Importance of large and small mammalian herbivores for the plant community structure in the forest tundra ecotone

Both theoretical arguments and empirical evidence suggests that herbivory in general and mammalian winter herbivory in particular is important in arctic-alpine ecosystems. Although knowledge of the effect of herbivores on specific plants and communities is quite extensive, little is known about the relative impact of large and small vertebrate herbivores and how it might vary among different habitats. To address this key issue, we established exclosures with two different mesh sizes in forest and nearby tundra at three different sites in four contrasting locations in the forest-tundra ecotone in northernmost Sweden and Norway. Plant community composition was recorded annually in three permanent plots within each exclosure and an unfenced control. Local densities of vertebrate herbivores were estimated in spring and autumn from 1998 to 2002. Reindeer (Rangifer tarandus) were the most abundant large vertebrate while Norwegian lemmings (Lemmus lemmus) and grey-sided voles (Clethrionomys rufocanus) were the most common small vertebrates. The study reveals that voles and lemmings have larger effects on the vegetation than reindeer in both habitats in all four locations, even though densities of reindeer differ between locations and only two locations experienced lemming peaks during the period of the experiment. The relative abundance of five of the fifteen most common species was significantly influenced by voles and lemmings whereas only a single species was significantly influenced by reindeer. Different analyses give contrasting results on the importance of herbivory in forest versus open heathlands. A principal component analyses revealed that herbivory influenced the vegetation more in open heathlands than in forests. However, an importance index of herbivores did not differ between forest and open heathlands. Moreover, none of the plant species responded differently in the two habitats, when herbivores were removed. Our results suggest that intense and localised selective foraging by small mammals may have a more marked effect on vegetation than transient feeding by reindeer.