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.

Phenotypic traits and their associated trade-offs have been shown to have globally consistent effects on individual plant physiological functions 1-3 , but how these effects scale up to influence competition, a key driver of community assembly in terrestrial vegetation, has remained unclear 4 . Here we use growth data from more than 3 million trees in over 140,000 plots across the world to show how three key functional traits-wood density, specific leaf area and maximum height-consistently influence competitive interactions. Fast maximum growth of a species was correlated negatively with its wood density in all biomes, and positively with its specific leaf area in most biomes. Low wood density was also correlated with a low ability to tolerate competition and a low competitive effect on neighbours, while high specific leaf area was correlated with a low competitive effect. Thus, traits generate trade-offs between performance with competition versus performance without competition, a fundamental ingredient in the classical hypothesis that the coexistence of plant species is enabled via differentiation in their successional strategies 5 . Competition within species was stronger than between species, but an increase in trait dissimilarity between species had little influence in weakening competition. No benefit of dissimilarity was detected for specific leaf area or wood density, and only a weak benefit for maximum height. Our traitbased approach to modelling competition makes generalization possible across the forest ecosystems of the world and their highly diverse species composition.Phenotypic traits are considered fundamental drivers of community assembly and thus species diversity 1,6 . The effects of traits on individual plant physiologies and functions are increasingly understood, and have been shown to be underpinned by well-known and globally consistent trade-offs 1-3 . For instance, traits such as wood density and specific leaf area capture trade-offs between the construction cost and longevity or strength of wood and leaf tissues 2,3 . By contrast, we still have a limited understanding of how such trait-based trade-offs translate into competitive interactions between species, particularly for long-lived organisms such as trees. Competition is a key filter through which ecological and evolutionary success is determined 4 . A long-standing hypothesis is that the intensity of competition decreases as two species diverge in trait values 7 (trait dissimilarity). The few studies [8][9][10][11][12][13] that have explored links between traits and competition have shown that linkages were more complex than this, as particular trait values may also confer competitive advantage independently from trait dissimilarity 9,13,14 . This distinction is fundamental for species coexistence and the local mixture of traits. If neighbourhood competition is driven mainly by trait dissimilarity, this will favour a wide spread of trait values at a local scale. By contrast, if neighbourhood interactions are mainly driven by the c...

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One of the most fundamental questions in ecology is how many species inhabit the Earth. However, due to massive logistical and financial challenges and taxonomic difficulties connected to the species concept definition, the global numbers of species, including those of important and well-studied life forms such as trees, still remain largely unknown. Here, based on global ground-sourced data, we estimate the total tree species richness at global, continental, and biome levels. Our results indicate that there are ∼73,000 tree species globally, among which ∼9,000 tree species are yet to be discovered. Roughly 40% of undiscovered tree species are in South America. Moreover, almost one-third of all tree species to be discovered may be rare, with very low populations and limited spatial distribution (likely in remote tropical lowlands and mountains). These findings highlight the vulnerability of global forest biodiversity to anthropogenic changes in land use and climate, which disproportionately threaten rare species and thus, global tree richness.

Hair lichens are strongly influenced by forest structure at local scales, but their broad-scale distributions are less understood. We compared the occurrence and length of Alectoria sarmentosa (Ach.) Ach., Bryoria spp., and Usnea spp. in the lower canopy of > 5000 Picea abies (L.) Karst. trees within the National Forest Inventory across all productive forest in Sweden. We used logistic regression to analyse how climate, nitrogen deposition, and forest variables influence lichen occurrence. Distributions overlapped, but the distribution of Bryoria was more northern and that of Usnea was more southern, with Alectoria's distribution being intermediate. Lichen length increased towards northern regions, indicating better conditions for biomass accumulation. Logistic regression models had the highest pseudo R 2 value for Bryoria, followed by Alectoria. Temperature and nitrogen deposition had higher explanatory power than precipitation and forest variables. Multiple logistic regressions suggest that lichen genera respond differently to increases in several variables. Warming decreased the odds for Bryoria occurrence at all temperatures. Corresponding odds for Alectoria and Usnea decreased in warmer climates, but in colder climates, they increased. Nitrogen addition decreased the odds for Alectoria and Usnea occurrence under high deposition, but under low deposition, the odds increased. Our analyses suggest major shifts in the broad-scale distribution of hair lichens with changes in climate, nitrogen deposition, and forest management.Key words: climate change, epiphytic lichens, forest structure, nitrogen deposition, temperature.