Moose and vole browsing patterns in experimentally assembled pure and mixed forest stands

Plants growing in diverse communities are believed to exhibit associational resistance to herbivores, but this hypothesis has seldom been tested experimentally for vertebrate herbivores in forest ecosystems. We examined browsing patterns of the two principal mammalian herbivores of Finnish boreal forests, moose and voles, in young stands where tree species diversity and composition were experimentally manipulated. The stands were composed either of monocultures or different 2–5 species mixtures of Norway spruce, Scots pine, Siberian larch, silver birch, and black alder. Voles and moose showed contrasting responses to stand diversity and species composition. In accordance with the predictions of the associational resistance hypothesis, vole damage was higher in tree monocultures than in mixed stands, although stand diversity effects were statistically significant only at one of the three study areas. Voles also damaged more trees in coniferous than in deciduous stands. In contrast, moose browsing tended to increase with the number of tree species in a stand and with the presence of the preferred tree species, birch, in a mixture. The observed differences in vole and moose responses to stand diversity and species composition are likely to be due to different feeding specialisation, foraging patterns, and movement ability of these herbivores. Voles switched to trees only when the supply of a more preferred food (grasses and forbs) was depleted and restricted their feeding choice only to the most palatable tree species regardless of the number of tree species present per stand. In contrast, tree branches and foliage represented an important part of moose diet throughout the year; moose may be able to tolerate secondary plant metabolites of different tree species better than voles and may thus benefit from diet broadening when more tree species are available. Furthermore, the home range size and foraging ability of these two very differently sized herbivores may partly explain the observed differences in utilisation of different tree species. Finally, both moose and voles showed high spatial and temporal variation in their feeding; in particular, vole damage was more influenced by tree species diversity in areas and years with high vole densities. Thus, diversification of forest stands may have very different effects on mammalian browsing depending on the herbivores present, their densities, and the tree species used in reforestation.     

Varying impacts of cervid, hare and vole browsing on growth and survival of boreal tree seedlings

The negative impacts of mammalian herbivores on plants have been studied quite extensively, but typically with only a single herbivore species at a time. We conducted a novel comparison of the browsing effects of voles, hares and cervids upon the growth and survival of boreal tree seedlings. This was done by excluding varying assemblages of these key mammalian herbivores from silver birch, Scots pine and Norway spruce seedlings for 3 years. We hypothesised that the pooled impacts of the herbivores would be greater than that of any individual group, while the cervids would be the group with the strongest impact. Growth of birch seedlings advanced when cervids were excluded whereas growth of seedlings accessible to cervids was hindered. Survival of all seedlings was lowest when they were accessible to voles and voles plus hares, whereas cervids seemed not to influence seedling survival. Our results show that the impact of herbivores upon woody plants can be potent in the boreal forests, but the mechanism and strength of this link depends on the tree and herbivore species in question. Risk of abated stand regeneration appears highest for the deciduous birch, though there is need for seedling protection also in coniferous stands. The clear cervid-mediated growth limitation of birch also indicates potential for a trophic cascade effect by mammalian top predators, currently returning to boreal ecosystems.     

Demographic responses to a mild winter in enclosed vole populations

Mild winter weather causing snow to melt and ice to accumulate on the ground has been proposed to cause the decreased survival of individuals, and less pronounced cyclicity, of small rodent populations in Fennoscandia. However, detailed data linking ice accumulation to decreased winter survival is lacking. We live-trapped and monitored with passive integrated transponders enclosed populations of root voles (Microtus oeconomus) exposed to different amounts of ice accumulation through a mild winter. We studied how social behaviour and survival responded to snow melt and ice accumulation. Voles avoided ground ice by moving their home ranges, thus increasing home range overlap in enclosed populations experiencing more extensive ice cover. Winter survival was not affected by the amount of ice accumulation, and was only slightly reduced during ice formation in early winter. The lowest survival rates were found at the onset of snow melt in early spring. These results suggest that ice accumulation does not cause lower survival during mild winters, probably because plastic social behaviour enables root voles to reduce the negative effects of varying winter weather on survival. The mechanisms for lower survival during mild winters may operate during spring and be related to spring floods or increased susceptibility to predators. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Allocation of resources within mountain birch canopy after simulated winter browsing

As a response to browsing, birches are known to produce fewer but larger, more nutritious leaves, with enhanced palatability for herbivores. We simulated winter browsing in ramets of mountain birch ( Betula pubescens ssp. czerepanovii ) to find out whether it decreases subsequent foliage biomass and alters the number and type of shoots. After removal of a considerable proportion of buds (up to 35%) in late winter, the birches were able to compensate for the lost leaf biomass in the following summer; there were no differences in total leaf biomass between winter-clipped and control ramets. This indicates that foliage growth was limited by the total amount of stored resources, not by the number of buds. Depending on the position of the buds removed, different mechanisms were responsible for the compensation. After removal of apical buds, the number of leaves decreased significantly but leaves were larger than in control ramets. Removal of the same mass of basal buds – containing similar amount of carbohydrates and proteins as in the treatment removing apical buds – activated dormant buds, especially in apical locations, so that leaf number was similar as in the controls; consequently, size of individual leaves increased only slightly. Thus, while the total leaf biomass in a tree seems to be limited by resources from source organs, the distribution of resources among different canopy sections is controlled by their relative sink strengths. In terms of leaf biomass, apical parts are able to compensate for bud loss by increasing shoot number, basal parts only by increasing leaf size.     

Density-dependent reproduction causes winter crashes in a common vole population

Common voles in western France exhibit three-year population cycles with winter crashes after large outbreaks. During the winter of 2011–2012, we monitored survival, reproduction, recruitment and population growth rate of common voles at different densities (from low to outbreak densities) in natura to better understand density dependence of demographic parameters. Between October and April, the number of animals decreased irrespective of initial density. However, the decline was more pronounced when October density was higher (loss of ≈54 % of individuals at low density and 95 % at high density). Using capture-mark-recapture models with Pradel's temporal symmetry approach, we found a negative effect of density on recruitment and reproduction. In contrast, density had a slightly positive effect on survival indicating that mortality did not drive the steeper declines in animal numbers at high density. We discuss these results in a population cycle framework, and suggest that crashes after outbreaks could reflect negative effects of density dependence on reproduction rather than changes in mortality rates.     

An experimental test of frequency-dependent food selection: winter browsing by moose

Penned moose Alces alces (L.) offered birch and rowan trees in various proportions in experimental tests showed a clear preference for birch both in terms of selection and use of trees. Both tree selection and tree use was frequency-independent, i.e. the preference for birches did not vary with either the number of birches, or total birch biomass available. The preference for birch was not absolute, i.e. rowans were also present in the diet. Birch were more profitable than rowans in terms of biomass intake rate, whereas rowans were more "nutritious" than birches. The results are consistent with both the optimal diet theory and the "nutrient mixture" hypothesis, although neither of the hypotheses are unambiguous regarding frequency-dependent food selection. It is argued, however, that tree selection and use by moose rather should be considered as a patch use problem and that trees should not be regarded as particulate food items as in most foraging models.     

Effects of nutrition and peripheral stimuli on preferences for familiar foods in the bank vole

Individual bank voles were given one of two nutritionally identical experimental diets which differed in taste and colour. Voles which were background-fed with lab diet during the experiment showed a long-lasting increase in preference for the recently-available experimental diet, while voles which were not background-fed showed a short-term (two-day) decrease in preference for the recently available experimental diet. The results showed that nutritional cues are important in determining which way the preference changes, and that learning is involved in associating peripheral cues such as colour or taste with the nutritional effects.     

MHC influences infection with parasites and winter survival in the root vole Microtus oeconomus

Selective pressure from parasites is thought to maintain the polymorphism of major histocompatibility complex (MHC) genes. Although a number of studies have shown a relationship between the MHC and parasitic infections, the fitness consequences of such associations are less well documented. In the present paper, we characterised the variation in exon 2 of MHC class II DRB gene in the root vole and examined the effects of that gene on parasite prevalence and winter survival. We identified 18 unique exon 2 sequences, which translated into 10 unique amino acid sequences. Phylogenetic analysis revealed the presence of three distinct clusters, and allele distributions among these individuals suggested that the clusters correspond to three different loci. Although the rate of synonymous substitutions (d_S) exceeded the rate of nonsynonymous substitutions (d_N) across sequences, implying purifying selection, d_N was significantly elevated at antigen-binding sites, suggesting that these sites could be under positive selection. Screening for parasites revealed a moderate prevalence of infection with gastrointestinal parasites (24 % infected), but a high infection rate for blood parasites (56 % infected). Infection with the blood parasite Babesia ssp. decreased survival almost twofold (25.7 vs. 13.9 %). Animals possessing the amino acid sequence AA*08 survived better than others (44.9 vs. 22 %), and they were infected with Babesia ssp. less often (13.9 vs 25.7 %). In contrast, individuals carrying allele AA*05 were infected more often (31.7 vs. 15.3 %). Heterozygosity at one of the putative loci was associated with a lower probability of infection with Babesia ssp., but at the other locus, the association was reversed. The unexpected latter result could be at least partly explained by the increased frequency of the susceptible allele AA*05 among heterozygotes. Overall, we demonstrate that infection with Babesia ssp. is a strong predictor of winter survival and that MHC genes are important predictors of infection status as well as survival in the root vole.     

Effects of simulated winter browsing on mountain birch foliar chemistry and on the performance of insect herbivores

Winter browsing by mammalian herbivores is known to induce a variety of morphological and physiological changes in plants. Browsing has been suggested to decrease the carbohydrate reserves in woody plants, which might lead to reduced tannin production in leaves during the following summer, and consequently, to increased herbivore damage on leaves. We conducted a clipping experiment with mature mountain birch trees and measured the effects of clipping on birch growth, leaf chemistry and toughness, as well as on the performance of insect herbivores. Leaves grew larger and heavier per unit area in the clipped ramets and had a higher content of proteins than leaves in the control trees. Clipping treatment did not affect the total content of sugars in the leaves (mg g^?1), suggesting that a moderate level of clipping did not significantly reduce the carbohydrate pools of fully‐grown mountain birch trees. Furthermore, the contents of proanthocyanidins (condensed tannins) and gallotannins were slightly higher in the leaves of clipped ramets, contrary to the hypothesis of reduced tannin production. The effects of clipping treatment on leaf and shoot growth and on foliar chemistry were mainly restricted to the clipped ramets, without spreading to untreated ramets within the same tree individual. The effects of clipping on leaf characters varied during the growing season; for instance, leaf toughness in clipped ramets was higher than toughness in control trees and ramets only when leaves were mature. Accordingly, clipping had inconsistent effects on insect herbivores feeding at different times of the growing season. The generally small impact of clipping on herbivore performance suggests that the low intensity of natural browsing at the study area, simulated by our clipping treatment, does not have strong consequences for the population dynamics of insect herbivores on mountain birch via enhanced population growth caused by browsing‐induced changes in food quality.     

Anemia at the onset of winter in the meadow vole (Microtus pennsylvanicus)

1. From 1981 to 1986, 6120 meadow voles (Microtus pennsylvanicus) were sampled for hematological indices in southeastern Manitoba, Canada. This survey revealed the sporadic occurrence of anemia in early winter at mean temperatures below about -5 degrees C. 2. Anemia was associated with leukocytosis and circulating normoblasts, suggesting a sudden, large blood loss. Individuals became anemic quickly, with no obvious predisposing factors. 3. Attempts were made to induce anemia by exposing voles in traps to various temperatures. Temperatures characteristic of most trapping sessions failed to induce anemia in both wild and laboratory-born voles. 4. Short-term exposure to more extreme temperatures (-20 to -30 degrees C) induced anemia. Voles lost blood through erosions of the epithelium of the glandular stomach, and developed other pathological lesions characteristic of hypothermia. 5. Although there was a strong association between cold weather and anemia, we could find no clear relationship between winter survival and winter weather. However, in 1984, extraordinarily cold temperatures were associated with anemia and a subsequent population decline. These events suggest a threshold mean daily temperature of about -15 degrees C, below which vole survival is grossly affected. 6. Deteriorating protein levels and energy reserves of small mammals in winter may make them particularly susceptible to cold stress. Hence, sporadic bouts of sustained cold may be responsible for some of the enigmatic winter declines seen in northern small mammals.     

Is habitat selection by the Cabrera vole (Microtus cabrerae) related to food preferences?

The Cabrera vole (Microtus cabrerae) is an Iberian endemism considered threatened both in Portugal and Spain. This vole is often referred to as a tall perennial grassland community specialist. However, a recent study revealed that the Cabrera vole exploits a wide variety of grasslands in south Portugal. The major aim of the present study was to test the palatability of 21 plant species of this region occurring within or outside typical perennial grassland colonies in order to assess vole's food preferences. Results showed a major consumption of monocotyledons and annual plants (such as Festuca ampla, Agrostis castellana, Brachypodium distachyon, B. phoenicoides and Vulpia geniculata), but no relationship was found between diet selection and perennial grassland plant communities. The preference of voles for annual species, occurring more abundantly outside the colonies, raises the hypothesis that the establishment of a colony might be determined by plant species abundance and persistence over the year, rather than by their palatability. This seems to be particularly evident in this region, where summer meteorological conditions make vegetation unavailable outside the colonies.     

Delayed induced responses of birch glandular trichomes and leaf surface lipophilic compounds to mechanical defoliation and simulated winter browsing

Changes in morphology and chemistry of leaf surface in response to herbivore damage may increase plant resistance to subsequent herbivore attack; however, there is lack of studies on induced responses of glandular trichomes and their exudates in woody plants and on effects of these changes on herbivores. We studied delayed induced responses in leaf surface traits of five clones of silver birch (Betula pendula Roth) subjected to various types of mechanical defoliation and simulated winter browsing. Glandular trichome density and concentrations of the majority of surface lipophilic compounds increased in trees defoliated during the previous summer. This induced response was systemic, since control branches in branch defoliated trees responded to the treatments similarly to defoliated branches, but differently from control trees. In contrast to defoliation treatments, simulated winter browsing reduced glandular trichome density on the following summer and had fewer effects on individual surface lipophilic compounds. Moreover, constitutive density of glandular trichomes was negatively correlated with induced total amount of lipophilic compounds per trichome, indicating a trade-off between constitutive and induced resistance in silver birch. Induced changes in leaf surface traits had no significant effect on leaf damage by chewers, miners and gall mites, but increased susceptibility of birch trees to aphids. However, leaf damage by chewers, miners and gall mites in defoliated (but not in control) trees was correlated with concentrations of some fatty acids and triterpenoids, although the direction of relationships varied among herbivore species. This indicates that induction of surface lipophilic compounds may influence birch resistance to herbivores. Our study thus demonstrated both specificity of elicitation of induced responses of birch leaf surface traits by different types of damage and specificity of the effects of these responses on different types of herbivores.Electronic Supplementary Material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Heightened sour preferences during childhood

Basic research has revealed that the chemical sensory world of children is different from that of adults, as evidenced by their heightened preferences for sweet and salty tastes. However, little is known about the ontogeny of sour taste preferences, despite the growing market of extreme sour candies. The present study investigated whether the level of sourness most preferred in a food matrix and the ability to discriminate differences in sour intensity differed between 5- to 9-year-old children and their mothers, by using a rank-by-elimination procedure embedded in the context of a game. Mothers also completed a variety of questionnaires and children were asked several questions to assess whether children's temperament and food preferences and habits related to sour preferences. The results indicated that, although every mother and all but two of the children (92%) were able to rank the gelatins from most to least sour, more than one-third (35%) of the children, but virtually none of the adults, preferred the high levels of sour taste (0.25 M citric acid) in gelatin. Those children who preferred the extreme sour tastes were significantly less food neophobic (P < 0.05) and tended to experience a greater variety of fruits when compared with the remaining children (P = 0.11). Moreover, the children's preference for sour tastes generalized to other foods, such as candies and lemons, as reported by both children and mothers. These findings are the first experimental evidence to demonstrate that sour taste preferences are heightened during childhood and that such preferences are related to children's food habits and preferences. Further research is needed to unfold the relationship between the level of sour taste preferred and the actual consumption of sour-tasting foods and flavors in children.     

Traditional and diversified crops in South Moravia (Czech Republic): Habitat preferences of common vole and mice species

Substantial changes in the composition of crops in Central Europe during the last two decades (increasing areas of maize, rape and sunflower fields) have significantly influenced the structure and dynamics of animal communities, though there is a lack of data available for small rodents. In this study, we assessed the importance of these three crops for rodents and compared it with traditional crops (alfalfa, barley, wheat). We observed that herbivorous species (especially the common vole) do not live in crops which do not have green leaves near the ground (e.g. sunflower, maize), while mobile granivorous species can inhabit all types of crop. We confirmed the presumed differences in habitat preferences; however we rejected the hypothesis of a general increase of rodent abundance during the vegetative season in managed fields. We found that (1) maize and sunflower had no importance for common voles, but they were favored habitats for wood mice; (2) numbers of wood mice in rape decreased during the season, while abundances of common voles increased; (3) common vole populations tended to increase during the season in all suitable crops (alfalfa, barley, wheat, tendency in maize and rape); (4) wood mice populations seemed stable in all crops; i.e. without a seasonal increase. It can be concluded that even if the new crop fields are an important part of the agricultural landscape, they are only a temporal habitat for small mammals, especially granivorous species.     

Surviving winter: Food,but not habitat structure,prevents crashes in cyclic vole populations

Vole population cycles are a major force driving boreal ecosystem dynamics in northwestern Eurasia. However, our understanding of the impact of winter on these cycles is increasingly uncertain, especially because climate change is affecting snow predictability, quality, and abundance. We examined the role of winter weather and snow conditions, the lack of suitable habitat structure during freeze‐thaw periods, and the lack of sufficient food as potential causes for winter population crashes. We live‐trapped bank voles Myodes glareolus on 26 plots (0.36 ha each) at two different elevations (representing different winter conditions) in southeast Norway in the winters 2013/2014 and 2014/2015. We carried out two manipulations: supplementing six plots with food to eliminate food limitation and six plots with straw to improve habitat structure and limit the effect of icing in the subnivean space. In the first winter, all bank voles survived well on all plots, whereas in the second winter voles on almost all plots went extinct except for those receiving supplemental food. Survival was highest on the feeding treatment in both winters, whereas improving habitat structure had no effect. We conclude that food limitation was a key factor in causing winter population crashes.     

Spatial correlations between browsing on balsam fir by white‐tailed deer and the nutritional value of neighboring winter forage

Associational effects, that is, the influence of neighboring plants on herbivory suffered by a plant, are an outcome of forage selection. Although forage selection is a hierarchical process, few studies have investigated associational effects at multiple spatial scales. Because the nutritional quality of plants can be spatially structured, it might differently influence associational effects across multiple scales. Our objective was to determine the radius of influence of neighbor density and nutritional quality on balsam fir (Abies balsamea) herbivory by white‐tailed deer (Odocoileus virginianus) in winter. We quantified browsing rates on fir and the density and quality of neighboring trees in a series of 10‐year‐old cutovers on Anticosti Island (Canada). We used cross‐correlations to investigate relationships between browsing rates and the density and nutritional quality of neighboring trees at distances up to 1,000 m. Balsam fir and white spruce (Picea glauca) fiber content and dry matter in vitro true digestibility were correlated with fir browsing rate at the finest extra‐patch scale (across distance of up to 50 m) and between cutover areas (300–400 m). These correlations suggest associational effects, that is, low nutritional quality of neighbors reduces the likelihood of fir herbivory (associational defense). Our results may indicate associational effects mediated by intraspecific variation in plant quality and suggest that these effects could occur at scales from tens to hundreds of meters. Understanding associational effects could inform strategies for restoration or conservation; for example, planting of fir among existing natural regeneration could be concentrated in areas of low nutritional quality.     

The biomechanics of browsing and grazing

Terrestrial plant leaves are complex structures of composite materials. Resistance to fracture is achieved by a number of mechanisms, which operate at the molecular, cell, tissue, and structural levels. Leaves of dicots have different venation patterns and cell wall volume fractions from those of grasses, and consequently, they potentially resist fracture in different ways. Animals mechanically process plants in order to rupture the cell wall in preparation for enzymic hydrolysis, for which the imperative is to maximize new surface area and/or to expedite access to cell contents, ideally by promoting elastic fracture. The two different plant types are fed on by two different groups of organisms of very different sizes, digestive physiologies, mechanical processing abilities and properties, and nutritional requirements. Small insects can feed in or on parts of leaves, while larger mammals generally have to feed on the whole leaf. The scale of feeding also differs for the two groups of herbivores, but how this interacts with the scale of the mechanical properties of the leaf is not well understood. Plant leaves are attacked at all scales and probably can only produce generalized responses to specialized herbivores. In addition, the opportunities that these different scales of interactions open for the different herbivores remain unexplored.     

Interaction between acquired preferences and developing predispositions during imprinting

Preferences of domestic chicks were studied either at approximately 2 h or approximately 24 h after exposure to a red box or a stuffed jungle fowl. Preferences of the chicks were measured in a simultaneous choice test involving these two stimuli. When tested at about 2 h, chicks preferred the object to which they had previously been exposed. Chicks tested at about 24 h showed a significantly greater preference for the jungle folw than those tested at about 2 h, regardless of the training stimulus. Birds that had been placed in a running wheel and exposed to diffuse light also showed an increasing preference for the fowl over time when tested in the same manner. These results may be accounted for in terms of an interaction between two processes, a developing predisposition for the fowl on the one hand and a preference acquired through learning on the other.