Forests are of major importance to human society, contributing several crucial ecosystem services. Biodiversity is suggested to positively influence multiple services but evidence from natural systems at scales relevant to management is scarce. Here, across a scale of 400,000 km2, we report that tree species richness in production forests shows positive to positively hump-shaped relationships with multiple ecosystem services. These include production of tree biomass, soil carbon storage, berry production and game production potential. For example, biomass production was approximately 50% greater with five than with one tree species. In addition, we show positive relationships between tree species richness and proxies for other biodiversity components. Importantly, no single tree species was able to promote all services, and some services were negatively correlated to each other. Management of production forests will therefore benefit from considering multiple tree species to sustain the full range of benefits that the society obtains from forests.

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Summary1 Transitions between major vegetation types, such as the tree line, are useful systems for monitoring the response of vegetation to climate change. Tree lines have, however, shown equivocal responses to such change. 2 Tree lines are considered to be primarily thermally controlled, although recent work has highlighted the importance of biotic factors. Dispersal limitation and the invasibility of the tundra matrix have been implicated and here we propose herbivory as an additional control at some tree lines. 3 We propose a conceptual model in which differing relative impacts of foliage consumption, availability of establishment sites, trampling, dispersal and seed predation can lead to very different tree-line responses. 4 The presence of large numbers of small trees above the current tree line at a site in northern Sweden that experiences limited reindeer ( Rangifer tarandus ) herbivory suggests range expansion. Other locations in the same region with higher reindeer populations have considerably fewer small trees, suggesting that range expansion is occurring much more slowly, if at all. 5 The use of tree lines as indicators of climate change is confounded by the activity of herbivores, which may either strengthen or nullify the impacts of a changed climate. Similar arguments are likely to be applicable to other ecotones.

Background: The reindeer (or caribou, Rangifer tarandus L.) has a natural range extending over much of Eurasia's and North America's arctic, alpine and boreal zones, yet its impact on vegetation is still unclear. This lack of a common understanding hampers both the management of wild and semi-domesticated reindeer populations and the preservation of biodiversity. To achieve a common platform, we have undertaken a systematic review of published studies that compare vegetation at sites with different reindeer densities. Besides biodiversity, we focused on effects on major plant growth forms. Methods: Searches for literature were made using online publication databases, search engines, specialist websites and bibliographies of literature reviews. Search terms were developed in English, Finnish, Norwegian, Russian and Swedish. Identified articles were screened for relevance based on titles, abstracts and full text using inclusion criteria set out in an a priori protocol. Relevant articles were then subject to critical appraisal of susceptibility to bias. Data on outcomes such as abundance, biomass, cover and species richness of vegetation were extracted together with metadata on site properties and other potential effect modifiers.Results: Our searches identified more than 6,000 articles. After screening for relevance, 100 of them remained. Critical appraisal excluded 60 articles, leaving 40 articles with 41 independent studies. Almost two thirds of these studies had been conducted in Fennoscandia. Meta-analysis could be made of data from 31 of the studies. Overall, effects of reindeer on species richness of vascular plants depended on temperature, ranging from negative at low temperature to positive at high temperature. Effects on forbs, graminoids, woody species, and bryophytes were weak or non-significant, whereas the effect on lichens was negative. However, many individual studies showed clear positive or negative effects, but the available information was insufficient to explain this context dependence. Conclusions:We see two pressing matters emerging from our study. First, there is a lack of research with which to build a circumpolar understanding of grazing effects, which calls for more studies using a common protocol to quantify reindeer impacts. Secondly, the highly context-dependent outcomes suggest that research and management have to consider local conditions. For instance, predictions of what a management decision would mean for the effects of reindeer on vegetation will have to take the variation of vegetation types and dominant growth forms, productivity, and grazing history into account. Policy and management have to go hand-in-hand with research in individual cases if the dynamics between plants, animals, and humans are to be sufficiently understood.

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