Functional identity drives tree species richness-induced increases in litterfall production and forest floor mass in young tree communities.

Forest floor accumulation is a key process that influences ecosystem carbon cycling. Despite evidence suggesting that tree diversity and soil carbon are positively correlated, most soil carbon studies typically omit the response of the forest floor carbon to tree diversity loss. Here, we evaluated how tree species richness affects forest floor mass and how this effect is mediated by litterfall production and forest floor decay rate in a tree diversity experiment in a subtropical forest. We observed that greater tree species richness leads to higher forest floor accumulation at the soil surface through increasing litterfall production - positively linked to functional trait identity (i.e. community-weighted mean functional trait) rather than functional diversity - and unchanged forest floor decay. Interestingly, structural equation modelling revealed that this lack of overall significant tree species richness effect on forest floor decay rate was due to two indirect and opposite effects cancelling each other out. Indeed, tree species richness increased forest floor decay rate through increasing litterfall production while decreasing forest floor decay rate by increasing litter species richness. Our reports of greater organic matter accumulation in the forest floor in species-rich forests suggest that tree diversity may have long-term and important effect on ecosystem carbon cycling and services.

Resolving the intricate role of climate in litter decomposition.

With approximately 60 Pg of carbon (C) released as CO2 annually, the decomposition of dead organic matter feeds the major terrestrial global CO2 flux to the atmosphere. Macroclimate control over this critical C flux facilitates the parametrization of the C cycle in Earth system models and the understanding of climate change effects on the global C balance. Yet, the long-standing paradigm of climate control was recently challenged by the so far underestimated environmental heterogeneity at local scales, questioning the conceptual framework of thousands of decomposition studies and accuracy of current predictive models. Using three complementary decomposition experiments at a European scale, we showed that macroclimate and litter characteristics largely control plant litter decomposition, reaffirming the role of macroclimate as an integrative decomposition driver through direct environmental control and by influencing co-evolving local plant and decomposer communities. Neglecting this latter indirect effect, commonly used standard litter types overrated micro-environmental control and failed to predict local decomposition of plot-specific litter. Our data help clarify a key question on the regulation of the global C cycle by identifying the relative role of control factors over decomposition and the scales at which they matter and by highlighting sources of confusion in the literature.

Synergistic interactions between detritivores disappear under reduced rainfall.

Understanding the consequences of altered rainfall patterns on litter decomposition is critical to predicting the feedback effect of climate change on atmospheric CO2 concentrations. Although their effect on microbial decomposition has received considerable attention, their effect on litter fragmentation by detritivores, the other dominant decomposition pathway, remains largely unexplored. Particularly, it remains unclear how different detritivore species and their interactions respond to changes in rainfall quantity and frequency. To fill this knowledge gap, we determined the contribution to litter decomposition of two detritivore species (millipede and isopod), separately and in combination, under contrasting rainfall quantity and frequency in a temperate forest. Although halving rainfall quantity and frequency decreased topsoil moisture by 7.8 and 13.1%, respectively, neither millipede- nor isopod-driven decomposition were affected by these changes. In contrast, decomposition driven by both detritivore species in combination was 65.5% higher than expected based on monospecific treatments under high rainfall quantity, but unchanged or even lower under low rainfall quantity. This indicates that although detritivore activity is relatively insensitive to changes in rainfall patterns, large synergistic interactions between detritivore species may disappear under future rainfall patterns. Incorporating interspecific interactions between decomposers thus seems critical to evaluate the sensitivity of decomposition to altered rainfall patterns.

Detritivore conversion of litter into faeces accelerates organic matter turnover.

Litter-feeding soil animals are notoriously neglected in conceptual and mechanistic biogeochemical models. Yet, they may be a dominant factor in decomposition by converting large amounts of plant litter into faeces. Here, we assess how the chemical and physical changes occurring when litter is converted into faeces alter their fate during further decomposition with an experimental test including 36 combinations of phylogenetically distant detritivores and leaf litter of contrasting physicochemical characteristics. We show that, across litter and detritivore species, litter conversion into detritivore faeces enhanced organic matter lability and thereby accelerated carbon cycling. Notably, the positive conversion effect on faeces quality and decomposition increased with decreasing quality and decomposition of intact litter. This general pattern was consistent across detritivores as different as snails and woodlice, and reduced differences in quality and decomposition amongst litter species. Our data show that litter conversion into detritivore faeces has far-reaching consequences for the understanding and modelling of the terrestrial carbon cycle.

Comparison of cellulose vs. plastic cigarette filter decomposition under distinct disposal environments.

It is estimated that 4.5 trillion cigarette butts are discarded annually, making them numerically the most common type of litter on Earth. To accelerate their disappearance after disposal, a new type of cigarette filters made of cellulose, a readily biodegradable compound, has been introduced in the market. Yet, the advantage of these cellulose filters over the conventional plastic ones (cellulose acetate) for decomposition, remains unknown. Here, we compared the decomposition of cellulose and plastic cigarettes filters, either intact or smoked, on the soil surface or within a composting bin over a six-month field decomposition experiment. Within the compost, cellulose filters decomposed faster than plastic filters, but this advantage was strongly reduced when filters had been used for smoking. This indicates that the accumulation of tars and other chemicals during filter use can strongly affect its subsequent decomposition. Strikingly, on the soil surface, we observed no difference in mass loss between cellulose and plastic filters throughout the incubation. Using a first order kinetic model for mass loss of for used filters over the short period of our experiment, we estimated that conventional plastic filters take 7.5-14 years to disappear, in the compost and on the soil surface, respectively. In contrast, we estimated that cellulose filters take 2.3-13 years to disappear, in the compost and on the soil surface, respectively. Our data clearly showed that disposal environments and the use of cellulose filters must be considered when assessing their advantage over plastic filters. In light of our results, we advocate that the shift to cellulose filters should not exempt users from disposing their waste in appropriate collection systems.

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.

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.

Pulse frequency and soil-litter mixing alter the control of cumulative precipitation over litter decomposition.

Macroclimate has traditionally been considered the predominant driver of litter decomposition. However, in drylands, cumulative monthly or annual precipitation typically fails to predict decomposition. In these systems, the windows of opportunity for decomposer activity may rather depend on the precipitation frequency and local factors affecting litter desiccation, such as soil-litter mixing. We used a full-factorial microcosm experiment to disentangle the relative importance of cumulative precipitation, pulse frequency, and soil-litter mixing on litter decomposition. Decomposition, measured as litter carbon loss, saturated with increasing cumulative precipitation when pulses were large and infrequent, suggesting that litter moisture no longer increased and/or microbial activity was no longer limited by water availability above a certain pulse size. More frequent precipitation pulses led to increased decomposition at high levels of cumulative precipitation. Soil-litter mixing consistently increased decomposition, with greatest relative increase (+194%) under the driest conditions. Collectively, our results highlight the need to consider precipitation at finer temporal scale and incorporate soil-litter mixing as key driver of decomposition in drylands.

Tree species diversity affects decomposition through modified micro-environmental conditions across European forests.

Different tree species influence litter decomposition directly through species-specific litter traits, and indirectly through distinct modifications of the local decomposition environment. Whether these indirect effects on decomposition are influenced by tree species diversity is presently not clear. We addressed this question by studying the decomposition of two common substrates, cellulose paper and wood sticks, in a total of 209 forest stands of varying tree species diversity across six major forest types at the scale of Europe. Tree species richness showed a weak but positive correlation with the decomposition of cellulose but not with that of wood. Surprisingly, macroclimate had only a minor effect on cellulose decomposition and no effect on wood decomposition despite the wide range in climatic conditions among sites from Mediterranean to boreal forests. Instead, forest canopy density and stand-specific litter traits affected the decomposition of both substrates, with a particularly clear negative effect of the proportion of evergreen tree litter. Our study suggests that species richness and composition of tree canopies modify decomposition indirectly through changes in microenvironmental conditions. These canopy-induced differences in the local decomposition environment control decomposition to a greater extent than continental-scale differences in macroclimatic conditions.

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.

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.

Two widespread green Neottia species (Orchidaceae) show mycorrhizal preference for Sebacinales in various habitats and ontogenetic stages.

Plant dependence on fungal carbon (mycoheterotrophy) evolved repeatedly. In orchids, it is connected with a mycorrhizal shift from rhizoctonia to ectomycorrhizal fungi and a high natural (13)C and (15)N abundance. Some green relatives of mycoheterotrophic species show identical trends, but most of these remain unstudied, blurring our understanding of evolution to mycoheterotrophy. We analysed mycorrhizal associations and (13)C and (15)N biomass content in two green species, Neottia ovata and N. cordata (tribe Neottieae), from a genus comprising green and nongreen (mycoheterotrophic) species. Our study covered 41 European sites, including different meadow and forest habitats and orchid developmental stages. Fungal ITS barcoding and electron microscopy showed that both Neottia species associated mainly with nonectomycorrhizal Sebacinales Clade B, a group of rhizoctonia symbionts of green orchids, regardless of the habitat or growth stage. Few additional rhizoctonias from Ceratobasidiaceae and Tulasnellaceae, and ectomycorrhizal fungi were detected. Isotope abundances did not detect carbon gain from the ectomycorrhizal fungi, suggesting a usual nutrition of rhizoctonia-associated green orchids. Considering associations of related partially or fully mycoheterotrophic species such as Neottia camtschatea or N. nidus-avis with ectomycorrhizal Sebacinales Clade A, we propose that the genus Neottia displays a mycorrhizal preference for Sebacinales and that the association with nonectomycorrhizal Sebacinales Clade B is likely ancestral. Such a change in preference for mycorrhizal associates differing in ecology within the same fungal taxon is rare among orchids. Moreover, the existence of rhizoctonia-associated Neottia spp. challenges the shift to ectomycorrhizal fungi as an ancestral pre-adaptation to mycoheterotrophy in the whole Neottieae.