survival in the Arctic, given the rapidity of change and limited dispersal options.

Rangifer management controls a climate-sensitive tundra state transition.

Rangifer (caribou/reindeer) management has been suggested to mitigate the temperature-driven transition of Arctic tundra into a shrubland state, yet how this happens is uncertain. Here we study this much focused ecosystem state transition in riparian areas, where palatable willows (Salix) are dominant tall shrubs and highly responsive to climate change. For the state transition to take place, small life stages must become tall and abundant. Therefore we predicted that the performance of small life stages (potential recruits) of the tall shrubs were instrumental to the focal transition, where Rangifer managed at high population density would keep the small-stage shrubs in a "browse trap" independent of summer temperature. We used a large-scale quasi-experimental study design that included real management units that spanned a wide range of Rangifer population densities and summer temperatures in order to assess the relative importance of these two driving variables. Ground surveys provided data on density and height of the small shrub life stages, while the distributional limit (shrubline) of established shrublands (the tall shrub life stage) was derived from aerial photographs. Where Rangifer densities were above a threshold of approximately 5 animals/km2 , we found, in accordance with the expectation of a "browse trap," that the small life stages of shrubs in grasslands were at low height and low abundance. At Rangifer densities below this threshold, the small life stages of shrubs were taller and more abundant indicating Rangifer were no longer in control of the grassland state. For the established shrubland state, we found that the shrubline was at a 100-m lower elevation in the management units where Rangifer had been browsing in summer as opposed to the migratory ranges with no browsing in summer. In both seasonal ranges, the shrubline increased 100 m per 1°C increment in temperature. Our study supports the proposal that Rangifer management within a sustainable range of animal densities can mitigate the much-focused transition from grassland to shrubland in a warming Arctic.

An Arctic predator-prey system in flux: climate change impacts on coastal space use by polar bears and ringed seals.

Climate change is impacting different species at different rates, leading to alterations in biological interactions with ramifications for wider ecosystem functioning. Understanding these alterations can help improve predictive capacity and inform management efforts designed to mitigate against negative impacts. We investigated how the movement and space use patterns of polar bears (Ursus maritimus) in coastal areas in Svalbard, Norway, have been altered by a sudden decline in sea ice that occurred in 2006. We also investigated whether the spatial overlap between polar bears and their traditionally most important prey, ringed seals (Pusa hispida), has been affected by the sea-ice decline, as polar bears are dependent on a sea-ice platform for hunting seals. We attached biotelemetry devices to ringed seals (n = 60, both sexes) and polar bears (n = 67, all females) before (2002-2004) and after (2010-2013) a sudden decline in sea ice in Svalbard. We used linear mixed-effects models to evaluate the association of these species to environmental features and an approach based on Time Spent in Area to investigate changes in spatial overlap between the two species. Following the sea-ice reduction, polar bears spent the same amount of time close to tidal glacier fronts in the spring but less time in these areas during the summer and autumn. However, ringed seals did not alter their association with glacier fronts during summer, leading to a major decrease in spatial overlap values between these species in Svalbard's coastal areas. Polar bears now move greater distances daily and spend more time close to ground-nesting bird colonies, where bear predation can have substantial local effects. Our results indicate that sea-ice declines have impacted the degree of spatial overlap and hence the strength of the predator-prey relationship between polar bears and ringed seals, with consequences for the wider Arctic marine and terrestrial ecosystems. Shifts in ecological interactions are likely to become more widespread in many ecosystems as both predators and prey respond to changing environmental conditions induced by global warming, highlighting the importance of multi-species studies.

From individuals to population cycles: the role of extrinsic and intrinsic factors in rodent populations.

Rodent population cycles have fascinated scientists for a long time. Among various hypotheses, an interaction of an extrinsic factor (predation) with intrinsic factors (e.g., sociality and dispersal) was suggested to lead to the generation of population cycles. Here, we tested this hypothesis with an individual-based model fully parameterized with an exceptionally rich empirical database on vole life histories. We employed a full factorial design that included models with the following factors: predation only, predation and sociality, predation and dispersal, and predation and both sociality and dispersal. A comprehensive set of metrics was used to compare results of these four models with the long-term population dynamics of natural vole populations. Only the full model, which included both intrinsic factors and predation, yielded cycle periods, amplitudes, and autumn population sizes closest to those observed in nature. Our approach allows to model, as emergent properties of individual life histories, the sort of nonlinear density- and phase-dependence that is expected to destabilize population dynamics. We suggest that the individual-based approach is useful for addressing the effects of other mechanisms on rodent populations that operate at finer temporal and spatial scales than have been explored with models so far.

Why Do the Boreal Forest Ecosystems of Northwestern Europe Differ from Those of Western North America?

The boreal forest is one of the largest terrestrial biomes on Earth. Conifers normally dominate the tree layer across the biome, but other aspects of ecosystem structure and dynamics vary geographically. The cause of the conspicuous differences in the understory vegetation and the herbivore-predator cycles between northwestern Europe and western North America presents an enigma. Ericaceous dwarf shrubs and 3- to 4-year vole-mustelid cycles characterize the European boreal forests, whereas tall deciduous shrubs and 10-year snowshoe hare-lynx cycles characterize the North American ones. We discuss plausible explanations for this difference and conclude that it is bottom-up: Winter climate is the key determinant of the dominant understory vegetation that then determines the herbivore-predator food-web interactions. The crucial unknown for the twenty-first century is how climate change and increasing instability will affect these forests, both with respect to the dynamics of individual plant and animal species and to their community interactions.

Continental-scale travelling waves in forest geometrids in Europe: an evaluation of the evidence.

A recent paper claims the existence of one of the most large-scale travelling waves ever recorded for any animal population. Here we address why conceptual and methodological pitfalls may have served to exaggerate or even impose the spatial patterns reported. Photo credit: Jane U. Jepsen.

Predictions replaced by facts: a keystone species' behavioural responses to declining arctic sea-ice.

Since the first documentation of climate-warming induced declines in arctic sea-ice, predictions have been made regarding the expected negative consequences for endemic marine mammals. But, several decades later, little hard evidence exists regarding the responses of these animals to the ongoing environmental changes. Herein, we report the first empirical evidence of a dramatic shift in movement patterns and foraging behaviour of the arctic endemic ringed seal (Pusa hispida), before and after a major collapse in sea-ice in Svalbard, Norway. Among other changes to the ice-regime, this collapse shifted the summer position of the marginal ice zone from over the continental shelf, northward to the deep Arctic Ocean Basin. Following this change, which is thought to be a 'tipping point', subadult ringed seals swam greater distances, showed less area-restricted search behaviour, dived for longer periods, exhibited shorter surface intervals, rested less on sea-ice and did less diving directly beneath the ice during post-moulting foraging excursions. In combination, these behavioural changes suggest increased foraging effort and thus also likely increases in the energetic costs of finding food. Continued declines in sea-ice are likely to result in distributional changes, range reductions and population declines in this keystone arctic species.

Identification and Evaluation of 21 Novel Microsatellite Markers from the Autumnal Moth (Epirrita autumnata) (Lepidoptera: Geometridae).

The autumnal moth (Epirrita autumnata) is a cyclically outbreaking forest Lepidoptera with circumpolar distribution and substantial impact on Northern ecosystems. We have isolated 21 microsatellites from the species to facilitate population genetic studies of population cycles, outbreaks, and crashes. First, PCR primers and PCR conditions were developed to amplify 19 trinucleotide loci and two tetranucleotide loci in six multiplex PCR approaches and then analyzed for species specificity, sensitivity and precision. Twelve of the loci showed simple tandem repeat array structures while nine loci showed imperfect repeat structures, and repeat numbers varied in our material between six and 15. The application in population genetics for all the 21 microsatellites were further validated in 48 autumnal moths sampled from Northern Norway, and allelic variation was detected in 19 loci. The detected numbers of alleles per locus ranged from two to 13, and the observed and expected heterozygosities varied from 0.04 to 0.69 and 0.04 to 0.79, respectively. Evidence for linkage disequilibrium was found for six loci as well as indication of one null allele. We find that these novel microsatellites and their multiplex-PCR assays are suitable for further research on fine- and large-scale population-genetic studies of Epirrita autumnata.

Demographic responses of a site-faithful and territorial predator to its fluctuating prey: long-tailed skuas and arctic lemmings.

Environmental variability, through interannual variation in food availability or climatic variables, is usually detrimental to population growth. It can even select for constancy in key life-history traits, though some exceptions are known. Changes in the level of environmental variability are therefore important to predict population growth or life-history evolution. Recently, several cyclic vole and lemming populations have shown large dynamical changes that might affect the demography or life-histories of rodent predators. Skuas constitute an important case study among rodent predators, because of their strongly saturating breeding productivity (they lay only two eggs) and high degree of site fidelity, in which they differ from nomadic predators raising large broods in good rodent years. This suggests that they cannot capitalize on lemming peaks to the same extent as nomadic predators and might be more vulnerable to collapses of rodent cycles. We develop a model for the population dynamics of long-tailed skuas feeding on lemmings to assess the demographic consequences of such variable and non-stationary prey dynamics, based on data collected in NE Greenland. The model shows that populations of long-tailed skua sustain well changes in lemming dynamics, including temporary collapses (e.g. 10 years). A high floater-to-breeder ratio emerges from rigid territorial behaviour and a long-life expectancy, which buffers the impact of adult abundance's decrease on the population reproductive output. The size of the floater compartment is affected by changes in both mean and coefficient of variation of lemming densities (but not cycle amplitude and periodicity per se). In Greenland, the average lemming density is below the threshold density required for successful breeding (including during normally cyclic periods). Due to Jensen's inequality, skuas therefore benefit from lemming variability; a positive effect of environmental variation. Long-tailed skua populations are strongly adapted to fluctuating lemming populations, an instance of demographic lability in the reproduction rate. They are also little affected by poor lemming periods, if there are enough floaters, or juveniles disperse to neighbouring populations. The status of Greenland skua populations therefore strongly depends upon floater numbers and juvenile movements, which are not known. This reveals a need to intensify colour-ringing efforts on the long-tailed skua at a circumpolar scale.

Bears are simply voles writ large: social structure determines the mechanisms of intrinsic population regulation in mammals.

The literature reveals opposing views regarding the importance of intrinsic population regulation in mammals. Different models have been proposed; adding importance to contrasting life histories, body sizes and social interactions. Here we evaluate current theory based on results from two Scandinavian projects studying two ecologically different mammal species with contrasting body sizes and life history traits: the root vole Microtus oeconomus and the brown bear Ursus arctos. We emphasize four inter-linked behavioral aspects-territoriality, dispersal, social inhibition of breeding, and infanticide-that together form a density-dependent syndrome with potentially regulatory effects on population growth. We show that the two species are similar in all four behaviors and thus the overall regulatory syndrome. Females form matrilineal assemblages, female natal dispersal is negatively density dependent and breeding is suppressed in philopatric young females. In both species, male turnover due to extrinsic mortality agents cause infanticide with negative effects on population growth. The sex-biased and density-dependent dispersal patterns promote the formation of matrilineal clusters which, in turn, leads to reproductive suppression with potentially regulatory effects. Hence, we show that intrinsic population regulation interacting with extrinsic mortality agents may occur irrespective of taxon, life history and body size. Our review stresses the significance of a mechanistic approach to understanding population ecology. We also show that experimental model populations are useful to elucidate natural populations of other species with similar social systems. In particular, such experiments should be combined with methodical innovations that may unravel the effects of cryptic intrinsic mechanisms such as infanticide.

Determinants of lemming outbreaks.

Population outbreaks in tundra rodents have intrigued scientists for a century as a result of their spectacular appearances and their general lessons in ecology. One outstanding question that has led to competing hypotheses is why sympatric lemmings and voles differ in regularity and shape of their outbreaks. Lemming outbreaks may be lost for decades while vole populations maintain regular population cycles. Moreover, when lemming populations eventually irrupt, they do so more steeply than the vole populations. Norwegian lemmings exhibited a large-scale outbreak synchronously with gray-sided voles in Finnmark, northern Fennoscandia, during 2006 to 2007 for the first time in two decades. Analyses of spatial variability of this outbreak across altitudinal gradients allowed us to identify determinants of the contrasting lemming and vole dynamics. The steeper lemming outbreak trajectories were caused by breeding and population growth during winter, when nonbreeding vole populations consistently declined. The differently shaped lemming and vole outbreaks appear to result from a particular demographic tactic of lemmings that evolved as an adaptation to the long and cold Arctic-Alpine winters. The lemming outbreak amplitude increased with altitude and vole density, indicating that lemming outbreaks are jointly facilitated by low temperatures and apparent mutualism with voles mediated by shared predators. High sensitivity to variation in climate and predation is likely to be the reasons why lemmings have more erratic population dynamics than sympatric voles. The combination of continued climatic warming and dampened vole cycles is expected to further decrease the frequency, amplitude, and geographic range of lemming outbreaks in tundra ecosystems.

Is the diet cyclic phase-dependent in boreal vole populations?

Herbivorous rodents in boreal, alpine and arctic ecosystems are renowned for their multi-annual population cycles. Researchers have hypothesised that these cycles may result from herbivore-plant interactions in various ways. For instance, if the biomass of preferred food plants is reduced after a peak phase of a cycle, rodent diets can be expected to become dominated by less preferred food plants, leading the population to a crash. It could also be expected that the taxonomic diversity of rodent diets increases from the peak to the crash phase of a cycle. The present study is the first to use DNA metabarcoding to quantify the diets of two functionally important boreal rodent species (bank vole and tundra vole) to assess whether their diet changed systematically in the expected cyclic phase-dependent manner. We found the taxonomic diet spectrum broad in both vole species but with little interspecific overlap. There was no evidence of systematic shifts in diet diversity metrics between the phases of the population cycle in either species. While both species' diet composition changed moderately between cycle phases and seasons, these changes were small compared to other sources of diet variation-especially differences between individuals. Thus, the variation in diet that could be attributed to cyclic phases is marginal relative to the overall diet flexibility. Based on general consumer-resource theory, we suggest that the broad diets with little interspecific overlap render it unlikely that herbivore-plant interactions generate their synchronous population cycles. We propose that determining dietary niche width should be the first step in scientific inquiries about the role of herbivore-plant interactions in cyclic vole populations.

Indirect food web interactions mediated by predator-rodent dynamics: relative roles of lemmings and voles.

Production cycles in birds are proposed as prime cases of indirect interactions in food webs. They are thought to be driven by predators switching from rodents to bird nests in the crash phase of rodent population cycles. Although rodent cycles are geographically widespread and found in different rodent taxa, bird production cycles appear to be most profound in the high Arctic where lemmings dominate. We hypothesized that this may be due to arctic lemmings inducing stronger predator responses than boreal voles. We tested this hypothesis by estimating predation rates in dummy bird nests during a rodent cycle in low-Arctic tundra. Here, the rodent community consists of a spatially variable mix of one lemming (Lemmus lemmus) and two vole species (Myodes rufocanus and Microtus oeconomus) with similar abundances. In consistence with our hypothesis, lemming peak abundances predicted well crash-phase nest predation rates, whereas the vole abundances had no predictive ability. Corvids were found to be the most important nest predators. Lemmings appear to be accessible to the whole predator community which makes them particularly powerful drivers of food web dynamics.

The role of colonization in the dynamics of patchy populations of a cyclic vole species.

The crash phase of vole populations with cyclic dynamics regularly leads to vast areas of uninhabited habitats. Yet although the capacity for cyclic voles to re-colonize such empty space is likely to be large and predicted to have become evolved as a distinct life history trait, the processes of colonization and its effect on the spatio-temporal dynamics have been little studied. Here we report from an experiment with root voles (Microtus oeconomus) specifically targeted at quantifying the process of colonization of empty patches from distant source patches and its resultant effect on local vole deme size variation in a patchy landscape. Three experimental factors: habitat quality, predation risk and inter-patch distance were employed among 24 habitat patches in a 100 × 300-m experimental area. The first-born cohort in the spring efficiently colonized almost all empty patches irrespective of the degree of patch isolation and predation risk, but this was dependent on habitat quality. Just after the initial colonization wave the deme sizes in patches of the same quality were underdispersed relative to Poisson variance, indicating regulated (density-dependent) settlement. Towards the end of the breeding season local demographic processes acted to smooth out the initial post-colonization differences among source and colonization patches, and among patches of initially different quality. However, at this time demographic stochasticity had also given rise to a large (overdispersed) variation in deme sizes that may have contributed to an overshadowing of the effect of other factors. The results of this experiment confirmed our expectation that the space-filling capacity of voles is large. The costs associated with transience appeared to be so low, at least at the spatial scale considered in this experiment, that such costs are not likely to substantially constrain habitat selection and colonization in the increase phase of cyclic patchy populations.