Biotic predictors complement models of bat and bird responses to climate and tree diversity in European forests.

Bats and birds are key providers of ecosystem services in forests. How climate and habitat jointly shape their communities is well studied, but whether biotic predictors from other trophic levels may improve bird and bat diversity models is less known, especially across large bioclimatic gradients. Here, we achieved multi-taxa surveys in 209 mature forests replicated in six European countries from Spain to Finland, to investigate the importance of biotic predictors (i.e. the abundance or activity of defoliating insects, spiders, earthworms and wild ungulates) for bat and bird taxonomic and functional diversity. We found that nine out of 12 bird and bat diversity metrics were best explained when biotic factors were added to models including climate and habitat variables, with a mean gain in explained variance of 38% for birds and 15% for bats. Tree functional diversity was the most important habitat predictor for birds, while bats responded more to understorey structure. The best biotic predictors for birds were spider abundance and defoliating insect activity, while only bat functional evenness responded positively to insect herbivory. Accounting for potential biotic interactions between bats, birds and other taxa of lower trophic levels will help to understand how environmental changes along large biogeographical gradients affect higher-level predator diversity in forest ecosystems.

The functional role of temperate forest understorey vegetation in a changing world.

Temperate forests cover 16% of the global forest area. Within these forests, the understorey is an important biodiversity reservoir that can influence ecosystem processes and functions in multiple ways. However, we still lack a thorough understanding of the relative importance of the understorey for temperate forest functioning. As a result, understoreys are often ignored during assessments of forest functioning and changes thereof under global change. We here compiled studies that quantify the relative importance of the understorey for temperate forest functioning, focussing on litter production, nutrient cycling, evapotranspiration, tree regeneration, pollination and pathogen dynamics. We describe the mechanisms driving understorey functioning and develop a conceptual framework synthesizing possible effects of multiple global change drivers on understorey-mediated forest ecosystem functioning. Our review illustrates that the understorey's contribution to temperate forest functioning is significant but varies depending on the ecosystem function and the environmental context, and more importantly, the characteristics of the overstorey. To predict changes in understorey functioning and its relative importance for temperate forest functioning under global change, we argue that a simultaneous investigation of both overstorey and understorey functional responses to global change will be crucial. Our review shows that such studies are still very scarce, only available for a limited set of ecosystem functions and limited to quantification, providing little data to forecast functional responses to global change.

Biotic homogenization can decrease landscape-scale forest multifunctionality.

Many experiments have shown that local biodiversity loss impairs the ability of ecosystems to maintain multiple ecosystem functions at high levels (multifunctionality). In contrast, the role of biodiversity in driving ecosystem multifunctionality at landscape scales remains unresolved. We used a comprehensive pan-European dataset, including 16 ecosystem functions measured in 209 forest plots across six European countries, and performed simulations to investigate how local plot-scale richness of tree species (α-diversity) and their turnover between plots (ß-diversity) are related to landscape-scale multifunctionality. After accounting for variation in environmental conditions, we found that relationships between α-diversity and landscape-scale multifunctionality varied from positive to negative depending on the multifunctionality metric used. In contrast, when significant, relationships between ß-diversity and landscape-scale multifunctionality were always positive, because a high spatial turnover in species composition was closely related to a high spatial turnover in functions that were supported at high levels. Our findings have major implications for forest management and indicate that biotic homogenization can have previously unrecognized and negative consequences for large-scale ecosystem multifunctionality.

Continental mapping of forest ecosystem functions reveals a high but unrealised potential for forest multifunctionality.

Humans require multiple services from ecosystems, but it is largely unknown whether trade-offs between ecosystem functions prevent the realisation of high ecosystem multifunctionality across spatial scales. Here, we combined a comprehensive dataset (28 ecosystem functions measured on 209 forest plots) with a forest inventory dataset (105,316 plots) to extrapolate and map relationships between various ecosystem multifunctionality measures across Europe. These multifunctionality measures reflected different management objectives, related to timber production, climate regulation and biodiversity conservation/recreation. We found that trade-offs among them were rare across Europe, at both local and continental scales. This suggests a high potential for 'win-win' forest management strategies, where overall multifunctionality is maximised. However, across sites, multifunctionality was on average 45.8-49.8% below maximum levels and not necessarily highest in protected areas. Therefore, using one of the most comprehensive assessments so far, our study suggests a high but largely unrealised potential for management to promote multifunctional forests.

Monitoring the Impact of Hedgerows and Grass Strips on the Performance of Multiple Ecosystem Service Indicators.

The importance of semi-natural vegetation elements in the agricultural landscape is increasingly recognized because they have the potential to enhance multiple ecosystem service delivery and biodiversity. However, there is great variability in the observed effects within and between studies. Also, little is known about the simultaneous delivery of multiple ecosystem services and biodiversity because most studies focus on monitoring one service at a time and in conditions specifically suited to observe this one service. In this study, the results are presented of 1 year of monitoring of a set of parcel-level and simplistic ecosystem service and biodiversity indicators on parcels with grass strips or hedgerows. In the grass strips, an increase in soil organic carbon stock, a decrease in soil mineral nitrogen content, a different carabid species composition and a higher spider activity density were found, compared to the adjacent arable parcel. These results indicate a contribution of grass strips to climate regulation, the regulation of water quality, an increase of beta diversity and potential for pest control. Next to hedgerows, crop yield was reduced and winter wheat thousand kernel weight, soil organic carbon stock and spider activity density were increased. These indicators show an effect of the hedgerow on food production, climate regulation and potential for pest control. The study concludes that both grass strips and hedgerows have the potential to increase multiple ecosystem service delivery, but that an increase of every service is not assured and that multifunctionality is affected by management choices. Also, an improved experimental setup in order to enhance ecosystem service monitoring is suggested.

Biodiversity and ecosystem functioning relations in European forests depend on environmental context.

The importance of biodiversity in supporting ecosystem functioning is generally well accepted. However, most evidence comes from small-scale studies, and scaling-up patterns of biodiversity-ecosystem functioning (B-EF) remains challenging, in part because the importance of environmental factors in shaping B-EF relations is poorly understood. Using a forest research platform in which 26 ecosystem functions were measured along gradients of tree species richness in six regions across Europe, we investigated the extent and the potential drivers of context dependency of B-EF relations. Despite considerable variation in species richness effects across the continent, we found a tendency for stronger B-EF relations in drier climates as well as in areas with longer growing seasons and more functionally diverse tree species. The importance of water availability in driving context dependency suggests that as water limitation increases under climate change, biodiversity may become even more important to support high levels of functioning in European forests.

Bat and bird diversity along independent gradients of latitude and tree composition in European forests.

Species assemblages are shaped by local and continental-scale processes that are seldom investigated together, due to the lack of surveys along independent gradients of latitude and habitat types. Our study investigated changes in the effects of forest composition and structure on bat and bird diversity across Europe. We compared the taxonomic and functional diversity of bat and bird assemblages in 209 mature forest plots spread along gradients of forest composition and vertical structure, replicated in 6 regions spanning from the Mediterranean to the boreal biomes. Species richness and functional evenness of both bat and bird communities were affected by the interactions between latitude and forest composition and structure. Bat and bird species richness increased with broadleaved tree cover in temperate and especially in boreal regions but not in the Mediterranean where they increased with conifer abundance. Bat species richness was lower in forests with smaller trees and denser understorey only in northern regions. Bird species richness was not affected by forest structure. Bird functional evenness increased in younger and denser forests. Bat functional evenness was also influenced by interactions between latitude and understorey structure, increasing in temperate forests but decreasing in the Mediterranean. Covariation between bat and bird abundances also shifted across Europe, from negative in southern forests to positive in northern forests. Our results suggest that community assembly processes in bats and birds of European forests are predominantly driven by abundance and accessibility of feeding resources, i.e., insect prey, and their changes across both forest types and latitudes.

Diversifying forest communities may change Lyme disease risk: extra dimension to the dilution effect in Europe.

Lyme disease is caused by bacteria of the Borrelia burgdorferi genospecies complex and transmitted by Ixodid ticks. In North America only one pathogenic genospecies occurs, in Europe there are several. According to the dilution effect hypothesis (DEH), formulated in North America, nymphal infection prevalence (NIP) decreases with increasing host diversity since host species differ in transmission potential. We analysed Borrelia infection in nymphs from 94 forest stands in Belgium, which are part of a diversification gradient with a supposedly related increasing host diversity: from pine stands without to oak stands with a shrub layer. We expected changing tree species and forest structure to increase host diversity and decrease NIP. In contrast with the DEH, NIP did not differ between different forest types. Genospecies diversity however, and presumably also host diversity, was higher in oak than in pine stands. Infected nymphs tended to harbour Borrelia afzelii infection more often in pine stands while Borrelia garinii and Borrelia burgdorferi ss. infection appeared to be more prevalent in oak stands. This has important health consequences, since the latter two cause more severe disease manifestations. We show that the DEH must be nuanced for Europe and should consider the response of multiple pathogenic genospecies.

Importance of overstorey attributes for understorey litter production and nutrient cycling in European forests.

BACKGROUND: In contrast with the negligible contribution of the forest understorey to the total aboveground phytobiomass of a forest, its share in annual litter production and nutrient cycling may be more important. Whether and how this functional role of the understorey differs across forest types and depends upon overstorey characteristics remains to be investigated. METHODS: We sampled 209 plots of the FunDivEUROPE Exploratory Platform, a network of study plots covering local gradients of tree diversity spread over six contrasting forest types in Europe. To estimate the relative contribution of the understorey to carbon and nutrient cycling, we sampled non-lignified aboveground understorey biomass and overstorey leaf litterfall in all plots. Understorey samples were analysed for C, N and P concentrations, overstorey leaf litterfall for C and N concentrations. We additionally quantified a set of overstorey attributes, including species richness, proportion of evergreen species, light availability (representing crown density) and litter quality, and investigated whether they drive the understorey's contribution to carbon and nutrient cycling. RESULTS AND CONCLUSIONS: Overstorey litter production and nutrient stocks in litterfall clearly exceeded the contribution of the understorey for all forest types, and the share of the understorey was higher in forests at the extremes of the climatic gradient. In most of the investigated forest types, it was mainly light availability that determined the contribution of the understorey to yearly carbon and nutrient cycling. Overstorey species richness did not affect the contribution of the understorey to carbon and nutrient cycling in any of the investigated forest types.

Year-to-year variation in the density of Ixodes ricinus ticks and the prevalence of the rodent-associated human pathogens Borrelia afzelii and B. miyamotoi in different forest types.

The human pathogens Borrelia afzelii, which causes Lyme borreliosis and B. miyamotoi, which causes relapsing fever, both circulate between Ixodes ricinus ticks and rodents. The spatiotemporal dynamics in the prevalence of these pathogens have not yet been fully elucidated, but probably depend on the spatiotemporal population dynamics of small rodents. We aimed to evaluate the effect of different forest types on the density of infected nymphs in different years and to obtain more knowledge about the spatial and temporal patterns of ticks and tick-borne pathogens. We analysed unfed nymphal ticks from 22 stands of four different forest types in Belgium in 2009, 2010, 2013 and 2014 and found that the density of nymphs in general and the density of nymphs infected with B. afzelii and B. miyamotoi varied yearly, but without temporal variation in the infection prevalence. The yearly variation in density of infected nymphs in our study thus seems to be caused most by the variation in the density of nymphs, which makes it a good predictor of disease risk. The risk for rodent-associated tick-borne diseases also varied between forest types. We stress the need to elucidate the contribution of the host community composition to tick-borne disease risk.

Low probability of a dilution effect for Lyme borreliosis in Belgian forests.

An increasing number of studies have investigated the consequences of biodiversity loss for the occurrence of vector-borne diseases such as Lyme borreliosis, the most common tick-borne disease in the northern hemisphere. As host species differ in their ability to transmit the Lyme borreliosis bacteria Borrelia burgdorferi s.l. to ticks, increased host diversity can decrease disease prevalence by increasing the proportion of dilution hosts, host species that transmit pathogens less efficiently. Previous research shows that Lyme borreliosis risk differs between forest types and suggests that a higher diversity of host species might dilute the contribution of small rodents to infect ticks with B. afzelii, a common Borrelia genospecies. However, empirical evidence for a dilution effect in Europe is largely lacking. We tested the dilution effect hypothesis in 19 Belgian forest stands of different forest types along a diversity gradient. We used empirical data and a Bayesian belief network to investigate the impact of the proportion of dilution hosts on the density of ticks infected with B. afzelii, and identified the key drivers determining the density of infected ticks, which is a measure of human infection risk. Densities of ticks and B. afzelii infection prevalence differed between forest types, but the model indicated that the density of infected ticks is hardly affected by dilution. The most important variables explaining variability in disease risk were related to the density of ticks. Combining empirical data with a model-based approach supported decision making to reduce tick-borne disease risk. We found a low probability of a dilution effect for Lyme borreliosis in a north-western European context. We emphasize that under these circumstances, Lyme borreliosis prevention should rather aim at reducing tick-human contact rate instead of attempting to increase the proportion of dilution hosts.

Jack-of-all-trades effects drive biodiversity-ecosystem multifunctionality relationships in European forests.

There is considerable evidence that biodiversity promotes multiple ecosystem functions (multifunctionality), thus ensuring the delivery of ecosystem services important for human well-being. However, the mechanisms underlying this relationship are poorly understood, especially in natural ecosystems. We develop a novel approach to partition biodiversity effects on multifunctionality into three mechanisms and apply this to European forest data. We show that throughout Europe, tree diversity is positively related with multifunctionality when moderate levels of functioning are required, but negatively when very high function levels are desired. For two well-known mechanisms, 'complementarity' and 'selection', we detect only minor effects on multifunctionality. Instead a third, so far overlooked mechanism, the 'jack-of-all-trades' effect, caused by the averaging of individual species effects on function, drives observed patterns. Simulations demonstrate that jack-of-all-trades effects occur whenever species effects on different functions are not perfectly correlated, meaning they may contribute to diversity-multifunctionality relationships in many of the world's ecosystems.