Evy Ampoorter

Biotic homogenization can decrease landscape-scale forest multifunctionality

Significance Numerous studies have demonstrated the importance of biodiversity in maintaining multiple ecosystem functions and services (multifunctionality) at local spatial scales, but it is unknown whether similar relationships are found at larger spatial scales in real-world landscapes. Here, we show, for the first time to our knowledge, that biodiversity can also be important for multifunctionality at larger spatial scales in European forest landscapes. Both high local (α-) diversity and a high turnover in species composition between locations (high β-diversity) were found to be potentially important drivers of ecosystem multifunctionality. Our study provides evidence that it is important to conserve the landscape-scale biodiversity that is being eroded by biotic homogenization if ecosystem multifunctionality is to be maintained. 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 (β-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 β-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.

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.

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.

Diversity of secondary woody species in relation to species richness and cover of dominant trees in thermophilous deciduous forests

ABSTRACT One of the prominent compositional features of thermophilous deciduous forests of southern Europe is their richness in secondary woody species (AWS). To date, no studies have focused on the diversity and contribution to the ecosystem functions and socio-economic benefits provided by these species. Here, we first characterized species richness and diversity for three main functional groups of AWS, sporadic trees, shrubs and vines, in 53 representative stands of central Italy. Secondly, we explored variations in AWS diversity and composition along a four-level gradient of species richness of dominant trees under comparable site conditions. The 40 AWS taxa recorded showed a remarkable taxonomic singularity at both the genus and the family rank. Dominant tree species richness had no significant effect on AWS species richness and composition, while Shannon diversity of AWS, especially shrubs, was associated with cover of the dominant trees. Each AWS possessed, on average, ca. five ability traits over a total of 13 considered that contributed to important ecosystem functions or expected socio-economic benefits such as wood and food production, conservation of genetic resources (wild crop relatives) and ornamentals. This highlights the key role of AWS, and in particular of sporadic trees, for enhancing multifunctionality of thermophilous deciduous 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.

Does neighbourhood tree diversity affect the crown arthropod community in saplings

Mixed forest with multiple tree species is expected to create heterogeneous habitat and diverse niches for the canopy arthropod community. We assessed arthropod abundance, order richness, and community composition in the crowns of saplings of nine temperate tree species in two plantations of a recently established tree diversity experiment in Belgium, and looked for relationships with the diversity and structure of the sapling’s local neighbourhood. The crown arthropod community differed between the two study sites, both in terms of abundances and composition. More arthropods were found in the post-agricultural site; the arthropod community was more complex in the formerly forested site. The tree species identity of a sapling, its apparency, and the phylogenetic diversity of its local neighbourhood all affected the crown arthropod community. Our study suggests that mixing phylogenetically distant tree species creates niches for a complex crown arthropod community.

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.

Functional Composition of Tree Communities Changed Topsoil Properties in an Old Experimental Tropical Plantation

Forest biogeochemistry is strongly determined by the interaction between the tree community and the topsoil. Functional strategies of tree species are coupled to specific chemical leaf traits, and thus also to litter composition, which affects mineral soil characteristics. The limited understanding on this interaction is mainly based on shorter-term common garden experiments in temperate forest, and needs to be extended to other forest types and climates if we want to understand the universality of this linkage. In particular, for highly diverse tropical forests, our understanding of this interaction remains limited. Using an old experimental plantation within the central Congo basin, we examined the relationship between leaf and litter chemical composition and topsoil properties. Canopy, litter and topsoil characteristics were measured and we determined how the community-level leaf and litter chemical composition altered the topsoil carbon, major plant nutrients and exchangeable cation concentration, acidity and pH over the last eight decades. We found that functional composition strongly affected topsoil pH. In turn, topsoil pH strongly determined the soil total carbon and available phosphorus, total nitrogen and exchangeable potassium. Our results indicate that, as observed in temperate common garden experiments, trees alter chemical topsoil properties primarily through soil acidification, differently induced by functional composition of the tree community. The strong link between this community-level composition and topsoil characteristics, on a highly representative soil type for the tropics, improves our understanding of tropical forests biogeochemistry.

Functional identity explains carbon sequestration in a 77-year-old experimental tropical plantation

Planting forests is an important practice for climate change mitigation, especially in the tropics where the carbon (C) sequestration potential is high. Successful implementation of this mitigation practice requires knowledge of the role of species identity and diversity on carbon accrual of plantations. Despite this need, solid data on the long-term development of forest plantations are still very scarce. Monospecific and two species mixture plots of a 77-year-old tree diversity experiment in Yangambi in the Congo basin were fully inventoried. We calculated above-ground C stocks using allometric equations, and soil C stocks by analyzing soil samples at multiple depths. Linear mixed effects models were used to analyze the effect of taxonomic and functional identity and diversity on the aboveground and soil carbon stocks. A high variability in aboveground C stocks across tree species combinations was observed. Apart from a species identity effect, the proportion of planted species in the total stand basal area (BA(pl)) and effective species richness were identified as compositional parameters with a significant effect on the aboveground carbon (AGC), with BApl being more important. Both AGC and BA(pl) were coupled to the functional identity of the planted species; the planting of short-lived pioneers led to low AGC. We found no clear benefits, but also no drawbacks, for AGC of two species mixture plots over monospecific plots or including nitrogen fixing species in the plantation scheme. However, the latter was the only compositional parameter with a significant positive effect on the soil carbon stock up to 1 m depth. We conclude that the different plantation configurations gave rise to a wide range in carbon stocks. This was predominantly caused by large differences in AGC sequestration over the past 77 years. Altogether, short-lived pioneer species had a low BApl resulting in low carbon sequestration, while partial shade tolerant species achieved the highest AGC stocks. Tolerating spontaneous ingrowth during the plantation development can further increase the AGC stock, given that the appropriate functional type is planted.

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.