Phenological diversity in the interactions between winter moth (Operophtera brumata) larvae and parasitoid wasps in sub-arctic mountain birch forest.

Population cycles of the winter moth (Operophtera brumata) in sub-arctic coastal birch forests show high spatiotemporal variation in amplitude. Peak larval densities range from levels causing little foliage damage to outbreaks causing spatially extensive defoliation. Moreover, outbreaks typically occur at or near the altitudinal treeline. It has been hypothesized that spatiotemporal variation in O. brumata cycle amplitude results from climate-induced variation in the degree of phenological matching between trophic levels, possibly between moth larvae and parasitoids. The likelihood of mismatching phenologies between larvae and parasitoids is expected to depend on how specialized parasitoids are, both as individual species and as a guild, to attacking specific larval developmental stages (i.e. instars). To investigate the larval instar-specificity of parasitoids, we studied the timing of parasitoid attacks relative to larval phenology. We employed an observational study design, with sequential sampling over the larval period, along an altitudinal gradient harbouring a pronounced treeline outbreak of O. brumata. Within the larval parasitoid guild, containing seven species groups, the timing of attack by different groups followed a successional sequence throughout the moth's larval period and each group attacked 1-2 instars. Such phenological diversity within parasitoid guilds may lower the likelihood of climate-induced trophic mismatches between victim populations and many/all of their enemies. Parasitism rates declined with increasing altitude for most parasitoid groups and for the parasitoid guild as a whole. However, the observed spatiotemporal parasitism patterns provided no clear evidence for or against altitudinal mismatch between larval and parasitoid phenology.

Intestinal parasites of the Arctic fox in relation to the abundance and distribution of intermediate hosts.

The intestinal parasite community of Arctic foxes (Vulpes lagopus) on the Svalbard archipelago in the High Arctic was investigated in relation to the abundance and distribution of intermediate hosts. Five species of cestodes (Echinococcus multilocularis, Taenia crassiceps, Taenia polyacantha, Taenia krabbei and Diphyllobothrium sp.), ascaridoid nematodes and one unidentified acanthocephalan species were found. The cestodes E. multilocularis, T. crassiceps and T. polyacantha all showed a decreasing prevalence in the fox population with increasing distance from their spatially restricted intermediate host population of sibling voles (Microtus levis). In addition, the prevalence of E. multilocularis in a sample from the vole population was directly related to the local vole abundance. The cestode T. krabbei uses reindeer as intermediate host, and its prevalence in female foxes was positively related to the density of reindeer (Rangifer tarandus platyrhyncus). Finally, the prevalence of the ascaridoid nematodes also decreased with increasing distance from the vole population, a finding that is consistent with the idea that voles are involved in transmission, most likely as paratenic hosts. The prevalence of the remaining species (Diphyllobothrium sp. and an unidentified acanthocephalan) was very low. We conclude that the distribution and abundance of intermediate host structure the gastrointestinal parasite community of the Arctic fox on the Svalbard archipelago.

Weak compensation of harvest despite strong density-dependent growth in willow ptarmigan.

Ptarmigan and grouse species (Lagopus spp.) are thought to be able to compensate for a modest harvest because there is a surplus of breeding birds that are prevented from breeding by territory holders. To estimate the degree of harvest-mortality compensation reliably we experimentally harvested 0%, 15% and 30% of the willow ptarmigan (Lagopus lagopus) on 13 estates ranging from 20 to 54 km2 in size during four hunting seasons in Norway according to a regional block design. Population overwinter growth rate was strongly negatively density dependent, but despite this, and contrary to earlier findings, only 33% of the harvest was compensated for. The lack of compensation was probably caused by long-distance juvenile dispersal that was unaffected by the harvest. The need for large-scale management experiments to detect the effects of harvest was clearly demonstrated: lack of compensation was found only when we used the whole dataset and not when the data were analysed by year or block. Our study shows that it is very difficult to demonstrate a population's lack of harvest compensation and warns against using small-scale, out-of-season or poorly replicated studies as a basis for future harvest-management decisions.

Echinococcus multilocularis on Svalbard: introduction of an intermediate host has enabled the local life-cycle.

The taeniid tapeworm Echinococcus multilocularis is here reported for the first time at the Svalbard Archipelago in the Norwegian Arctic. This new finding is interesting because the establishment of E. multilocularis is due to a recent anthropogenic introduction of its intermediate host--the sibling vole Microtus rossiaemeridionalis at Svalbard. The parasite itself has probably become naturally transferred to Svalbard due to migratory movements of its final host--the arctic fox Alopex lagopus between source areas for E. multilocularis in Siberia and Svalbard. We report macroscopically determined prevalence of E. multilocularis from a sample of 224 voles trapped in August in 1999 and 2000. The prevalence was among the highest ever recorded in intermediate hosts and was dependent on age and sex of the hosts approaching 100% in overwintered males. The high prevalence and the simplicity of the vole-arctic fox-E. multilocularis system at Svalbard makes it an eminent model system for further epidemiological studies.

Spatial synchronization of vole population dynamics by predatory birds.

Northern vole populations exhibit large-scale, spatially synchronous population dynamics. Such cases of population synchrony provide excellent opportunities for distinguishing between local intrinsic and regional extrinsic mechanisms of population regulation. Analyses of large-scale survey data and theoretical modelling have indicated several plausible synchronizing mechanisms. It is difficult, however, to determine the most important one without detailed data on local demographic processes. Here we combine results from two field studies in southeastern Norway--one identifies local demographic mechanisms and landscape-level annual synchrony among 28 enclosed experimental populations and the other examines region-level multi-annual synchrony in open natural populations. Despite fences eliminating predatory mammals and vole dispersal, the growth rates of the experimental populations were synchronized and moreover, perfectly linked with vole abundance in the region. The fates of 481 radio-marked voles showed that bird predation was the synchronizing mechanism. A higher frequency of risky dispersal movements in slowly growing populations appeared to accelerate predation rate. Thus, dispersal may induce a feedback-loop between predation and population growth that enhances synchrony.

Genetic differentiation among populations of the beetle Bolitophagus reticulatus (Coleoptera: tenebrionidae) in a fragmented and a continuous landscape.

The effect of habitat fragmentation on genetic differentiation among local populations of the fungivorous beetle Bolitophagus reticulatus (Coleoptera: Tenebrionidae) was studied in two contrasting landscapes: one heavily fragmented with forest fragments of variable size surrounded by inhabitable agricultural fields, the other an old forest providing a continuous habitat. The genetic structure of the beetle within each of the two contrasting areas was investigated by means of protein electrophoresis, screening four polymorphic loci in 20 populations from each area. In both areas there were significant genetic differences among local populations, but on average differentiation in the fragmented area was three times greater than in the continuous one, strongly indicating a genetic isolation effect of habitat fragmentation. These genetic results are in accordance with previous studies on dispersal in this species.

The effects of spatial habitat configuration on recruitment, growth and population structure in arctic Collembola.

The population density and demography of five species of arctic Collembola were studied in a naturally patchy habitat, consisting of Carex ursinae tussocks with varying degrees of isolation. Focal predictor variables were those describing the spatial configuration of tussocks, including tussock size and isolation and the amount of habitat (cover) at a 1-m2 scale surrounding each tussock population. The Collembola populations were heavily influenced by environmental stochasticity in the form of winter mortality and summer drought, and the influence of patchiness on population characteristics was evaluated in this context. The five species showed very different responses to the structuring effect of the habitat, depending on life history characteristics, mobility and habitat requirements. Population density was highly variable in both time and space. Spring densities indicated larger winter mortality compared to observations from a previous study, and the snow- and ice-free season from June to August only resulted in population growth for Folsomia sexoculata. In the other species, adult mortality must have been high as there was no net population growth despite observed reproduction. The exception was Hypogastrura viatica, whose population decline was more likely to have been the result of migration out of the study area. Cover was the most important variable explaining density. No pure area or isolation effects at the tussock level were detected, even in areas with very low habitat cover. Drought was probably an important mortality factor, as July was particularly warm and dry. Due to qualitative differences in the tussocks and the matrix substrate, desiccation risk would be higher during dispersal between tussocks. We suggest that increased dispersal mortality gave the observed pattern of increased density in relation to cover, both in general and in F. quadrioculata, an opportunistic species otherwise known for rapid population growth. Onychiurus groenlandicus, which had a similar density response to cover, may also be influenced by a rescue effect sustaining densities in areas with high cover. The cover effect can be viewed as a large-scale factor which encompasses the general spatial neighbourhood of each tussock, where inter-population processes are important, as opposed to internal patch dynamics.

Bank voles in linear habitats show restricted gene flow as revealed by mitochondrial DNA (mtDNA).

Genetic structure of bank vole populations in linear river bank habitat in southeast Norway was determined from analyses of DNA sequences for the mitochondrial D-loop. Animals were sampled at sites separated by 1 km, along two forested river banks separated by approximately 100 m of open water. Twenty-six distinct haplotypes were found among 120 voles. The voles showed significant deviation from panmixis on both sides of the river. Animals from the same site or from sites 1 km apart were more likely to share haplotypes than animals 2 km apart or more. Common haplotypes were widespread on both river banks, and had a wider distribution than relatively rare haplotypes. Some rare haplotypes were found on both banks, but most were restricted to a single bank. The results suggest that short-term gene flow may be restricted for female bank voles in linear habitats. Female territorial behaviour may vary with habitat geometry. In the linear habitat described here, females defend only two territorial borders and may effectively limit female dispersal. Results were compared to a previous study of bank voles from this region in a two-dimensional habitat. Gene flow in the linear habitat was much more restricted than gene flow in the two-dimensional habitat. Probable mechanisms underlying this difference are discussed.

The effects of experimental habitat destruction and patch isolation on space use and fitness parameters in female root vole Microtus oeconomus.

The aim of this study was to test whether experimentally induced destruction and fragmentation of habitat and varying degrees of patch isolation applied to 11 experimental populations of root voles affected space use and fitness parameters in radio-tracked females. It was predicted that the habitat destruction would disrupt the spatiosocial relations between breeding individuals, which, in turn, would cause lowered survival and reproductive success. Different degrees of patch isolation were expected to affect the rate of risky and energetically costly interpatch movements. The experimental habitat destruction comprised the removal of more than half of the habitat area (meadow vegetation) in each of seven populations that had been established in a large continous habitat block. This treatment yielded six small, habitat patches arranged in two clusters with long and short interfragment distances, respectively. Measures of individual space use, survival rates (predation and unknown causes) and litter production were contrasted with equivalent measures obtained from four control populations. The control populations had been established in permanently fragmented plots with the same fragment configuration as the post-destruction treatment populations. The effect of different interpatch distance was tested by comparing movement rates, space use and demographic parameters between the two types of patch cluster. Home ranges were larger in the continuous predestruction populations compared to the permanently fragmented control populations. There was a large decrease in home range area of individual females as a result of habitat destruction, and they became temporally smaller than in the control plots. The degree of overlap only increased on home range core areas. The degree of space sharing between matrilineal related females, a main characteristic of Microtus social organization, was higher in the permanently fragmented populations (controls) than in the treatment populations. This difference was also maintained after habitat destruction. Thus, the basic spatio-social organization of reproductive females when first established, seems to be very resistant to habitat destruction. There were no effects of habitat destruction on female fitness parameters when tested for either at the cluster or the population level. There were less between-patch movements when interpatch distances were large than when they were small, irrespective of the fragmentation history of the population. Predation by avian predators, the main mortality cause, was highest in fragment clusters with long interpatch distances, probably because long distance movements between fragments increased the predation risk. There was a large intrapopulation variation with respect to how severely individuals were affected by the habitat destruction. The farther a female had to relocate her home range because of habitat destruction, the higher was the predation risk. Thus, this study predicts that the most influential aspect of habitat fragmentation on demography may be that causing long interpatch movements.

Lack of concordance between mtDNA gene flow and population density fluctuations in the bank vole.

The genetic structure of bank voles Clethrionomys glareolus was determined from analyses of mitochondrial DNA (mtDNA) sequences, and compared with previous data on geographical synchrony in population density fluctuations. From 31 sample sites evenly spaced out along a 256-km transect in SE Norway a total of 39 distinct mtDNA haplotypes were found. The geographical distribution of the haplotypes was significantly non-random, and a cladistic analysis of the evolutionary relationship among haplotypes shows that descendant types were typically limited to a single site, whereas the ancestral types were more widely distributed geographically. This geographical distribution pattern of mtDNA haplotypes strongly indicates that the range and amount of female dispersal is severely restricted and insufficient to account for the previously observed synchrony in population density fluctuations. We conclude that geographical synchrony in this species must be caused by factors that are external to the local population, such as e.g. mobile predators.

The ecology and evolution of reproductive synchrony.

The temporal pattern of breeding in populations is often characterized by a pronounced temporal clustering of births, flowering or seed set. It has long been suspected that this phenomenon is not caused by climatic seasonality alone but that reproductive synchrony represents a strategy that individuals adopt to maximize reproductive success. The classical hypotheses predicting an adaptive advantage of reproductive synchrony incorporate both sociobiological and ecological explanations. However, new theoretical and empirical analyses have shown that the predicted advantage of reproductive synchrony depends on the ecological setting in which populations reproduce, and processes earlier thought to be responsible only for synchrony may under some ecological conditions lead to asynchronous reproduction being the best strategy.

Spatial clumping of sexually receptive females induces space sharing among male voles.

The spatial organization of individuals in populations (their spacing system) can be highly variable even among populations of the same species. As spacing systems have important consequences for ecological processes such as population regulation, competition and mating systems, there have been many attempts to explore factors that may cause this variation. For mammals, it has been argued that the spatial distribution of sexually receptive females is the most important factor determining the spacing system of males, whereas habitat characteristics are most important to females. This has been difficult to test experimentally as it requires manipulations of the spatial distribution of the opposite sex without changing other properties of the environment. Here, I present a novel experimental procedure that can achieve this and demonstrate that the spatial distribution of the opposite sex in a population of voles is indeed an important determinant of the spacing system of males, but not of females. However, the effects on males are different from those predicted by many theoretical studies.