Sporadic cases of chronic wasting disease in old moose - an epidemiological study.

Transmissible spongiform encephalopathies or prion diseases comprise diseases with different levels of contagiousness under natural conditions. The hypothesis has been raised that the chronic wasting disease (CWD) cases detected in Nordic moose (Alces alces) may be less contagious, or not contagious between live animals under field conditions. This study aims to investigate the epidemiology of CWD cases detected in moose in Norway, Sweden and Finland using surveillance data from 2016 to 2022.In total, 18 CWD cases were detected in Nordic moose. All moose were positive for prion (PrPres) detection in the brain, but negative in lymph nodes, all were old (mean 16 years; range 12-20) and all except one, were female. Age appeared to be a strong risk factor, and the sex difference may be explained by few males reaching high age due to hunting targeting calves, yearlings and males.The cases were geographically scattered, distributed over 15 municipalities. However, three cases were detected in each of two areas, Selbu in Norway and Arjeplog-Arvidsjaur in Sweden. A Monte Carlo simulation approach was applied to investigate the likelihood of such clustering occurring by chance, given the assumption of a non-contagious disease. The empirical P-value for obtaining three cases in one Norwegian municipality was less than 0.05, indicating clustering. However, the moose in Selbu were affected by different CWD strains, and over a 6 year period with intensive surveillance, the apparent prevalence decreased, which would not be expected for an ongoing outbreak of CWD. Likewise, the three cases in Arjeplog-Arvidsjaur could also indicate clustering, but management practices promotes a larger proportion of old females and the detection of the first CWD case contributed to increased awareness and sampling.The results of our study show that the CWD cases detected so far in Nordic moose have a different epidemiology compared to CWD cases reported from North America and in Norwegian reindeer (Rangifer tarandus tarandus). The results support the hypothesis that these cases are less contagious or not contagious between live animals under field conditions. To enable differentiation from other types of CWD, we support the use of sporadic CWD (sCWD) among the names already in use.

Factors promoting hunting groups' sustainable harvest of moose in a co-management system.

Collaboration between and within management levels and involvement of local communities (co-management) increases sustainable management of natural resources. In Finland, moose (Alces alces) are harvested by hunting groups within a co-management system, providing meat and social benefits. We computed the 14-year change in moose harvest (2007-2020) for 4320 hunting groups. Moose harvest declined on average 1.1% per year, but with substantial variation in moose harvest changes between the hunting groups. We extracted information describing the collaboration between the hunting groups, their democratic status as well as leader dynamics, and the year of establishment. A hunting group's moose harvest was more stable (i.e. declined less) when the hunting group was (1) established a longer time ago; (2), had more changes in leadership over time, but did not depend on collaboration with other local hunting groups (in terms of jointly holding moose hunting licenses), whether the hunting group was a registered society (presumed to be more democratic than a non-registered one) or had consecutive leaders that shared a surname (presumed to be related). We conclude that encouraging resource users' early establishment in groups and groups' long-term persistence and promoting democratic leadership roles improves stable benefits from a natural resource in a co-management system.

Density-habitat relationships of white-tailed deer (Odocoileus virginianus) in Finland.

In heterogeneous landscapes, resource selection constitutes a crucial link between landscape and population-level processes such as density. We conducted a non-invasive genetic study of white-tailed deer in southern Finland in 2016 and 2017 using fecal DNA samples to understand factors influencing white-tailed deer density and space use in late summer prior to the hunting season. We estimated deer density as a function of landcover types using a spatial capture-recapture (SCR) model with individual identities established using microsatellite markers. The study revealed second-order habitat selection with highest deer densities in fields and mixed forest, and third-order habitat selection (detection probability) for transitional woodlands (clear-cuts) and closeness to fields. Including landscape heterogeneity improved model fit and increased inferred total density compared with models assuming a homogenous landscape. Our findings underline the importance of including habitat covariates when estimating density and exemplifies that resource selection can be studied using non-invasive methods.

Calf/female ratio and population dynamics of wild forest reindeer in relation to wolf and moose abundances in a managed European ecosystem.

BACKGROUND: The alternative prey hypothesis describes the mechanism for apparent competition whereby the mortality of the secondary prey species increases (and population size decreases decreases) by the increased predation by the shared predator if the population size of the primary prey decreases. Apparent competition is a process where the abundance of two co-existing prey species are negatively associated because they share a mutual predator, which negatively affects the abundance of both prey Here, we examined whether alternative prey and/or apparent competition hypothesis can explain the population dynamics and reproductive output of the secondary prey, wild forest reindeer (Rangifer tarandus fennicus) in Finland, in a predator-prey community in which moose (Alces alces) is the primary prey and the wolf (Canis lupus) is the generalist predator. METHODS: We examined a 22-year time series (1996-2017) to determine how the population size and the calf/female ratio of wild forest reindeer in Eastern Finland were related to the abundances of wolf and moose. Only moose population size was regulated by hunting. Summer predation of wolves on reindeer focuses on calves. We used least squares regression (GLS) models (for handling autocorrelated error structures and resulting pseudo-R2s) and generalized linear mixed (GLMs) models (for avoidance of negative predictions) to determine the relationships between abundances. We performed linear and general linear models for the calf/female ratio of reindeer. RESULTS AND SYNTHESIS: The trends in reindeer population size and moose abundance were almost identical: an increase during the first years and then a decrease until the last years of our study period. Wolf population size in turn did not show long-term trends. Change in reindeer population size between consecutive winters was related positively to the calf/female ratio. The calf/female ratio was negatively related to wolf population size, but the reindeer population size was related to the wolf population only when moose abundance was entered as another independent variable. The wolf population was not related to moose abundance even though it is likely to consist the majority of the prey biomass. Because reindeer and moose populations were positively associated, our results seemed to support the alternative prey hypothesis more than the apparent competition hypothesis. However, these two hypotheses are not mutually exclusive and the primary mechanism is difficult to distinguish as the system is heavily managed by moose hunting. The recovery of wild forest reindeer in eastern Finland probably requires ecosystem management involving both habitat restoration and control of species abundances.

Estimating preharvest density, adult sex ratio, and fecundity of white-tailed deer using noninvasive sampling techniques.

Adult sex ratio and fecundity (juveniles per female) are key population parameters in sustainable wildlife management, but inferring these requires abundance estimates of at least three age/sex classes of the population (male and female adults and juveniles). Prior to harvest, we used an array of 36 wildlife camera traps during 2 and 3 weeks in the early autumn of 2016 and 2017, respectively. We recorded white-tailed deer adult males, adult females, and fawns from the pictures. Simultaneously, we collected fecal DNA (fDNA) from 92 20 m × 20 m plots placed in 23 clusters of four plots between the camera traps. We identified individuals from fDNA samples with microsatellite markers and estimated the total sex ratio and population density using spatial capture-recapture (SCR). The fDNA-SCR analysis concluded equal sex ratio in the first year and female bias in the second year, and no difference in space use between sexes (fawns and adults combined). Camera information was analyzed in a spatial capture (SC) framework assuming an informative prior for animals' space use, either (a) as estimated by fDNA-SCR (same for all age/sex classes), (b) as assumed from the literature (space use of adult males larger than adult females and fawns), or (c) by inferring adult male space use from individually identified males from the camera pictures. These various SC approaches produced plausible inferences on fecundity, but also inferred total density to be lower than the estimate provided by fDNA-SCR in one of the study years. SC approaches where adult male and female were allowed to differ in their space use suggested the population had a female-biased adult sex ratio. In conclusion, SC approaches allowed estimating the preharvest population parameters of interest and provided conservative density estimates.

Modelling the effect of moose Alces alces population density and regional forest structure on the amount of damage in forest seedling stands.

BACKGROUND: Moose (Alces alces L.) populations and moose damage in forests are debated in Nordic countries with dense moose populations. Moose populations and food resources vary greatly, both spatially and temporally, and reliable data covering both variables simultaneously at the same scale have seldom been available. We modelled the effect of moose population density and forest resources on the area of moose damage at regional scale, referring to moose management areas (MMA). Forest data and moose damage data originated from the Finnish National Forest Inventory, and the moose population data came from a Bayesian moose model. For modelling, average values of moose population, damage and forest variables were calculated for the periods 2004-2008 and 2009-2013 for each MMA. The MMAs were further classified into one of four larger geographical zones. The area of moose damage was used as a dependent variable, and the proportions of different types of forests and moose population densities per land area or area of seedling stands as explanatory variables. The relationships were modelled with a linear mixed-effects model with an exponential spatial correlation structure. RESULTS: The area of moose damage was best explained by total forest area, proportions of plantations and mature forests, and moose population density per land area or the proportion of plantations. There were differences among the biogeographical zones in how different variables explained the amount of damage. CONCLUSION: The results provide tools for analyzing the regional effects of moose population density and the amount of food resources on the amount of moose damage. This information can be used in reconciling sustainable moose population levels and the amount of damage.

Is transport distance correlated with animal welfare and carcass quality of reindeer (Rangifer tarandus tarandus)?

BACKGROUND: Slaughter reindeer are exposed to stress caused by gathering, handling, loading and unloading, and by conditions in vehicles during transport. These stress factors can lead to compromised welfare and trauma such as bruises or fractures, aspiration of rumen content, and abnormal odour in carcasses, and causing condemnations in meat inspection and lower meat quality. We investigated the statistical association of slaughter transport distance with these indices using meat inspection data from years 2004-2016, including inspection of 669,738 reindeer originating from Finnish reindeer herding areas. RESULTS: Increased stress and decreased welfare of reindeer, as indicated by higher incidence of carcass condemnation due to bruises or fractures, aspiration of rumen content, or abnormal odour, were positively associated with systems involving shorter transport distances to abattoirs. Significant differences in incidence of condemnations were also detected between abattoirs and reindeer herding cooperatives. CONCLUSIONS: This study indicates that in particular the short-distance transports of reindeer merit more attention. While the results suggest that factors associated with long distance transport, such as driver education, truck design, veterinary supervision, and specialist equipment, may be favourable to reducing pre-slaughter stress in reindeer when compared with short distance transport systems, which occur in a variety of vehicle types and may be done by untrained handlers. Further work is required to elucidate the causal factors to the current results.

Temporal patterns of moose-vehicle collisions with and without personal injuries.

Collisions with wild ungulates are an increasing traffic safety issue in boreal regions. Crashes involving smaller-bodied deer species usually lead to vehicle damage only, whereas collisions with a large animal, such as the moose, increase the risk of personal injuries. It is therefore important to understand both the factors affecting the number of moose-vehicle collisions (MVCs) and the underlying causes that turn an MVC into an accident involving personal injuries or fatalities. As a basis for temporal mitigation measures, we examined the annual and monthly variation of MVCs with and without personal injuries. Using a 22-year-long (1990-2011) time series from Finland, we tested the effect of moose population density and traffic volume on the yearly number of all MVCs and those leading to personal injuries. We also examined the monthly distribution of MVCs with and without personal injuries, and contrasted the Finnish findings with collision data from Sweden (years 2008-2010) and Norway (years 2008-2011). Both moose population abundance indices and traffic volume were positively related to the yearly variation in the number of MVCs in Finland. The proportion of MVCs involving personal injuries decreased during our 22-year study period. The monthly distribution of all MVCs peaked during the autumn or winter depending on country, while MVCs involving personal injury peaked in summer. Our study indicates that efforts to reduce MVCs involving personal injuries need to address driver awareness and attitudes during summer, despite most MVCs occurring in autumn or winter.

Serological evidence of tick-borne encephalitis virus infection in moose and deer in Finland: sentinels for virus circulation.

BACKGROUND: The incidence of tick-borne encephalitis (TBE) in humans has increased in Finland, and the disease has emerged in new foci. These foci have been investigated to determine the circulating virus subtype, the tick host species and the ecological parameters, but countrywide epidemiological information on the distribution of TBEV has been limited. METHODS: In this study, we screened sera from hunter-harvested wild cervids for the presence of antibodies against tick-borne encephalitis virus (TBEV) with a hemagglutination inhibition test. The positive results were confirmed by a neutralisation assay. RESULTS: Nine (0.74%) of 1213 moose, one (0.74%) of 135 white-tailed deer, and none of the 17 roe deer were found seropositive for TBEV. A close geographical congruence between seropositive cervids and recently reported human TBE cases was observed: nine of the ten seropositive animals were from known endemic areas. CONCLUSIONS: Our results confirm the local circulation of TBEV in several known endemic areas. One seropositive moose had been shot in an area where human TBE cases have not been reported, suggesting a possible new focus. Moose appear to be a useful sentinel animal for the presence of TBEV in the taiga region.

Moose (Alces alces) reacts to high summer temperatures by utilizing thermal shelters in boreal forests - an analysis based on airborne laser scanning of the canopy structure at moose locations.

The adaptation of different species to warming temperatures has been increasingly studied. Moose (Alces alces) is the largest of the ungulate species occupying the northern latitudes across the globe, and in Finland it is the most important game species. It is very well adapted to severe cold temperatures, but has a relatively low tolerance to warm temperatures. Previous studies have documented changes in habitat use by moose due to high temperatures. In many of these studies, the used areas have been classified according to how much thermal cover they were assumed to offer based on satellite/aerial imagery data. Here, we identified the vegetation structure in the areas used by moose under different thermal conditions. For this purpose, we used airborne laser scanning (ALS) data extracted from the locations of GPS-collared moose. This provided us with detailed information about the relationships between moose and the structure of forests it uses in different thermal conditions and we were therefore able to determine and differentiate between the canopy structures at locations occupied by moose during different thermal conditions. We also discovered a threshold beyond which moose behaviour began to change significantly: as day temperatures began to reach 20 °C and higher, the search for areas with higher and denser canopies during daytime became evident. The difference was clear when compared to habitat use at lower temperatures, and was so strong that it provides supporting evidence to previous studies, suggesting that moose are able to modify their behaviour to cope with high temperatures, but also that the species is likely to be affected by warming climate.

Combination treatment of elevated UVB radiation, CO2 and temperature has little effect on silver birch (Betula pendula) growth and phytochemistry.

Elevations of carbon dioxide, temperature and ultraviolet-B (UBV) radiation in the growth environment may have a high impact on the accumulation of carbon in plants, and the different factors may work in opposite directions or induce additive effects. To detect the changes in the growth and phytochemistry of silver birch (Betula pendula) seedlings, six genotypes were exposed to combinations of ambient or elevated levels of CO2 , temperature and UVB radiation in top-closed chambers for 7 weeks. The genotypes were relatively similar in their responses, and no significant interactive effects of three-level climate factors on the measured parameters were observed. Elevated UVB had no effect on growth, nor did it alter plant responses to CO2 and/or temperature in combined treatments. Growth in all plant parts increased under elevated CO2 , and height and stem biomass increased under elevated temperature. Increased carbon distribution to biomass did not reduce its allocation to phytochemicals: condensed tannins, most flavonols and phenolic acids accumulated under elevated CO2 and elevated UVB, but this effect disappeared under elevated temperature. Leaf nitrogen content decreased under elevated CO2 . We conclude that, as a result of high genetic variability in phytochemicals, B. pendula seedlings have potential to adapt to the tested environmental changes. The induction in protective flavonoids under UVB radiation together with the positive impact of elevated CO2 and temperature mitigates possible UVB stress effects, and thus atmospheric CO2 concentration and temperature are the climate change factors that will dictate the establishment and success of birch at higher altitudes in the future.

Dark-leaved willow (Salix myrsinifolia) is resistant to three-factor (elevated CO2, temperature and UV-B-radiation) climate change.

Elevated carbon dioxide (CO2 ), temperature (T) and ultraviolet-B (UV-B) radiation may affect plant growth and secondary chemistry in different directions, but the effect of the combination of the three factors has seldom been tested. Here, we grew four dark-leaved willow (Salix myrsinifolia) clones under combinations of ambient or elevated CO2, T and UV-B radiation in top-closed chambers for 7 wk. Elevated UV-B had no effects on growth or phenolic compounds, and there were no significant interactions between UV-B, CO2 and T. CO2 alone increased most growth parameters, but the magnitude of the effect varied among the clones. Total phenolics increased at elevated CO2 , whereas they decreased at elevated T. The responses varied between the clones. The results imply that dark-leaved willow are fairly resistant to the applied three-factor climate change, probably because of high constitutive defense. However, the interactions between clone and climate change factors implies that some clones are more susceptible than the species as a whole.

Climate change promotes the emergence of serious disease outbreaks of filarioid nematodes.

Filarioid parasites represent major health hazards with important medical, veterinary, and economic implications, and considerable potential to affect the everyday lives of tens of millions of people globally (World Health Organization, 2007). Scenarios for climate change vary latitudinally and regionally and involve direct and indirect linkages for increasing temperature and the dissemination, amplification, and invasiveness of vector-borne parasites. High latitude regions are especially influenced by global climate change and thus may be prone to altered associations and dynamics for complex host-pathogen assemblages and emergence of disease with cascading effects on ecosystem structure. Although the potential for substantial ecological perturbation has been identified, few empirical observations have emanated from systems across the Holarctic. Coincidental with decades of warming, and anomalies of high temperature and humidity in the sub-Arctic region of Fennoscandia, the mosquito-borne filarioid nematode Setaria tundra is now associated with emerging epidemic disease resulting in substantial morbidity and mortality for reindeer and moose. We describe a host-parasite system that involves reindeer, arthropods, and nematodes, which may contribute as a factor to ongoing declines documented for this ungulate species across northern ecosystems. We demonstrate that mean summer temperatures exceeding 14 degrees C drive the emergence of disease due to S. tundra. An association between climate and emergence of filarioid parasites is a challenge to ecosystem services with direct effects on public health, sustainability of free-ranging and domestic ungulates, and ultimately food security for subsistence cultures at high latitudes.

Foraging patterns of voles at heterogeneous avian and uniform mustelid predation risk.

Temporal variation of antipredatory behavior and a uniform distribution of predation risk over refuges and foraging sites may create foraging patterns different from those anticipated from risk in heterogenous habitats. We studied the temporal variation in foraging behavior of voles exposed to uniform mustelid predation risk and heterogeneous avian predation risk of different levels induced by vegetation types in eight outdoor enclosures (0.25 ha). We manipulated mustelid predation risk with weasel presence or absence and avian predation risk by reducing or providing local cover at experimental food patches. Foraging at food patches was monitored by collecting giving-up densities at artificial food patches, overall activity was automatically monitored, and mortality of voles was monitored by live-trapping and radiotracking. Voles depleted the food to lower levels in the sheltered patches than in the exposed ones. In enclosures with higher avian predation risk caused by lower vegetation height, trays were depleted to lower levels. Unexpectedly, voles foraged in more trays and depleted trays to lower levels in the presence of weasels than in the absence. Weasels match their prey's body size and locomotive abilities and therefore increase predation risk uniformly over both foraging sites and refuge sites that can both be entered by the predator. This reduces the costs of missing opportunities other than foraging. Voles changed their foraging strategy accordingly by specializing on the experimental food patches with predictable returns and probably reduced their foraging in the matrix of natural food source with unpredictable returns and high risk to encounter the weasel. Moreover, after 1 day of weasel presence, voles shifted their main foraging activities to avoid the diurnal weasel. This behavior facilitated bird predation, probably by nocturnal owls, and more voles were killed by birds than by weasels. Food patch use of voles in weasel enclosures increased with time. Voles had to balance the previously missed feeding opportunities by progressively concentrating on artificial food patches.

Intraspecific competition, growth, chemistry, and susceptibility to voles in seedlings of Betula pendula.

We studied the effects of the intensity of intraspecific competition, as indicated by seedling density, and competitive success within populations, as indicated by seedling size, on the secondary chemistry of the stems of silver birch seedlings and their palatability to field voles. We found that the size of seedlings and their total phenolic concentrations were inversely related to stand density. Voles, however, did not discriminate between seedlings grown at the densities studied. Variation in competitive success was associated with several phenomena. Seedling size was positively related to the concentrations of several secondary compounds and to vole preferences, but negatively related to nitrogen concentration. We suggest that the ecological consequences of variation in seedling size cannot be predicted without knowledge of the mechanism behind the variation.

Effects of different organic farming methods on the concentration of phenolic compounds in sea buckthorn leaves.

The effects of different cultivation methods on the amount of phenolic compounds in leaves of 1-year-old seedlings of two Finnish sea buckthorn (Hippophae rhamnoides L. ssp. rhamnoides) cultivars 'Terhi' and 'Tytti' were studied in a field experiment established at coastal area in Merikarvia, western Finland. The cultivation methods included different fertilizers (suitable for organic cultivation), mulches (organic and plastic), and land contours (flat vs low hill surface). Two experiments were conducted. The first allowed the estimation of the effects of cultivar, fertilizer, surface contour, and all their interactions, while the other allowed the estimation of the effects of mulches, land contours, and their interactions for the cultivar 'Tytti'. Eleven different hydrolyzable tannins, pentagalloylglucose, and 14 other phenolic compounds were detected by chemical analysis with high-performance liquid chromatography (HPLC). The amount of phenolic compounds varied between different land contours and mulches. The concentrations of gallic acid, pentagalloylglucose, quercetin-3-rhamnoside, monocoumaroyl astragalin A, total hydrolyzable tannins, and condensed tannins were significantly higher on the flat surface than on the low hill surface. The plastic mulch decreased the concentration of gallic acid, hydrolyzable tannins, and condensed tannins compared to the other mulches used. These results suggest ways to cultivate sea buckthorn to produce large amounts of valuable chemicals, especially tannins in the leaves.

Variations in phenology and growth of European white birch (Betula pendula) clones.

Phenology can have a profound effect on growth and climatic adaptability of northern tree species. Although the large interannual variations in dates of bud burst and growth termination have been widely discussed, little is known about the genotypic and spatial variations in phenology and how these sources of variation are related to temporal variation. We measured bud burst of eight white birch (Betula pendula Roth) clones in two field experiments daily over 6 years, and determined the termination of growth for the same clones over 2 years. We also measured yearly height growth. We found considerable genetic variation in phenological characteristics among the birch clones. There was large interannual variation in the date of bud burst and especially in the termination of growth, indicating that, in addition to genetic effects, environmental factors have a strong influence on both bud burst and growth termination. Height growth was correlated with timing of growth termination, length of growth period and bud burst, but the relationships were weak and varied among years. We accurately predicted the date of bud burst from the temperature accumulation after January 1, and base temperatures between +2 and -1 degrees C. There was large clonal variation in the duration of bud burst. Interannual variation in bud burst may have important consequences for insect herbivory of birches.

Variation in predation risk and vole feeding behaviour: a field test of the risk allocation hypothesis.

Many prey animals experience temporal variation in the risk of predation and therefore face the problem of allocating their time between antipredator efforts and other activities like feeding and breeding. We investigated time allocation of prey animals that balanced predation risk and feeding opportunities. The predation risk allocation hypothesis predicts that animals should forage more in low- than in high-risk situations and that this difference should increase with an increasing attack ratio (i.e. difference between low- and high-risk situations) and proportion of time spent at high risk. To test these predictions we conducted a field test using bank voles ( Clethrionomys glareolus) as a prey and the least weasel ( Mustela nivalis nivalis) as a predator. The temporal pattern and intensity of predation risk were manipulated in large outdoor enclosures and the foraging effort and patch use of voles were measured by recording giving-up densities. We did not observe any variation in feeding effort due to changes in the level of risk or the proportion of time spent under high-risk conditions. The only significant effect was found when the attack ratio was altered: the foraging effort of voles was higher in the treatment with a low attack ratio than in the treatment with a high attack ratio. Thus the results did not support the predation risk allocation hypothesis and we question the applicability of the hypothesis to our study system. We argue that the deviation between the observed pattern of feeding behaviour of bank voles and that predicted by the predation risk allocation hypothesis was mostly due to the inability of voles to accurately assess the changes in the level of risk. However, we also emphasise the difficulties of testing hypotheses under outdoor conditions and with mammals capable of flexible behavioural patterns.

Genotypic variation in growth and resistance to insect herbivory in silver birch (Betula pendula) seedlings.

If herbivory is unevenly distributed among different types of plants, or if individual plants differ in their response to herbivory, herbivores may affect seedling growth and survival, and consequently plant population structure. In this study we examined variation in resistance to insect herbivory and in growth responses to feeding among 20 silver birch (Betula pendula Roth) genotypes representing variation within a natural population. Birch seedlings were grown in dense stands in random arrangement so that seedlings of different genotypes competed with each other. On insect exposure plots natural colonization of insects was allowed, and insect removal plots were sprayed with insecticide. Resistance to insect herbivory was measured as the inverse of leaf damage, and growth responses of seedlings to feeding were determined as the change in seedling height relative to the amount of damage. Resistance varied significantly among genotypes, but growth responses to feeding did not. In fertilized seedlings, resistance correlated negatively with height, indicating a trade-off between resistance and growth. The absence of such a correlation in non-fertilized seedlings is a sign of environmental effects on the cost of resistance. Growth responses to feeding did not correlate with either resistance or growth. Nevertheless, different effects of the actual damage on height increase changed the positions of the genotypes in the size hierarchy of the experimental populations. Thus, even moderate levels of insect herbivory can change the outcome of competitive interactions between birch genotypes.

Choice of voles among genotypes of birch seedlings: its relationship with seedling quality and preference of insects.

Selective feeding by herbivores on establishing seedlings has been suggested to affect genotype frequencies in several plant populations. The existence of genotypes susceptible to herbivores calls for an explanation in such populations. In the present study we assessed the choice of multiple herbivores, field voles (Microtus agrestis) and insects, among genotypes of silver birch (Betula pendula) representing variation occurring in a naturally regenerated stand. We examined how food choice of voles and insects is related to each other, competitive ability among the seedling genotypes and variation in soil fertility. We set up a field experiment and randomly assigned seedling populations, composed of mixed genotypes, to fully crossed insect exposure and fertilization treatments. After the first growing season we exposed a half of the seedling populations to vole herbivory. Voles selected clearly among the genotypes: they preferred the fastest growing seedlings as well as those with a low density of resin droplets on their stems. The preference of voles and insects among the genotypes was tightly correlated. We conclude that the effects of herbivory compensate those of intraspecific competition in this system and thus favor coexistence of genotypes differing in their susceptibility to herbivores.