Community diversity outweighs effect of warming on plant colonization

Abiotic environmental change, local species extinctions and colonization of new species often co‐occur. Whether species colonization is driven by changes in abiotic conditions or reduced biotic resistance will affect community functional composition and ecosystem management. We use a grassland experiment to disentangle effects of climate warming and community diversity on plant species colonization. Community diversity had dramatic impacts on the biomass, richness and traits of plant colonists. Three times as many species colonized the monocultures than the high diversity 17 species communities (~30 vs. 10 species), and colonists collectively produced 10 times as much biomass in the monocultures than the high diversity communities (~30 vs. 3 g/m²). Colonists with resource‐acquisitive strategies (high specific leaf area, light seeds, short heights) accrued more biomass in low diversity communities, whereas species with conservative strategies accrued most biomass in high diversity communities. Communities with higher biomass of resident C4 grasses were more resistant to colonization by legume, nonlegume forb and C3 grass colonists, but not by C4 grass colonists. Compared with effects of diversity, 6 years of 3°C‐above‐ambient temperatures had little impact on plant colonization. Warmed subplots had ~3 fewer colonist species than ambient subplots and selected for heavier seeded colonists. They also showed diversity‐dependent changes in biomass of C3 grass colonists, which decreased under low diversity and increased under high diversity. Our findings suggest that species colonization is more strongly affected by biotic resistance from residents than 3°C of climate warming. If these results were extended to invasive species management, preserving community diversity should help limit plant invasion, even under climate warming.     

Leaf litter species identity alters the structure of pond communities

The input of leaf litter resources is a major driver of ecosystem processes in terrestrial and freshwater habitats. Although variation exists in the quantity and composition of litter inputs due to natural and anthropogenic causes, few studies have examined how such variation influences the structure and composition of aquatic food webs. Using outdoor mesocosms, we examined the bottom–up effects of 10 chemically distinct tree litter species on microbial, algal, invertebrate and vertebrate fauna found in temperate ponds. We hypothesized that individual litter species, which differ in their traits, would differentially and predictably affect abiotic and biotic elements of pond communities. We further hypothesized that the presence of leaf litter, regardless of species, would elevate resource supply and increase the biomass of community members. Finally, we hypothesized that a mixture of litter species would have non‐additive effects on community responses. We followed the system for > 4 months and measured > 30 abiotic and biotic responses related to primary and secondary production. The different species of leaf litter had major effects on abiotic and biotic responses, including phytoplankton, periphyton, zooplankton, snails, amphipods and tadpoles. Most biological responses were negatively associated with soluble carbon content of litter, or litter decay rate. Other litter traits, including phenolic concentrations and litter C:N were of secondary importance but did exhibit both positive and negative associations with several responses. The absence of litter had pervasive effects on abiotic attributes, but did not promote substantial changes in organism biomass. Most responses to the litter mixture were additive. Our results suggest that changes in temperate forest composition can strongly affect pond communities.     

Species divergence and trait convergence in experimental plant community assembly

Despite decades of research, it remains controversial whether ecological communities converge towards a common structure determined by environmental conditions irrespective of assembly history. Here, we show experimentally that the answer depends on the level of community organization considered. In a 9‐year grassland experiment, we manipulated initial plant composition on abandoned arable land and subsequently allowed natural colonization. Initial compositional variation caused plant communities to remain divergent in species identities, even though these same communities converged strongly in species traits. This contrast between species divergence and trait convergence could not be explained by dispersal limitation or community neutrality alone. Our results show that the simultaneous operation of trait‐based assembly rules and species‐level priority effects drives community assembly, making it both deterministic and historically contingent, but at different levels of community organization.     

Plant traits, litter quality and decomposition in a Mediterranean old-field succession

Human-induced changes in land use lead to major changes in plant community composition which have strong effects on ecosystem processes. Here, we tested the hypothesis that changes in traits of living plants induced by such changes resulted in changes in the quality and decay properties of the litter produced by the different communities. This was done in the context of a secondary succession following land abandonment in the Mediterranean region of Southern France. During the course of succession, species with high specific leaf area (the ratio of leaf area to leaf mass), low leaf dry matter content (the ratio of leaf dry mass to leaf fresh mass) and high leaf nitrogen concentration were progressively replaced by species with opposite characteristics. Accordingly, the initial litter concentrations of carbon (C) and nitrogen (N) decreased, while their C:N ratio and their hemicellulose concentration increased with time after abandonment. Early-successional communities had faster rates of litter decay and N release from litter, but these differences damped out with decomposition time. Nitrogen release from litter was related to initial litter chemical composition, particularly to its N concentration. This also held for litter decay rate, but only during the first 18 months of decomposition. Community functional parameters (i.e. trait values weighed according to the relative abundance of species) were tightly linked to initial litter N concentration, and thereby to litter decay and N loss rates. The strongest correlations were found with leaf dry matter content, which therefore appears as a powerful marker of litter properties. This provides further evidence that characteristics of living leaves persist in litter, and that some ecosystem processes can be inferred from plant functional traits. Responsible Editor: Alfonso Escudero     

Positive feedbacks between decomposition and soil nitrogen availability along fertility gradients

Background and aims

We determined the relationship between site N supply and decomposition rates with respect to controls exerted by environment, litter chemistry, and fungal colonization.

Methods

Two reciprocal transplant decomposition experiments were established, one in each of two long-term experiments in oak woodlands in Minnesota, USA: a fire frequency/vegetation gradient, along which soil N availability varies markedly, and a long-term N fertilization experiment. Both experiments used native Quercus ellipsoidalis E.J. Hill and Andropogon gerardii Vitman leaf litter and either root litter or wooden dowels.

Results

Leaf litter decay rates generally increased with soil N availability in both experiments while belowground litter decayed more slowly with increasing soil N. Litter chemistry differed among litter types, and these differences had significant effects on belowground (but not aboveground) decay rates and on aboveground litter N dynamics during decomposition. Fungal colonization of detritus was positively correlated with soil fertility and decay rates.

Conclusions

Higher soil fertility associated with low fire frequency was associated with greater leaf litter production, higher rates of fungal colonization of detritus, more rapid leaf litter decomposition rates, and greater N release in the root litter, all of which likely enhance soil fertility. During decomposition, both greater mass loss and litter N release provide mechanisms through which the plant and decomposer communities provide positive feedbacks to soil fertility as ultimately driven by decreasing fire frequency in N-limited soils and vice versa.     

The spatial scaling of saprotrophic fungal beta diversity in decomposing leaves

Assembly of fungal communities remains poorly understood in part because of the daunting range of spatial scales that may be involved in this process. Here, we use individual leaves as a natural sampling unit, comprising spatially distinct habitat and/or resource patches with unique histories and suites of resources. Spatial patterns in fungal beta diversity were tested for consistency with the metacommunity paradigms of species sorting and neutral dynamics. Thirty senesced leaves were collected from the forest floor (O horizon) in replicate upland forest, riparian forest and vernal pool habitats. We quantified spatial distance between leaves, and fungal community composition was assayed by terminal restriction fragment length polymorphism. Significant distance‐decay relationships were detected at all but one upland site. This is the first study where changes in fungal community composition were quantified across discrete adjacent habitat patches, providing evidence that fungal distance decay is operational at a scale of centimetres. Although leaves of differing lignin contents were sampled from each site, leaf type was not consistently important in explaining variation in fungal community composition. However, depth of a leaf within the forest floor significantly influenced community composition at five of six sites. Environmental heterogeneity associated with depth could include moisture gradients, relative influence of soil or spore colonization, and impact of forest floor biotic community (i.e. collembola and earthworms). Because the influence of spatial distance and depth on fungal community composition could not be disentangled, both species‐sorting and neutral processes may be embedded within the distance‐decay relationships that we found.     

Benefits of increased colonist quantity and genetic diversity for colonization depend on colonist identity

Larger numbers of colonists can be more likely to establish and spread due to the benefits provided by either more individuals (quantity) or a greater diversity of genotypes or phenotypes (genetic diversity). However, the value of higher colonist quantity or genetic diversity varies widely across studies, leaving a great deal of uncertainty in how these respective mechanisms affect colonization success. This variability is potentially driven by differences in which traits are present in respective colonist pools (‘colonist identity’). Studies with high‐performing colonizers (e.g. genotypes pre‐adapted to the colonizing environment) may find increasing quantity or diversity to be beneficial because it increases the chance high‐performers are sampled, while studies with no high‐performers may find no effects of quantity or diversity. Alternatively, quantity and genetic diversity may play little to no role if the smallest populations already contain high‐performing colonists because there is no scope for a sampling effect to operate. We conducted a field mesocosm experiment to determine if variability in the benefits provided by increased quantity or genetic diversity relates to colonist traits. Nine distinct genotypes of Daphnia pulex characterized also by phenotype, were introduced in ‘single’ (one individual) or ‘many’ (nine individuals) introduction quantities and at ‘low’ (monoclonal) and ‘high’ (mixed genotypes) genetic diversities. We found that larger‐bodied D. pulex genotypes benefited less from increased colonist quantity, while increasing genetic diversity tended to have a lower effect on higher growth rate genotypes. Our results show that the trait values of the colonists can determine the benefits gained when colonist quantity or genetic diversity are increased, with potential applications to future research and practical efforts to promote, or prevent, population establishment.     

The influence of initial colonization by hydropsychid caddisfly larvae on the development of stream invertebrate assemblages

Many aspects of the colonization history of a disturbed site can influence the development of a biological community. Initial colonization is known to play a significant role in community development because of the facilitative or inhibitory effects that `pioneer' species can have on subsequently arriving taxa. We performed an experiment to assess how initial colonization by two species of benthic invertebrates (Trichoptera: Hydropsychidae) might influence the development of stream faunal assemblages. Substrate baskets initially colonized by either Hydropsyche depravata or Ceratopsyche bronta were placed alongside control baskets in a recently flooded stream. After baskets had colonized for 30 days, we found that species composition in treatment baskets was identical to that in control baskets, indicating that the caddisfly taxa had no selective effects on colonization of other macroinvertebrate species. We did, however, find that C. bronta facilitated the recruitment of all species in the colonist pool leading to greater overall abundance and biomass of macroinvertebrates in the final assemblages. In contrast, H. depravata had no effect on the abundance or biomass of colonizing invertebrates. The differential effects of these two taxa on abundance and biomass may have been related to differences in microhabitat complexity created by the construction of their retreats and catchnets. The results of this study support the growing recognition that colonization history does influence the structure of lotic communities, but they also suggest that even closely related taxa can play different roles as initial colonists in community development.     

Solar UV-B decreases decomposition in herbaceous plant litter in Tierra del Fuego, Argentina: potential role of an altered decomposer community

Tierra del Fuego, Argentina (55°S), receives increased solar ultraviolet‐B radiation (UV‐B) as a result of Antarctic stratospheric ozone depletion. We conducted a field study to examine direct and indirect effects of solar UV‐B radiation on decomposition of Gunnera magellanica, a native perennial herb, and on the native community of decomposer organisms. In general, indirect effects of UV‐B mostly occur due to changes in the chemical composition of litter, whereas direct effects during decomposition result from changes in decomposer organisms and/or differences in the photochemical breakdown of litter. We designed a full‐factorial experiment using senescent leaves that had received either near‐ambient or attenuated UV‐B during growth. The leaves were distributed in litterbags and allowed to decompose under near‐ambient or reduced solar UV‐B during the growing season. We evaluated initial litter quality, mass loss, and nutrient release of decomposing litter, and microbial colonization of both initial litter and decomposed litter. We found that litter that decomposed under near‐ambient UV‐B had significantly less mass loss than litter that decomposed under reduced UV‐B. The UV‐B conditions received by plants during growth, which did not affect mass loss and nutrient composition of litter, affected fungal species composition but in different ways throughout the decomposition period. Before the decomposition trial, Beauveria bassiana and Penicillium frequentans were higher under reduced UV‐B, whereas Cladosporium herbarum and pigmented bacteria were more common under the near‐ambient compared to the reduced UV‐B treatment. After the decomposition period, leaves that had grown under reduced UV‐B showed higher frequency of Penicillium thomii and lower frequency of Trichoderma polysporum than leaves that had grown under near‐ambient conditions. The UV‐B condition received during decomposition also affected fungal colonization, with Penicillium chrysogenum being more frequent in leaves that had decomposed under reduced UV‐B, while the other species were not affected. Our results demonstrate that, in this ecosystem, the effects of UV‐B radiation on decomposition apparently occurred mostly through changes in the fungal community, while changes in photochemical breakdown appeared to be less important.     

Community monopolization: local adaptation enhances priority effects in an evolving metacommunity

The diversity and composition of biological communities might often depend on colonization history because early colonists can exclude future colonists through a priority effect. These priority effects, which have been observed across a wide variety of ecosystems, often arise because early colonists have sufficient time to use available resources efficiently and subsequently withhold them from invaders. Here, we explore the extent to which rapid local adaptive evolution contributes to the pervasiveness of these priority effects. Using an individual-based simulation, we show that early colonization allows the descendants of colonists to adapt to novel conditions and reduce the establishment success of an initially ecologically equivalent competing species. Our model predicts that slight differences in colonization timing and adaptive capacity between species can substantially alter the dynamics and diversity of communities. We also show that priority effects and gene flow can generate a novel mechanism for the expansion and retraction of species distributions in a metacommunity. Our results suggest that local adaptation combined with stochastic colonization events can obscure direct relationships between species distributions and environmental gradients. Given the increasing recognition of rapid, microgeographic evolution in natural populations, we expect that evolutionary priority effects could affect the structure and dynamics of many natural metacommunities.     

Placing the Effects of Leaf Litter Diversity on Saprotrophic Microorganisms in the Context of Leaf Type and Habitat

Because of conflicting results in previous studies, it is unclear whether litter diversity has a predictable impact on microbial communities or ecosystem processes. We examined whether effects of litter diversity depend on factors that could confound comparisons among previous studies, including leaf type, habitat type, identity of other leaves in the mixture, and spatial covariance at two scales within habitats. We also examined how litter diversity affects the saprotrophic microbial community using terminal restriction fragment length polymorphism to profile bacterial and fungal community composition, direct microscopy to quantify bacterial biomass, and ergosterol extraction to quantify fungal biomass. We found that leaf mixture diversity was rarely significant as a main effect (only for fungal biomass), but was often significant as an interaction with leaf type (for ash-free dry mass recovered, carbon-to-nitrogen ratio, fungal biomass, and bacterial community composition). Leaf type and habitat were significant as main effects for all response variables. The majority of variance in leaf ash-free dry mass and C/N ratio was explained after accounting for treatment effects and spatial covariation at the meter (block) and centimeter (litterbag) scales. However, a substantial amount of variability in microbial communities was left unexplained and must be driven by factors at other spatial scales or more complex spatiotemporal dynamics. We conclude that litter diversity effects are primarily dependent on leaf type, rather than habitat type or identity of surrounding leaves, which can guide the search for mechanisms underlying effects of litter diversity on ecosystem processes.     

Macroinvertebrate identity mediates the effects of litter quality and microbial conditioning on leaf litter recycling in temperate streams

Plant litter decomposition is an essential ecosystem function that contributes to carbon and nutrient cycling in streams. Aquatic shredders, mainly macroinvertebrates, can affect this process in various ways; they consume leaf litter, breaking it down into fragments and creating suitable habitats or resources for other organisms through the production of fine particulate organic matter (FPOM). However, measures of litter‐feeding traits across a wide range of aquatic macroinvertebrates are still rare. Here, we assessed the contributions of 11 species of freshwater macroinvertebrates to litter decomposition, by measuring consumption rate, FPOM production, and assimilation rate of highly decomposable (Alnus glutinosa) or poorly decomposable (Quercus robur) leaf litter types. In general, an increase in the quality of litter improved the litter consumption rate, and fungal conditioning of the leaf litter increased both the litter consumption rate and FPOM production. Macroinvertebrates specializing in leaf litter consumption also appeared to be the most sensitive to shifts in litter quality and the conditioning process. Contrary to expectations, the conditioning process did not increase the assimilation of low‐quality litter. There was a strong correlation between the relative consumption rate (RCR) of the two litter types, and the relative FPOM production (RFP) was strongly correlated to the RCR. These findings suggest a consistent relationship between RCR and macroinvertebrate identity that is not affected by litter quality, and that the RFP could be inferred from the RCR. The varying responses of the macroinvertebrate feeding traits to litter quality and the conditioning process suggest that the replacement of a shredder invertebrate species by another species could have major consequences for the decomposition process and the detritus‐based food web in streams. Further studies onto the importance of invertebrate identity and the effects of litter quality in a variety of freshwater ecosystems are needed to understand the whole ecosystem functioning and to predict its response to environmental changes.     

Leaf litter traits drive community structure and functioning in a natural aquatic microcosm

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Pesticide alters habitat selection and aquatic community composition

Anthropogenic chemical contamination is an important issue for conservation of aquatic ecosystems. While recent research highlights that community context can mediate the consequences of contaminant exposure, little is known about how contaminants themselves might determine this context by altering habitat selection and thus initial community composition. Here we show that the insecticide carbaryl and its commercial counterpart Sevin can affect aquatic community composition by differentially altering oviposition and colonization of experimental pools by amphibians and insects. On average, contaminated pools received 20-fold more adult beetle and heteropteran colonists and 12-fold more Culex mosquito and chironomid midge egg masses. On the other hand, ovipositing Anopheles mosquitoes and cricket frogs showed no preference and we have shown previously that gray treefrogs strongly avoid contaminated pools. Overall, initial richness doubled in contaminated pools compared with controls. By affecting colonizing taxa differently and increasing richness, the contaminant may alter the ecological context under which subsequent effects of exposure will unfold. Given that community context is important for evaluating toxicity effects, understanding the net effects of contaminants in natural systems requires an understanding of their effects on community assembly via shifts in habitat selection.     

Litter and soil biodiversity jointly drive ecosystem functions

The decomposition of litter and the supply of nutrients into and from the soil are two fundamental processes through which the above- and belowground world interact. Microbial biodiversity, and especially that of decomposers, plays a key role in these processes by helping litter decomposition. Yet the relative contribution of litter diversity and soil biodiversity in supporting multiple ecosystem services remains virtually unknown. Here we conducted a mesocosm experiment where leaf litter and soil biodiversity were manipulated to investigate their influence on plant productivity, litter decomposition, soil respiration, and enzymatic activity in the littersphere. We showed that both leaf litter diversity and soil microbial diversity (richness and community composition) independently contributed to explain multiple ecosystem functions. Fungal saprobes community composition was especially important for supporting ecosystem multifunctionality (EMF), plant production, litter decomposition, and activity of soil phosphatase when compared with bacteria or other fungal functional groups and litter species richness. Moreover, leaf litter diversity and soil microbial diversity exerted previously undescribed and significantly interactive effects on EMF and multiple individual ecosystem functions, such as litter decomposition and plant production. Together, our work provides experimental evidence supporting the independent and interactive roles of litter and belowground soil biodiversity to maintain ecosystem functions and multiple services.     

Fungal endophyte‐infected leaf litter alters in‐stream microbial communities and negatively influences aquatic fungal sporulation

Endophytes are ubiquitous plant‐associated microbes and although they have the potential to alter the decomposition of infected leaf litter, this has not been well‐studied. The endophyte Rhytisma punctatum infects the leaves of Acer macrophyllum (bigleaf maple), causing the appearance of black ‘tar spots’ that persist in senesced leaves. Other foliar fungi also cause visible damage in healthy tissues of this host plant system including an unidentified bullseye‐shaped lesion, common in western Washington. Using three treatments of endophyte infection status in leaf tissue (R. punctatum‐infected, bullseye‐infected, lesion‐free), leaf litter discs were submerged in a third‐order temperate stream using mesh litter bags and harvested periodically over two months to determine the effects of litter treatment and incubation time on litter mass loss, fungal sporulation, and microbial community colonization. Litter containing symptomatic endophyte infections (Rhytisma or bullseye) had reduced sporulation of aquatic hyphomycetes, but decomposed significantly faster than lesion‐free or bullseye‐infected litter. Using amplicon‐based sequencing, we found a significant difference in bacterial communities colonizing Rhytisma‐infected and bullseye‐infected leaf litter, a significant difference in fungal communities colonizing Rhytisma‐infected leaf litter compared to the two other treatments, and a change in both community structure and relative abundances of bacterial and fungal taxa throughout the study period. Indicator Species Analysis clarified the drivers of these community shifts at the genus level. Our results show that endophyte‐associated, in‐stream sporulation and microbial community effects are observable within one species of leaf litter.     

Comparison of leaf decomposition and macroinvertebrate colonization between exotic and native trees in a freshwater ecosystem

One of the most important sources of energy in aquatic ecosystems is the allochthonous input of detritus. Replacement of native tree species by exotic ones affects the quality of detritus entering freshwater ecosystems. This replacement can alter nutrient cycles and community structure in aquatic ecosystems. The aims of our study were (1) to compare leaf litter decomposition of two widely distributed exotic species (Ailanthus altissima and Robinia pseudoacacia) with the native species they coexist with (Ulmus minor and Fraxinus angustifolia), and (2) to compare macroinvertebrate colonization among litters of the invasive and native species. Litter bags of the four tree species were placed in the water and collected every 2, 25, 39, 71, and 95 days in a lentic ecosystem. Additionally, the macroinvertebrate community on litter bags was monitored after 25, 39, and 95 days. Several leaf chemistry traits were measured at the beginning (% lignin; lignin:N, C:N, LMA) and during the study (leaf total nitrogen). We detected variable rates of decomposition among species (k values of 0.009, 0.008, 0.008, and 0.005 for F. angustifolia, U. minor, A. altissima and R. pseudoacacia, respectively), but we did not detect an effect of litter source (from native/exotic). In spite of its low decay, the highest leaf nitrogen was found in R. pseudoacacia litter. Macroinvertebrate communities colonizing litter bags were similar across species. Most of them were collectors (i.e., they feed on fine particulate organic matter), suggesting that leaf material of either invasive or native trees was used as substrate both for the animals and for the organic matter they feed on. Our results suggest that the replacement of the native Fraxinus by Robinia would imply a reduction in the rate of leaf processing and also a slower release of leaf nitrogen to water.     

The effect of sunlight on leaf litter quality reduces growth of the shredder Klapopteryx kuscheli

• 1 The direct effect of sunlight on the conditioning, breakdown and incorporation of leaf litter in stream food webs has not yet been considered. The aim here was to evaluate the effects of light intensity on the colonization of leaf litter by microorganisms and its resulting quality as food for the stonefly shredder Klapopteryx kuscheli.
• 2 Leaf litter was conditioned for 2 months in an open reach of a second‐order stream in litter bags either exposed to or shaded from direct sunlight. Subsequently, we performed laboratory experiments to test larval consumption, growth, growth efficiency and feeding preference fed on both leaf litter treatments.
• 3 Leaf litter in the unshaded treatment had three times more chlorophyll‐a (Chl‐a) than that in the shaded treatment, 50% lower fungal biomass and similar bacterial abundance. Although larvae did not prefer either food and fed at the same rate on both leaf litter treatments, they grew twice as fast on the shade‐conditioned leaves and attained a two‐fold higher growth efficiency.
• 4 Sunlight can have significant effects on detritus‐based food webs. Riparian modification induced by human activities in forested catchments increases the potential for sunlight to influence detritus dynamics.

     

Enhanced summer warming reduces fungal decomposer diversity and litter mass loss more strongly in dry than in wet tundra

Many Arctic regions are currently experiencing substantial summer and winter climate changes. Litter decomposition is a fundamental component of ecosystem carbon and nutrient cycles, with fungi being among the primary decomposers. To assess the impacts of seasonal climatic changes on litter fungal communities and their functioning, Betula glandulosa leaf litter was surface‐incubated in two adjacent low Arctic sites with contrasting soil moisture regimes: dry shrub heath and wet sedge tundra at Disko Island, Greenland. At both sites, we investigated the impacts of factorial combinations of enhanced summer warming (using open‐top chambers; OTCs) and deepened snow (using snow fences) on surface litter mass loss, chemistry and fungal decomposer communities after approximately 1 year. Enhanced summer warming significantly restricted litter mass loss by 32% in the dry and 17% in the wet site. Litter moisture content was significantly reduced by summer warming in the dry, but not in the wet site. Likewise, fungal total abundance and diversity were reduced by OTC warming at the dry site, while comparatively modest warming effects were observed in the wet site. These results suggest that increased evapotranspiration in the OTC plots lowered litter moisture content to the point where fungal decomposition activities became inhibited. In contrast, snow addition enhanced fungal abundance in both sites but did not significantly affect litter mass loss rates. Across sites, control plots only shared 15% of their fungal phylotypes, suggesting strong local controls on fungal decomposer community composition. Nevertheless, fungal community functioning (litter decomposition) was negatively affected by warming in both sites. We conclude that although buried soil organic matter decomposition is widely expected to increase with future summer warming, surface litter decay and nutrient turnover rates in both xeric and relatively moist tundra are likely to be significantly restricted by the evaporative drying associated with warmer air temperatures.     

Tree litter functional diversity and nitrogen concentration enhance litter decomposition via changes in earthworm communities

Biodiversity is a major driver of numerous ecosystem functions. However, consequences of changes in forest biodiversity remain difficult to predict because of limited knowledge about how tree diversity influences ecosystem functions. Litter decomposition is a key process affecting nutrient cycling, productivity, and carbon storage and can be influenced by plant biodiversity. Leaf litter species composition, environmental conditions, and the detritivore community are main components of the decomposition process, but their complex interactions are poorly understood. In this study, we tested the effect of tree functional diversity (FD) on litter decomposition in a field experiment manipulating tree diversity and partitioned the effects of litter physiochemical diversity and the detritivore community. We used litterbags with different mesh sizes to separate the effects of microorganisms and microfauna, mesofauna, and macrofauna and monitored soil fauna using pitfall traps and earthworm extractions. We hypothesized that higher tree litter FD accelerates litter decomposition due to the availability of complementary food components and higher activity of detritivores. Although we did not find direct effects of tree FD on litter decomposition, we identified key litter traits and macrodetritivores that explained part of the process. Litter mass loss was found to decrease with an increase in leaf litter carbon:nitrogen ratio. Moreover, litter mass loss increased with an increasing density of epigeic earthworms, with most pronounced effects in litterbags with a smaller mesh size, indicating indirect effects. Higher litter FD and litter nutrient content were found to increase the density of surface‐dwelling macrofauna and epigeic earthworm biomass. Based on structural equation modeling, we conclude that tree FD has a weak positive effect on soil surface litter decomposition by increasing the density of epigeic earthworms and that litter nitrogen‐related traits play a central role in tree composition effects on soil fauna and decomposition.