Variation in quality of mountain birch and tea-leaved willow for mammal and insect herbivores: differences among trees, among sites and between tree species

We evaluated palatability of winter dormant tree twigs to the mountain hare Lepus tunidus and the grey-sided vote Clethrionomys rufocanus with captive animals We tested differences among trees within sites, among sites, and between two tree species (the mountain birch Betula pubescens ssp czerepanovii and the tea-leaved willow Salix phylicifolia ) In one of the sites, we also measured growth rates of autumnal moth Epirrita autumnata larvae on the same trees that were used m preference trials with the hares and voles The differences m palatability to hares and voles were greatest at the level of tree Species, both hares and voles preferred birches over willows, but with the hare there was some overlap in palatability between the tree species There were also large and significant differences among sites and among trees within sites Within sites, variation in tree palatability seemed to be larger among willows than among birches Hares and voles may select willows at least partly on the same basis, but there was no correlation between palatability of the trees to the mammals and the growth rate of the larvae of the autumnal moth on the same trees Variation in twig palatability to the hares and voles was so large at all levels (among trees within sites, among sites and between tree species) that we suggest it also has implications for food selection of these mammalian herbivores under natural conditions     

Regulation of boreal forest understory vegetation: the roles of resources and herbivores

To understand inter-trophic linkages between components of the boreal forest understory vegetation, three hypotheses were tested: survival, growth and abundance of grasses and legumes were controlled by (i) resource availability alone, (ii) by herbivores alone, and (iii) by both resource availability and herbivores. The hypotheses were tested using three experimental treatments – fertilization, herbivore exclusion, and fertilization plus herbivore exclusion – in three areas having different densities of resident herbivores, mostly snowshoe hares and ground squirrels. The highest density of snowshoe hares is comparable to natural levels during peaks in the snowshoe hare cycle. As the density of herbivores increased so too did the level of response by the measured variables – survival, growth of transplants and leaf area index of established vegetation. In general, fertilization resulted in a decrease in survival and growth of transplants, and fences increased survival and growth; both responses were more noticeable at higher herbivore densities. Fertilizer and herbivore exclosure fences had only negligible effects on established grass and legume abundance at all hare densities. We have shown that some hypotheses of vegetation regulation are over-simplified because different species groups (i.e., grasses and legumes) are regulated by different factors, at different life history stages, and sometimes these factors act in opposing directions. We argue that during the increase phase and peak of the snowshoe hare cycle (high herbivore density), growth and survival of establishing plants is regulated by herbivores. During the decline and low phases of the snowshoe hare cycle herbivores will have little impact on early life stages, whereas the established, mature, vegetation will be resource-regulated. Because of the variability in responses to the same manipulations we may begin to understand which plant life history stages are most vulnerable to consumer and resource regulation, the magnitudes of these sources of regulation at each of these stages, and how these vary among species groups and types of environments.     

Foliage value,apparency and defence investment in birch seedlings and trees

Summary Two factors determining plant anti-herbivore defence investment fitness loss due to herbivory and the probability of herbivory occurring in the field were quantified for birch seedlings and trees. Fitness loss due to defoliation (assumed to be related to loss of growth increment compared to controls) appeared to be greater in seedlings compared to trees, but the result was equivocal. In contrast, seedling foliage at the field site — a typical habitat for birch — suffered much less natural defoliation than tree foliage, suggesting that seedlings are markedly less apparent to most birch herbivores than trees. This low apparency should result in lower investment in anti-herbivore defences by seedlings compared to trees — and being a strong effect, should outweigh the possibly greater growth loss suffered by seedlings, which in isolation would tend to increase their optimum defence investment compared to trees. This prediction was tested using palatability trials with a wide range of common birch herbivores and by direct quantification of anti-herbivore defences. Problems and assumptions inherent in these approaches are discussed, but it seems that birch seedlings are genuinely unapparent to herbivores, and consequently do not need the degree of defence investment required by trees.     

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.     

Delayed local responses of downy birch to damage by leafminers and leafrollers

Recent findings suggest that impacts of endemic herbivory on forest ecosystems over the long term may exceed impacts of herbivore outbreaks. However, responses of trees to minor and local damage imposed by small arthropod herbivores, especially by those mining or skeletonising individual leaves, remain poorly understood. We studied the delayed effects of injuries by several leafmining and leafrolling insects on the performance of downy birch shoots. Insect feeding did not affect survival of shoots or survival of individual axillary buds in long shoots. In the year following the damage, shoots produced an average of 13.8% more biomass than undamaged shoots of the same tree. The magnitude of this effect increased with an increase in the leaf area injured during the previous year, but it did not differ among four localities in subarctic and boreo‐nemoral forests, between herbivore feeding guilds, or among herbivores imposing damage in early, mid and late summer. We also found that herbivores attacked the next‐year foliage produced by damaged shoots less frequently than they attacked the next‐year foliage produced by undamaged shoots of the same tree. Thus, our study demonstrated delayed local compensatory growth and increased antiherbivore defence in downy birch shoots following local damage by insect feeding. We suggest that this pattern reflects evolutionary adaptations of plants to permanently acting minor, dispersed and spatially unpredictable damage imposed by endemic herbivory. Local responses are less costly and represent a more sustainable strategy to maintain plant fitness under low levels of herbivory than constitutive resistance or systemic responses.     

Tree species diversity influences herbivore abundance and damage: meta-analysis of long-term forest experiments

Plant monocultures are commonly believed to be more susceptible to herbivore attacks than stands composed of several plant species. However, few studies have experimentally tested the effects of tree species diversity on herbivory. In this paper, we present a meta-analysis of uniformly collected data on insect herbivore abundance and damage on three tree species (silver birch, black alder and sessile oak) from seven long-term forest diversity experiments in boreal and temperate forest zones. Our aim was to compare the effects of forest diversity on herbivores belonging to different feeding guilds and inhabiting different tree species. At the same time we also examined the variation in herbivore responses due to tree age and sampling period within the season, the effects of experimental design (plot size and planting density) and the stability of herbivore responses over time. Herbivore responses varied significantly both among insect feeding guilds and among host tree species. Among insect feeding guilds, only leaf miner densities were consistently lower and less variable in mixed stands as compared to tree monocultures regardless of the host tree species. The responses of other herbivores to forest diversity depended largely on host tree species. Insect herbivory on birch was significantly lower in mixtures than in birch monocultures, whereas insect herbivory on oak and alder was higher in mixtures than in oak and alder monocultures. The effects of tree species diversity were also more pronounced in older trees, in the earlier part of the season, at larger plots and at lower planting density. Overall our results demonstrate that forest diversity does not generally and uniformly reduce insect herbivory and suggest instead that insect herbivore responses to forest diversity are highly variable and strongly dependent on the host tree species and other stand characteristics as well as on the type of the herbivore.     

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.     

Large and small herbivores have strong effects on tundra vegetation in Scandinavia and Alaska

1. Large and small mammalian herbivores are present in most vegetated areas in the Arctic and often have large impacts on plant community composition and ecosystem functioning. The relative importance of different herbivores and especially how their specific impact on the vegetation varies across the Arctic is however poorly understood.
2. Here, we investigate how large and small herbivores influence vegetation density and plant community composition in four arctic vegetation types in Scandinavia and Alaska. We used a unique set of exclosures, excluding only large (reindeer and muskoxen) or all mammalian herbivores (also voles and lemmings) for at least 20 years.
3. We found that mammalian herbivores in general decreased leaf area index, NDVI, and abundance of vascular plants in all four locations, even though the strength of the effect and which herbivore type caused these effects differed across locations. In three locations, herbivore presence caused contrasting plant communities, but not in the location with lowest productivity. Large herbivores had a negative effect on plant height, whereas small mammalian herbivores increased species diversity by decreasing dominance of the initially dominating plant species. Above‐ or belowground disturbances caused by herbivores were found to play an important role in shaping the vegetation in all locations.
4. Synthesis: Based on these results, we conclude that both small and large mammalian herbivores influence vegetation in Scandinavia and Alaska in a similar way, some of which can mitigate effects of climate change. We also see important differences across locations, but these depend rather on local herbivore and plant community composition than large biogeographical differences among continents.

     

Herbivory and plant growth rate determine the success of El Niño Southern Oscillation-driven tree establishment in semiarid South America

While climatic extremes are predicted to increase with global warming, we know little about the effect of climatic variability on biome distribution. Here, we show that rainy El Niño Southern Oscillation (ENSO) events can enhance tree recruitment in the arid and semiarid ecosystems of north‐central Chile and northwest Peru. Tree‐ring studies in natural populations revealed that rainy El Niño episodes have triggered forest regeneration in Peru. Field experiments indicate that tree seedling recruitment in Chile is much less successful than in Peru due mostly to larger mortality caused by herbivores. The dramatic impact of herbivores in Chile was derived from the combined result of slower plant growth and the presence of exotic herbivores (European rabbits and hares). The interplay of herbivory and climatic effects we demonstrated implies that rainy ENSO events may represent ‘windows of opportunity’ for forest recovery if herbivore pressure is minimized at the right moment.     

Seedling Mortality and Herbivory Damage in Subtropical and Temperate Populations: Testing the Hypothesis of Higher Herbivore Pressure Toward the Tropics

Herbivory rates are generally thought to be higher in tropical than in temperate forests. Nevertheless, tests of this biogeographic prediction by comparing a single plant species across a tropical-temperate range are scarce. Here, we compare herbivore damage between subtropical and temperate populations of the evergreen tree Aextoxicon punctatum (Olivillo), distributed between 30° and 43° S along the Pacific margin of Chile. To assess the impact of herbivory on Olivillo seedlings, we set up 29 experimental plots, 1.5 × 3 m: 16 in forests of Fray Jorge National Park (subtropical latitude), and 13 in Guabún, Chiloé Island (temperate latitude). Half of each plot was fenced around with chicken wire, to exclude small mammals, and the other half was left unfenced. In each half of the plots we planted 16 seedlings of Olivillo in December 2003, with a total of 928 plants. Seedling survival, leaf production and herbivory by invertebrates were monitored over the next 16 mo. Small mammal herbivores killed ca 30 percent of seedlings in both sites. Nevertheless, invertebrate herbivory was greater in the temperate forest, thus contradicting the expected trend of increasing herbivore impact toward the tropics. Seedling growth was greater in subtropical forest suggesting better conditions for tree growth or that higher invertebrate herbivory depressed seedling growth in the temperate forest. Invertebrate herbivory increased toward temperate latitudes while small mammal herbivory was similar in both sites. We suggest that comparison of single species can be useful to test generalizations about latitudinal patterns and allow disentangling factors controlling herbivory patterns across communities.     

Spatial autocorrelation and the interpretation of patterns of tree seed and seedling predation by rodents in old-fields

Spatial autocorrelation is ubiquitous in ecological systems. However, consideration of this phenomenon in seed and seedling predation studies has been limited, perhaps explaining some of the variation observed between studies and the difficulty in generalizing about the impacts of herbivores on successional processes. I examined the prevalence and patterns of spatial autocorrelation in seed and seedling survival between neighboring tree propagules in old-fields. All experiments were conducted within enclosures in which I manipulated the density of meadow voles ( Microtus pennsylvanicus ). Voles, and competitively inferior white-footed mice ( Peromyscus leucopus ), are the dominant consumers of tree seedlings and seeds in old-fields in the northeastern United States, respectively. An intermediate level of spatial autocorrelation was present in seed and seedling predation trials. Furthermore, the frequency of spatial autocorrelation increased under low-vole density conditions and in study sites with greater heterogeneity in vegetation cover. Survival of neighboring tree propagules was generally positively autocorrelated, suggesting that foraging by mice and voles contributes to a clumped pattern of tree propagule establishment in old-fields. A review of published predation studies found that only 8.3% attempted to test for spatial autocorrelation and that the minimum average spacing used between replicates is well below the average autocorrelation distance of 8.1±6.3 m observed in the current study. Patterns of autocorrelation appear somewhat predictable based on knowledge of the ecology of mice and voles and to provide insights into the factors influencing their foraging behavior and potential impacts on old-field succession.     

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.     

Interactions among dwarf bamboo,litter and consumption by small vertebrates place multiple constraints on the establishment of native tree seedlings in a Japanese agricultural landscape

Aims Faunal assemblage alterations due to anthropogenic impacts have changed herbivore–vegetation relationships in various ecosystems, but the influences of small vertebrates on revegetation processes remain unclear. In a peri-urban agricultural landscape in eastern Japan that lacks large ungulates but supports small generalist herbivores, fewer native seedlings have become established in thickets dominated by native dwarf bamboo,Pleioblastus chino(Franch. et Sav.) Makino. The mechanisms limiting tree seedling establishment are unknown. Our aim here was to evaluate the influence of interactions among the dwarf bamboo, its litter cover and small vertebrate herbivores on the microsite conditions governing the establishment of native tree seedlings from different successional stages in an old field in temperate Japan.     

The effects of elevated CO2 and temperature on the resistance of winter-dormant birch seedlings (Betula pendula) to hares and voles

The effects of elevated CO₂ and temperature on the resource allocation pattern and resistance against mammalian herbivores of silver birch (Betula pendula Roth) were studied. Birch seedlings were grown through two growing seasons in closed‐top chambers exposed to four different treatments: ambient CO₂ and temperature, elevated atmospheric CO₂ (700 ppm) and ambient temperature, elevated temperature (+3°C above ambient) and ambient CO₂, and a combination of elevated CO₂ and temperature. After winter hardening of the seedlings, the growth of the seedlings was measured and the concentration of secondary compounds such as phenolics and papyriferic acid determined. The top parts of the stem were fed to hares, and the basal parts of the same stems were offered to voles. Elevated CO₂ increased the height and basal diameter of the shoots, shoot biomass and total biomass of the seedlings but did not have any effect on secondary chemistry. Elevated temperature increased the height and shoot biomass, but did not have a significant effect on the total biomass of the seedlings. Elevated temperature decreased the concentration of condensed tannins and their precursor, (+)‐catechin, in the top part of the stems, but only the concentration of (+)‐catechin in the basal part of the stems. There were no significant interactive effects between CO₂ and temperature on phenolics in the stems, while the concentration of papyriferic acid showed significant interaction in the top part of the stems. This indicates high accumulation of papyriferic acid in ambient CO₂ under increased temperature. Consequently, elevated temperature increased the resistance of birch against hares, but did not affect the resistance of the basal parts of the same birches to voles. Our results indicate that the predicted climatic change will not necessarily lead to increased browsing damage by the mountain hare and the field vole to silver birch.     

Increasing vole numbers cause more lethal damage to saplings in tree monocultures than in mixed stands

Overall, mammalian herbivores are more harmful in mixed plantations than in monocultures, but the effect of herbivore abundance has not been experimentally tested in this context. It has been proposed that there is a critical threshold density where herbivore pressure spreads from preferred plants to everything edible, leading to non-linear density effects on low-quality plants. We experimentally investigated whether survival of an unpalatable plant is similarly related to herbivore density in both monocultures and mixed stands. This we did by establishing monocultures of unpalatable black alder (Alnus glutinosa) and mixed stands of black alder and five more palatable tree species in enclosures, where Microtus voles were introduced and their abundances monitored.The effect of stand diversity tended to depend on vole abundance. Vole damage of tree saplings did not differ between monocultures and mixed stands, but at higher vole abundances attacks had a stronger effect on sapling survival in the monocultures. Sapling survival showed a significant drop in the monocultures at peak abundance of approximately 300 voles ha^?1. In monocultures herbivores do not have alternatives and therefore are forced to become deadlier consumers.     

Effects of Deer Browsing, Fabric Mats, and Tree Shelters on Quercus rubra Seedlings

We examined the effects of deer browsing, fabric mats, and tree shelters on the growth and survival of Quercus rubra (northern red oak) seedlings planted as part of a reforestation project in southcentral Minnesota. Browsing by white-tailed deer occurred on 68.6% of the seedlings growing without tree shelters, whereas only 3.6% of the trees were browsed by mice or voles and 2.5% were browsed by rabbits. Fabric mats, used to control competition from herbaceous plants for roughly half of the seedlings included in the study, had a detrimental effect overall. Seedlings grown with mats had a greater frequency of deer browsing and a greater chance of dying than seedlings grown without mats. Stem height for seedlings browsed by deer was less overall than for nonbrowsed seedlings, although this pattern varied with use of fabric mats and plot location. The use of plastic tree shelters effectively prevented deer browsing and reduced the mortality rate from 34.6% to 3.2%. Our results indicate that fabric mats should not be used in restoration projects with large deer populations. They also suggest that planting seedlings away from existing forest edges and using seedling protection measures such as tree shelters will increase seedling survival and growth in future restoration projects.     

The confounding effects of logging on tree seedling growth and herbivory in Central Amazon

Light availability is an important modulator of seedling growth and plant–herbivore dynamics. Logging increases light levels in forests, potentially altering herbivore–plant interactions that drive seedling establishment. We conducted a transplant experiment to evaluate how logging and herbivory affect seedling growth and survival in three shade‐tolerant tree species, at paired canopy gap and understory sites in logged forest and an adjacent unlogged area in central Amazonia (Brazil). Seedlings were either left exposed to naturally occurring insect herbivores or protected from insects by a fine netting structure. We measured the herbivore damage and growth rate of seedlings after 18 mo. In general, logged areas received more light than unlogged sites. Growth and herbivory rates were positively influenced by light, and herbivory was also influenced positively by logging. In gaps, increased growth mitigated foliar damage. Logging resulted in a loss of foliar tissue due to increased herbivory. Herbivory rates were higher in the understory of logged sites than in that of unlogged understory sites, but growth was similar in these areas. Thus, the understory of logged areas provided the least favorable sites for shade‐tolerant tree regeneration, due to higher herbivory rates. The effect of logging on biotic interactions can extend beyond the gaps it creates into untouched understory sites. To our knowledge, this is the first time such a pattern has been observed, highlighting the importance of evaluating the impact of logging on biotic interactions.     

Herbivory on Temperate Rainforest Seedlings in Sun and Shade: Resistance,Tolerance and Habitat Distribution

Differential herbivory and/or differential plant resistance or tolerance in sun and shade environments may influence plant distribution along the light gradient. Embothrium coccineum is one of the few light-demanding tree species in the temperate rainforest of southern South America, and seedlings are frequently attacked by insects and snails. Herbivory may contribute to the exclusion of E. coccineum from the shade if 1) herbivory pressure is greater in the shade, which in turn can result from shade plants being less resistant or from habitat preferences of herbivores, and/or 2) consequences of damage are more detrimental in the shade, i.e., shade plants are less tolerant. We tested this in a field study with naturally established seedlings in treefall gaps (sun) and forest understory (shade) in a temperate rainforest of southern Chile. Seedlings growing in the sun sustained nearly 40% more herbivore damage and displayed half of the specific leaf area than those growing in the shade. A palatability test showed that a generalist snail consumed ten times more leaf area when fed on shade leaves compared to sun leaves, i.e., plant resistance was greater in sun-grown seedlings. Herbivore abundance (total biomass) was two-fold greater in treefall gaps compared to the forest understory. Undamaged seedlings survived better and showed a slightly higher growth rate in the sun. Whereas simulated herbivory in the shade decreased seedling survival and growth by 34% and 19%, respectively, damaged and undamaged seedlings showed similar survival and growth in the sun. Leaf tissue lost to herbivores in the shade appears to be too expensive to replace under the limiting light conditions of forest understory. Following evaluations of herbivore abundance and plant resistance and tolerance in contrasting light environments, we have shown how herbivory on a light-demanding tree species may contribute to its exclusion from shade sites. Thus, in the shaded forest understory, where the seedlings of some tree species are close to their physiological tolerance limit, herbivory could play an important role in plant establishment.     

Do carbon-based defences reduce foliar damage? Habitat-related effects on tree seedling performance in a temperate rainforest of Chiloé Island, Chile

Carbon-based secondary compounds (CBSCs), such as phenols or tannins, have been considered as one of the most important and general chemical barriers of woody plants against a diverse array of herbivores. Herbivory has been described as a critical factor affecting the growth and survival of newly established tree seedlings or juveniles then, the presence of secondary metabolites as defences against herbivores should be a primary strategy to reduce foliar damage. We examined whether light-induced changes in leaf phenolic chemistry affected insect herbivory on seedlings of two rainforest tree species, Drimys winteri (Winteraceae) and Gevuina avellana (Proteaceae). Seedlings of both species were planted under closed canopy and in a canopy gap within a large remnant forest patch. Half of the seedlings in each habitat were disinfected with a wide-spectrum systemic insecticide and the other half were used as controls. Seedling growth, survival, and foliar damage (estimated by an herbivory index) due to insect herbivores were monitored over a period of 16 months (December 2001–April 2003). The total leaf content of phenols and condensed tannins were assessed in seedlings from both habitats. As expected, access to light induced a greater production of CBSCs in seedlings of both tree species, but these compounds did not seem to play a significant defensive role, as seedlings grown in gaps suffered greater leaf damage than those planted in forest interior. In addition, in both habitats, seedlings without insecticide treatment suffered a greater foliar damage than those with insecticide, especially 16 months after the beginning of the experiment. Canopy openness and herbivory had positive and negative effects, respectively, on seedling growth and survival in both tree species. In conclusion, despite the higher levels of defence in tree-fall gap, the higher densities of herbivore override this and lead to higher damage levels.     

Mammalian predator scent, vegetation cover and tree seedling predation by meadow voles

Herbivores are thought to respond to the increased risk of attack by predators during foraging activities by concentrating feeding in safe habitats and by reducing feeding in the presence of predators. We tested these hypotheses by comparing tree seedling predation by meadow voles within large outdoor enclosures treated either with scent of large mammalian predators (red fox, bobcat, coyote) or a control scent (vinegar). In addition, we compared the distribution of voles in relation to naturally occurring variation in vegetation cover and the tendency of voles to attack tree seedlings planted in small patches with cover manipulation (intact, reduced or removed cover). Predator scent did not affect the rate or spatial distribution of tree seedling predation by voles, nor did it affect giving up densities (a surrogate of patch quitting harvest rate), survival rates, body size or habitat distribution of voles. In both predator scent and vinegar treatments voles preferred abundant vegetation providing good cover, which was also the site of almost all tree seedling predation. We conclude that large mammalian predator scent does not influence the perception by voles of the general safety of habitat, which is more strongly affected by the presence of cover.