Tree Species Richness Gradient Research Articles

Trophic interactions in soil micro-food webs drive ecosystem multifunctionality along tree species richness.

Rapid biodiversity losses under global climate change threaten forest ecosystem functions. However, our understanding of the patterns and drivers of multiple ecosystem functions across biodiversity gradients remains equivocal. To address this important knowledge gap, we measured simultaneous responses of multiple ecosystem functions (nutrient cycling, soil carbon stocks, organic matter decomposition, plant productivity) to a tree species richness gradient of 1, 4, 8, 16, and 32 species in a young subtropical forest. We found that tree species richness had negligible effects on nutrient cycling, organic matter decomposition, and plant productivity, but soil carbon stocks and ecosystem multifunctionality significantly increased with tree species richness. Linear mixed-effect models showed that soil organisms, particularly arbuscular mycorrhizal fungi (AMF) and soil nematodes, elicited the greatest relative effects on ecosystem multifunctionality. Structural equation models revealed indirect effects of tree species richness on ecosystem multifunctionality mediated by trophic interactions in soil micro-food webs. Specifically, we found a significant negative effect of gram-positive bacteria on soil nematode abundance (a top-down effect), and a significant positive effect of AMF biomass on soil nematode abundance (a bottom-up effect). Overall, our study emphasizes the significance of a multitrophic perspective in elucidating biodiversity-multifunctionality relationships and highlights the conservation of functioning soil micro-food webs to maintain multiple ecosystem functions.

Predation pressure by arthropods, birds, and rodents is interactively shaped by tree species richness, vegetation structure, and season

Tree species richness, forest structure, and seasonal fluctuations between rainy and dry seasons can strongly affect trophic interactions in forest ecosystems, but the inter- and scale dependence of these variables remains unclear. Using artificial caterpillars (~18,000 replicates), we analyzed predation pressure by arthropods, birds, and rodents along a tree species richness gradient across seasons in a subtropical tree diversity experiment (BEF-China). The aim of the study was to test if forest structure, in addition to tree species richness, has an effect on predation pressure and to further specify which structural variables are important in driving predation. We assessed the effects of tree species richness and forest structure at the plot and local neighborhood levels. We also included fine-scale placement covariates, plot size, and topographical covariates of the study site. Forest structure and tree species richness independently and interactively affected predation pressure. The spatial scale was an important determinant for tree species richness and structural effects, extending from within plot scales to the overall heterogeneity of the plots’ surrounding environment. For example, the effect of branch density in the local neighborhood depended on both surrounding tree species richness and plot-level vegetation density. Similarly, visibility-enhancing factors increased attacks by arthropods (lack of branches in close surroundings) and by birds (open area), depending on the surrounding vegetation. A comparison of structural measures showed that predation pressure can be addressed in much greater detail with multiple specific structural features than with overall forest complexity. Seasonal change also affected predation pressure, with foliage being a stronger attractant in spring, but also by presumable topography-driven study plot differences in sun exposure and humidity between rainy and dry seasons. Our study demonstrates that predation pressure is not simply a function of tree species richness or structure but is shaped by the interplay of structural elements, spatial scale, and seasonal dynamics along gradients of tree species richness and forest structure. The structural and seasonal effects are important to take into account when addressing how current and future biodiversity loss may change top‐down control of herbivory and overall ecosystem functioning.

Tree species richness around urban red maples reduces pest density but does not enhance biological control

Urban trees often host greater insect pest abundance than trees in rural forests. This may be due, in part, to differences in tree diversity and canopy cover between these settings. Urban trees are often planted in isolation or monoculture, which favors pest accumulation. Gloomy scale, Melanaspis tenebricosa Comstock, is a pest of urban red maples (Acer rubrum L.) that is abundant where impervious surfaces dominate the local landscape. Increasing tree diversity and canopy cover around urban red maples may reduce gloomy scale abundance by supporting natural enemy communities. We investigated the effect that surrounding tree species richness and tree canopy cover had on gloomy scale abundance, natural enemy abundance, and biological control in red maple trees in Raleigh, NC, USA. We collected scales and natural enemies from red maples that spanned a gradient of tree species richness, canopy cover, and impervious surface values. We also measured gloomy scale parasitism and predation of sentinel prey in red maple canopies. Greater tree species richness and canopy cover were associated with lower gloomy scale density. Red maples in diverse settings also hosted fewer scales per natural enemy. Parasitoids were less common in maples in diverse settings, but generalist predator abundance was unaffected by tree diversity. Finally, tree species richness and canopy cover did not increase biological control of scales or sentinel prey. Our findings suggest that higher tree diversity and greater canopy cover may reduce gloomy scale density, but this is not entirely explained by the effects of natural enemies and biological control.

Nitrogen-fixing tree species modulate species richness effects on soil aggregate-associated organic carbon fractions

Soil aggregation is an important mechanism shaping soil organic carbon (SOC) fractions, but the effects of tree species richness on soil aggregate-associated organic carbon fractions are poorly understood. In this study, a manipulative experiment was conducted to examine whether and how tree species richness (1, 2, 4 and 6) together with or without the presence of nitrogen (N)-fixing tree species affect soil aggregate-associated organic carbon. Soil aggregates were classified into four fractions: large macroaggregates (>2 mm), small macroaggregates (0.25–2 mm), microaggregates (0.053–0.25 mm), and silt and clay (<0.053 mm). And then large and small macroaggregates were further separated into coarse (c-iPOC), fine intra-aggregate particulate organic carbon (f-iPOC) and mineral associated organic carbon (MAOC), and microaggregates were divided into f-iPOC and MAOC. We observed that the effects of tree species richness on soil aggregate-associated organic carbon fractions were depended on the presence or absence of N-fixing tree species. In the presence of N-fixing tree species, organic carbon contents of small macroaggregate, microaggregate and silt and clay fractions were increased across the gradient of tree species richness (+10.4%, +31.3% and + 26.7% from monocultures to 6-species mixture), and the similar responses occurred for c-iPOC within small macroaggregates (+42.9%) and f-iPOC within microaggregates (+50.0%). In the absence of N-fixing tree species, however, only f-iPOC within large macroaggregates (+60.0%) had a positive linear relationship with tree species richness. Structural equation modeling was then used to dissect the mechanism underlying the positive effects of tree species richness on the aggregate-associated organic carbon fractions, indicating either direct or indirect effects by increasing root length density. Our study highlights the important roles of N-fixing tree species in shaping tree species richness effects on soil aggregate-associated organic carbon fractions, and therefore, N-fixing tree species should be preferentially considered in mixed-species plantations to enhance long-term SOC stabilization.

Nitrogen-fixing tree species rather than tree species diversity shape soil nematode communities in subtropical plantations

Positive relationships between tree species diversity and soil communities are reported in many studies. However, the effects of tree species richness and nitrogen (N)-fixing tree species on soil fauna communities are not yet fully understood. Here, we conducted a field experiment to investigate changes in soil nematode communities under a gradient of tree species richness (e.g., 1, 2, 4 and 6 tree species) with N-fixing tree species (absence or presence). We found that N-fixing tree species strongly affected soil nematode trophic groups, and significantly increased the abundance of omnivore-predators and structure index via promoting more trophic links and multi-trophic interactions, leading to a more stable and complex soil food web. In the presence of N-fixing tree species, tree species richness significantly increased the abundance and relative abundance of fungivores and the channel index, with the shift in soil energy channel through increased the C/N ratio of leaf litter. In contrast, tree species richness detrimentally affected the abundance of plant parasites due to the decline of leaf litter N content under the presence of N-fixing tree species. The results revealed that the bottom-up effects of N-fixing tree species on soil nematode community structure were crucial drivers in shaping nematode communities in subtropical plantations. Our study suggests that N-fixing tree species is more important than tree species richness to regulate soil nematode community structure and soil energy channel, and therefore, should be preferentially considered in multispecies forest plantations.

Abiotic and biotic drivers of tree trait effects on soil microbial biomass and soil carbon concentration

AbstractForests are ecosystems critical to understanding the global carbon budget, due to their carbon sequestration potential in both aboveground and belowground compartments, especially in species‐rich forests. Soil carbon sequestration is strongly linked to soil microbial communities, and this link is mediated by the tree community, likely due to modifications of microenvironmental conditions (i.e., biotic conditions, soil properties, and microclimate). We studied soil carbon concentration and the soil microbial biomass of 180 local neighborhoods along a gradient of tree species richness ranging from 1 to 16 tree species per plot in a Chinese subtropical forest experiment (BEF‐China). Tree productivity and different tree functional traits were measured at the neighborhood level. We tested the effects of tree productivity, functional trait identity, and dissimilarity on soil carbon concentrations, and their mediation by the soil microbial biomass and microenvironmental conditions. Our analyses showed a strong positive correlation between soil microbial biomass and soil carbon concentrations. In addition, soil carbon concentration increased with tree productivity and tree root diameter, while it decreased with litterfall C:N content. Moreover, tree productivity and tree functional traits (e.g., fungal root association and litterfall C:N ratio) modulated microenvironmental conditions with substantial consequences for soil microbial biomass. We also showed that soil history and topography should be considered in future experiments and tree plantations, as soil carbon concentrations were higher at sites where historical (i.e., at the beginning of the experiment) carbon concentrations were high, themselves being strongly affected by the topography. Altogether, these results implied that the quantification of the different soil carbon pools is critical for understanding microbial community–soil carbon stock relationships and their dependence on tree diversity and microenvironmental conditions.

Soil Fungal Communities and Enzyme Activities along Local Tree Species Diversity Gradient in Subtropical Evergreen Forest

The majority of studies have found that an increase in tree species diversity can increase the productivity of forest stands thanks to complimentary effects with enhanced resource use efficiency or selection effects; however, it is unclear how tree species diversity affects the soil fungal community and enzyme activities in subtropical evergreen forests. In this study, we used soil high-throughput sequencing to investigate the soil fungal community structure and diversity in the central area of tree clusters in the gradient of tree species richness formed by four possible dominant tree species (Pinus massoniana Lamb., Choerospondias axillaris Roxb., Cyclobalanopsis glauca Thunb. and Lithocarpus glaber Thunb.) in subtropical evergreen broad-leaved forest. The results showed that soil organic carbon content and total nitrogen content were significantly higher in mixed tree clusters, and that soil fungal richness and diversity increased with the increase in tree species diversity (1–3 species). Soil acid phosphatase and urease activity were also enhanced with tree species diversity (p &lt; 0.05). The relative abundance of soil symbiotic fungi (ectomycorrhizal fungi) decreased, while the relative abundance of saprotrophic fungi increased. Redundancy analysis (RDA) revealed that soil acid phosphatase activity was the main factor affecting soil fungal communities and functional guilds, and that soil water content was the main driving force behind fungal trophic modes. In subtropical forests, changes in tree species diversity have altered the soil fungal community structure and trophic modes and functions, accelerating the decomposition of organic matter, increasing nutrient cycling, and perhaps also changing the nutrient absorption of trees.

Tree species richness promotes an early increase of stand structural complexity in young subtropical plantations

Abstract Forest structural complexity has been identified as an important driver for promoting simultaneously biodiversity across trophic levels and multiple ecosystem services. However, we still have a limited understanding of the processes that lead to structural complex stands and how they evolve over time. Using terrestrial laser scanning (TLS), we quantified a three‐dimensional (3D) stand structural complexity index (SSCI) in an experimental plantation with a long gradient of tree species richness (1–24 species). The plantation was established in 2009, and we made use of a multi‐temporal TLS dataset recorded during 2012–2019. We found a positive relationship between tree species richness and structural complexity. This relationship became stronger over time. Ten years after planting, SSCI was on average two‐fold higher in 16‐ and 24‐species mixtures than in monocultures. Furthermore, we demonstrate that tree species richness promotes 3D stand structural complexity indirectly by fostering a high vertical heterogeneity and thus greater spatial complementarity in canopy space. Synthesis and applications. Our findings indicate that tree species richness plays a crucial role in promoting stand structural complexity in young plantations, and this role becomes more important already during early stand development. Thus, afforestation measures would benefit from planting multiple native tree species to initiate structurally complex stands.

Tree species richness and water availability interact to affect soil microbial processes

The objective of this study was to examine the interactive influence of tree diversity and water availability on microbial functioning in surface soil (0–10 cm). The study was conducted at two field experiments (ages 4.2 and 9.0 years, respectively) of the TreeDivNet platform that established plantations with a gradient of tree species richness exposed to high and low water availability treatments. Soil microbial biomass, activity, functional diversity and functional identity were estimated using the MicroResp™ method. Soil organic carbon and several other variables potentially associated with soil processes (soil moisture, soil and air temperature, soil pH, soil C:N ratio, tree biomass) were also measured. Our results indicate that tree species richness significantly increased soil microbial biomass and, to a lesser extent, soil microbial activity in the high water availability treatment, at both trial sites. However, tree species richness effects on microbial processes were not expressed in the low water availability treatment, contrary to the stress-gradient hypothesis. Tree species richness and water treatment interaction impacted microbial biomass and activity through their effects on microbial functional diversity and identity at one site and air temperature at the other site. In conclusion, the mechanisms through which tree diversity creates bottom-up effects on biological soil processes depend on environmental conditions and time following the establishment of tree communities. Future research relating diversity and water availability effects on soil microbial processes should consider functional traits of tree species and a larger range of trophic levels including herbivores and soil fauna to better understand biodiversity-soil ecosystem function relationships in forest ecosystems.

Drivers of understorey biomass: tree species identity is more important than richness in a young forest

Abstract Aims Positive biodiversity–ecosystem functioning relationships have been widely reported, predominately from grassland ecosystems. However, this does not necessarily have to apply accordingly in more complex situations such as in forests across different vertical strata. For instance, overstorey tree species richness has been shown to be associated with a lower understorey productivity. Whether or not tree species richness effects add to understorey productivity by increasing (i.e. due to habitat heterogeneity) or reducing resource availability (i.e. through increasing competition) and whether understorey productivity is indeed being governed more strongly by tree species identity are likely to change over time. Moreover, studies also suggested that richness–productivity relationships change with the environmental context. Using an experimental forest plantation with manipulated tree species richness, this study examined these temporal and environmental dynamics across strata. Methods In the context of the Biodiversity-Ecosystem Functioning project in subtropical China (BEF-China), we made use of understorey biomass samples repeatedly collected over a time period of 3 years along a tree species richness gradient. The effects of tree species richness, tree species identities and time were studied across different environmental treatments for their impact on understorey biomass. Important Findings While we found significant and consistent tree layer identity effects on understorey biomass, no such effect was encountered for tree species richness. Our results also indicate that among structural layers in forests, there might not be a single, generalizable overstorey species richness–understorey productivity relationship, and that the extent as to which overstorey-related environmental factors such as light transmittance contribute to understorey productivity change with time. Overall, we demonstrate that temporal dynamics should be considered when studying relationship among structural layers in forests.

Tree species richness, tree identity and non-native tree proportion affect arboreal spider diversity, abundance and biomass

Sustainable forest management increasingly favors mixtures, but effects on many ecosystem characteristics, such as biocontrol by predators and their biodiversity, are still not well understood. This knowledge gap can be particularly problematic when these mixtures include non-native tree species, such as Douglas fir (Pseudotsuga menziesii (Mirb.) Franco), which are planted because of their high growth rates and climatic suitability but might have negative effects on forest biota. Available studies have mostly focused on epigeic communities, even though tree-associated predator communities can be assumed to make an important contribution to biocontrol that differs significantly from conditions on the forest floor.Here, we analyzed the responses of spiders, as generalist arboreal predators, along a tree species richness gradient in a young tree diversity experiment in Thuringia, Germany, utilizing four of the most economically important deciduous and coniferous tree species in Europe: European beech (Fagus sylvatica L.), Norway spruce (Picea abies (L.) H. Karst.), Douglas fir and sessile oak (Quercus petraea Liebl.), planted in stands of 1–4 tree species. We tested for effects of tree species richness, tree identity and composition (tree species proportion and position of individual trees) on arboreal spider species richness, functional diversity, abundance, biomass and community structure at both the tree and plot level.Tree species richness promoted arboreal spider species richness and functional richness at the plot level and community-weighted mean spider biomass at the tree level. Tree identity and composition had a stronger influence than tree species richness on structuring spider communities, with arboreal spiders having higher numbers per tree on conifers, especially on Douglas fir. However, at the plot level spider species richness, abundance and biomass decreased with an increasing proportion of conifers due to lower species turnover among coniferous trees. The observed effects of tree species richness, tree identity and tree species proportions early in the establishment of forest stands might influence the strength of top-down control of herbivores in critical stages of development of forest stands. Tree species selection and the proportion in which these trees are mixed with other species should be determined with care in order to concomitantly promote biodiversity and biocontrol. In this context, an important finding of our analyses is that potentially negative effects of non-native Douglas fir on arboreal arthropods were not more severe than effects of native conifers that are planted outside of their natural habitats.

Decoupling the Complementarity Effect and the Selection Effect on the Overyielding of Fine Root Production Along a Tree Species Richness Gradient in Subtropical Forests

The mechanism whereby tree species richness and identity affect the production of fine roots (≤ 2 mm) in forests remains controversial. Complementarity effects (via resource partitioning and facilitation, CEs) and selection effects (that is, dominant of species with particular traits, SEs) are the two hypotheses to explain biodiversity effects on ecosystem functions. This study aimed to (1) examine how tree species diversity affects fine root production and (2) disentangle the complementarity effect and the selection effect on the relationship between biodiversity and fine root production. A total of 60 tree clusters with 15 combinations of diversity gradients consisting of 1–4 tree species (Pinus massoniana, Choerospondias axillaris, Cyclobalanopsis glauca and Lithocarpus glaber) were established in subtropical forests. The sequential soil core and ingrowth core methods were used in each cluster to measure fine root biomass and productivity. Fine root production increased with increase in tree species richness. The biodiversity effects on fine root production mostly resulted from CEs. In the nongrowing season, in most cases, the CE on biomass was positive and became stronger as richness increased, but the opposite situation was observed in the growing season. The strong positive and negative effects of the proportions of C. glauca and L. glaber in the tree clusters on fine root biomass, CEs and SEs, suggest the coordinated action of species diversity and identity in modulating biodiversity effects on belowground processes.

Tree species identity determines wood decomposition via microclimatic effects.

Empirical evidence suggests that the rich set of ecosystem functions and nature's contributions to people provided by forests depends on tree diversity. Biodiversity–ecosystem functioning research revealed that not only species richness per se but also other facets of tree diversity, such as tree identity, have to be considered to understand the underlying mechanisms. One important ecosystem function in forests is the decomposition of deadwood that plays a vital role in carbon and nutrient cycling and is assumed to be determined by above‐ and belowground interactions. However, the actual influence of tree diversity on wood decay in forests remains inconclusive. Recent studies suggest an important role of microclimate and advocate a systematical consideration of small‐scale environmental conditions. We studied the influence of tree species richness, tree species identity, and microclimatic conditions on wood decomposition in a 12‐year‐old tree diversity experiment in Germany, containing six native species within a tree species richness gradient. We assessed wood mass loss, soil microbial properties, and soil surface temperature in high temporal resolution. Our study shows a significant influence of tree species identity on all three variables. The presence of Scots pine strongly increased wood mass loss, while the presence of Norway spruce decreased it. This could be attributed to structural differences in the litter layer that were modifying the capability of plots to hold the soil surface temperature at night, consequently leading to enhanced decomposition rates in plots with higher nighttime surface temperatures. Therefore, our study confirmed the critical role of microclimate for wood decomposition in forests and showed that soil microbial properties alone were not sufficient to predict wood decay. We conclude that tree diversity effects on ecosystem functions may include different biodiversity facets, such as tree identity, tree traits, and functional and structural diversity, in influencing the abiotic and biotic soil properties.

Competition, tree age and size drive the productivity of mixed forests of pedunculate oak, beech and red oak

In order to understand the relationship between tree diversity and forest productivity, it is crucial to understand what is going on at the level of individual trees, since each tree responds to the prevailing environmental conditions and interacts with his neighbours. We explored tree interactions in 53 forest plots in northern Belgium (TREEWEB platform) containing all possible combinations of pedunculate oak, beech and red oak. We cored 307 trees and determined the average annual basal area increment for the last 10 years for each sampled tree. We investigated the importance of neighbourhood competition, tree age an size for tree-level growth and then scaled up the species-specific tree growth to stand-level productivity. By exploring how competition, tree age and size varied along the studied gradient of tree species richness, we were able to explain the observed species-specific tree level and stand-level growth along the tree species richness gradient. We found no relationship between tree species richness and forest productivity, due to contrasting growth patterns of the three species along the richness gradient: the growth of pedunculate oak decreased, growth of beech was constant and growth of red oak increased along the tree species richness gradient. Increased heterospecific competition was the main reason for the observed lower growth of pedunculate oak in mixtures, whereas variation in tree age and size along the studied tree species richness gradient were the main drivers for the observed growth patterns of beech and red oak. Species-specific age differences along the tree species richness gradient could be linked with the species composition. We observed a multi-aged stand structure with the light-demanding pedunculate oak being older than the more shade-tolerant beech and red oak, probably a consequence of management interventions needed to enable long-term coexistence of light-demanding and shade-tolerant species. Our study highlights that the management history of mature forests may influence diversity-productivity relationships and should therefore be taken into account during the analysis.

Contrasting patterns of insect herbivory and predation pressure across a tropical rainfall gradient

AbstractOne explanation for the extraordinarily high tree diversity of tropical lowland forests is that it is maintained by specialized natural enemies such as insect herbivores, which cause distance and density dependent mortality. Insect herbivory could also explain the positive correlation between tree species richness and rainfall if herbivory increases with rainfall, is higher on locally abundant versus rare species, and is not limited by predation pressure at wet sites. To test these predictions, insect herbivory and predation pressure on insect herbivores were quantified across a Neotropical rainfall and tree species richness gradient, and herbivory was investigated in relation to local tree abundances. Insect herbivory on leaves (folivory) decreased strongly and significantly with rainfall, while predation pressure was significantly higher at the wetter site. Herbivores were more likely to attack abundant tree species, but herbivore damage levels were not related to tree species abundance. Insect folivores might contribute to local tree species coexistence in our system, but seem unlikely to drive the positive correlation between tree species richness and rainfall. The unexpected and contrasting patterns of herbivory and predation we observed support the need for a multi‐trophic perspective to understand fully the processes contributing to diversity and ecosystem functioning.

Tree species richness attenuates the positive relationship between mutualistic ant-hemipteran interactions and leaf chewer herbivory.

Interactions across trophic levels influence plant diversity effects on ecosystem functions, but the complexity of these interactions remains poorly explored. For example, the interplay between different interactions (e.g. mutualism, predation) might be an important moderator of biodiversity-ecosystem function relationships. We tested for relationships between trophobioses (facultative ant-hemipteran mutualism) and leaf chewer herbivory in a subtropical forest biodiversity experiment. We analysed trophobiosis and herbivory data of more than 10 000 trees along a tree species richness gradient. Against expectations, chewing damage was higher on trees with trophobioses. However, the net positive relationship between trophobioses and overall herbivory depended on tree species richness, being most pronounced at low richness. Our results point to indirect, positive effects of ant-tended sap suckers on leaf chewers, potentially by altering plant defences. Direct antagonistic relationships of trophobiotic ants and leaf-chewing herbivores-frequently reported to drive community-wide effects of trophobioses in other ecosystems-seemed less relevant. However, antagonistic interactions likely contributed to the attenuating effect of tree species richness, because trophobiotic ant and herbivore communities changed from monocultures to species-rich mixtures. Our findings, therefore, suggest that biodiversity loss might lead to complex changes in higher trophic level effects on ecosystem functions, mediated by both trophic and non-trophic interactions.

A role for the sampling effect in invaded ecosystems

Species loss and invasion of exotic species are two components of global biodiversity change that are expected to influence ecosystem functioning. Yet how they interact in natural settings remains unclear. Experiments have revealed two major mechanisms for the observed increase in primary productivity with plant species richness. Plant productivity may rise with species richness due to the increased amount of resources used by more diverse communities (niche complementarity) or through the increased probability of including a highly productive, dominant species in the community (sampling effect). Current evidence suggests that niche complementarity is the most relevant mechanism, whereas the sampling effect would only play a minor and transient role in natural systems. In turn, exotic species can invade by using untapped resources or because they possess a fitness advantage over resident species allowing them to dominate the community. We argue that the sampling effect can be a significant biodiversity mechanism in ecosystems invaded by dominant exotic species, and that the effect can be persistent even after decades of succession. We illustrate this idea by analyzing tree species richness–productivity relationships in a subtropical montane forest (NW Argentina) heavily invaded by Ligustrum lucidum, an evergreen tree from Asia. We found that the forest biomass increased along a natural gradient of tree species richness whether invaded by L. lucidum or not. Consistent with the sampling effect, L. lucidum invasion tripled total tree biomass irrespective of species richness, and monocultures of L. lucidum were more productive than any of the most species‐rich, uninvaded communities. Hence, the sampling effect may not be restricted to randomly assembled, synthetic communities. We emphasize that studying invaded ecosystems may provide novel insights on the mechanisms underlying the effect of biodiversity on ecosystem function.

Foliar fungi of Betula pendula: impact of tree species mixtures and assessment methods

Foliar fungi of silver birch (Betula pendula) in an experimental Finnish forest were investigated across a gradient of tree species richness using molecular high-throughput sequencing and visual macroscopic assessment. We hypothesized that the molecular approach detects more fungal taxa than visual assessment, and that there is a relationship among the most common fungal taxa detected by both techniques. Furthermore, we hypothesized that the fungal community composition, diversity, and distribution patterns are affected by changes in tree diversity. Sequencing revealed greater diversity of fungi on birch leaves than the visual assessment method. One species showed a linear relationship between the methods. Species-specific variation in fungal community composition could be partially explained by tree diversity, though overall fungal diversity was not affected by tree diversity. Analysis of specific fungal taxa indicated tree diversity effects at the local neighbourhood scale, where the proportion of birch among neighbouring trees varied, but not at the plot scale. In conclusion, both methods may be used to determine tree diversity effects on the foliar fungal community. However, high-throughput sequencing provided higher resolution of the fungal community, while the visual macroscopic assessment detected functionally active fungal species.

Positive effects of tree species diversity on litterfall quantity and quality along a secondary successional chronosequence in a subtropical forest

Litterfall, as an important link between aboveground and belowground processes, plays a key role in forest ecosystems. Here, we test for effects of tree species richness on litter production and litter quality in subtropical forest. The study further encompasses a factorial gradient of secondary succession that resulted from human exploitation. Given that a large percentage of subtropical forests are in secondary successional stages, understanding the role of biodiversity on forest re-growth after disturbance appears critical. From January 2009 to December 2014, we monitored forest litterfall in 27 Comparative Study Plots that spanned a gradient of tree species richness (3–20 species) and secondary successional ages (~20 to 120 years) in Gutianshan Natural Nature Reserve, Zhejiang Province, China. The experiment is part of the biodiversity–ecosystem functioning research platform ‘BEF-China’. Tree litterfall was collected in monthly intervals using litter traps. Samples were separated into leaf and non-leaf components. Leaf litter was further sorted into dominant and other species. Community level monthly leaf litter C and N contents were analysed through a full year. General linear mixed-effects models were applied to test for effects of tree species richness and successional age on litter quantity and leaf litter C/N. Litterfall increased with species richness among and within successional age and this effect was consistent across years. Successionally older stands had higher litterfall and this effect was related to increased tree species richness. However, species richness did not change the intra- and inter-annual temporal stability of litterfall. Increasing tree species richness increased leaf litter quality (decreased C/N), while successional age had no effect. Our study indicates that more diverse forest stands produce more leaf litter and that this litter has higher N concentrations, which could promote forest growth through accelerated nutrient re-cycling.

Driving mechanisms of overstorey–understorey diversity relationships in European forests

Forest ecosystem multifunctionality and the provision of ecosystem services benefit from high biodiversity, including from the understorey. Hence, it is crucial to get insight into the mechanisms that influence the composition and diversity in this forest layer. Understorey species have specific environmental requirements which are influenced by the overstorey in a species-specific way. Mixing tree species may create a heterogeneous pattern of environmental conditions similar to the component species’ monocultures, or induce new environmental conditions. We quantified overstorey–understorey taxonomic and functional diversity relationships at different spatial scales in an innovative large-scaled European diversity-oriented research platform that allows the separation of tree species identity and diversity effects. We considered our results in relation to the divergent patterns expected, depending on whether environmental heterogeneity or novel environment creation is the mechanism driving overstorey–understorey diversity relationships.We examined the understorey in mature forests in six European regions. Per region, three to five tree species were selected and 28–43 plots of 30m×30m were established (209 in total), covering a tree species richness gradient from monocultures to three- to five-species mixed plots. Within each plot, we performed vegetation surveys in three vegetation quadrats. We examined the understorey compositional variation among and within regions and the influence of overstorey taxonomic and functional diversity on understorey diversity at different spatial scales.The understorey composition showed clear interregional differences. We found lower compositional differentiation among quadrats with a higher tree species richness level (Finland, Germany and Italy). These quadrats share more tree species and probably have more similar environmental conditions. Neither overstorey taxonomic nor functional diversity affected the stand-level understorey species richness. At the forest-level, adding two-species mixtures to component monocultures clearly increased the understorey species richness, while adding mixtures with a higher tree species richness level to component monocultures had a negligible or negative impact.Overstorey–understorey diversity relationships depend on the spatial scale. At the stand-level, our results lacked support for the environmental heterogeneity hypothesis and revealed the possibility of novel environment creation in tree-by-tree mixed stands. At the forest-level, favouring the mixture of two-species mixed stands and their component monoculture stands offers the best prospects for a higher understorey species richness.