Effects Of Litter Mixing Research Articles

Incubation time, functional litter diversity, and habitat characteristics predict litter-mixing effects on decomposition

Plant diversity influences many fundamental ecosystem functions, including carbon and nutrient dynamics, during litter breakdown. Mixing different litter species causes litter mixtures to lose mass at different rates than expected from component species incubated in isolation. Such nonadditive litter-mixing effects on breakdown processes often occur idiosyncratically because their direction and magnitude change with incubation time, litter species composition, and ecosystem characteristics. Taking advantage of results from 18 litter mixture experiments in streams, we examined whether the direction and magnitude of nonadditive mixing effects are randomly determined. Across 171 tested litter mixtures and 510 incubation time-by-mixture combinations, nonadditive effects on breakdown were common and on average resulted in slightly faster decomposition than expected. In addition, we found that the magnitude of nonadditive effects and the relative balance of positive and negative responses in mixtures change predictably over time, and both were related to an index of functional litter diversity and selected environmental characteristics. Based on these, it should be expected that nonadditive effects are stronger for litter mixtures made of functionally dissimilar species especially in smaller streams. Our findings demonstrate that effects of litter diversity on plant mixture breakdown are more predictable than generally thought. We further argue that the consequences of current worldwide homogenization in the composition of plant traits on carbon and nutrient dynamics could be better inferred from long-duration experiments that manipulate both functional litter diversity and ecosystem characteristics in "hotspots of biodiversity effects," such as small streams.

Macro‐detritivore identity drives leaf litter diversity effects

The importance of leaf litter diversity for decomposition, an important process in terrestrial ecosystems, is much debated. Previous leaf litter‐mixing studies have shown that non‐additive leaf litter diversity effects can occur, but it is not clear why they occurred in only half of the studies and which underlying mechanisms can explain these conflicting results. We hypothesized that incorporating the role of macro‐detritivores could be important. Although often ignored, macro‐detritivores are known to strongly influence decomposition. To better understand the importance of macro‐detritivores for leaf litter mixing effects during decomposition, four common leaf litter species were added separately and in two and four species combinations to monocultures of three different macro‐detritivores and a control without fauna. Our results clearly show that leaf litter‐mixing effects occurred only in the presence of two macro‐detritivores (earthworms and woodlice). Application of the additive partitioning method revealed that in the specific combination of woodlice and the presence of a slow‐decomposing leaf litter species in the mixture, leaf litter mixing effects were strongly driven by a selection effect. This was caused by food preference of the isopod: the animals avoided the slow decomposing species when given the choice. However, most leaf litter mixing effects were caused by complementarity effects. The potential mechanisms underlying the complementarity effects are discussed. Our results clearly show that that both leaf litter and macro‐detritivore identity can affect litter diversity. This may help to explain the conflicting results obtained in previous experiments.

Nutrient dynamics in litter mixtures of four Mediterranean maquis species decomposing in situ

In natural conditions, litters shed from different species become mixed with each other, and decompose together. Most studies deal with decomposition of individual species; few studies investigate the influence of litter mixing on decomposition and nutrient dynamics; the results are contradictory as positive, negative, or no effect, of litter mixing have been observed. In this study we test the hypothesis: i) that litter mixing in the Mediterranean maquis, a nutrient poor, high diversity ecosystem, produces non-additive effects on nutrient dynamics; ii) that the effects vary with the composition in species of the mixture and with the relative amount of the species component the mixture. Two types of 3-species mixtures were set up; one contained three sclerophylls, Phillyrea angustifolia, Pistacia lentiscus and Quercus ilex; the other contained the first two species with the mesophyll Cistus. Litterbags, containing monospecific litters and even and uneven mixtures, were incubated under natural condition in situ; even mixtures had the 3 species in equal proportion, whereas uneven mixtures had one of the species as dominant (50%) and the other two species in equal proportion (25%:25%). Litterbags were retrieved after 92, 188 and 403 days; litters from the mixtures were separately analyzed for mass loss and content of nitrogen (N), potassium (K), sodium (Na), calcium (Ca), magnesium (Mg), manganese (Mn), iron (Fe), copper (Cu), and zinc (Zn). Results indicate that mixing influences the dynamics of N, Mn, Ca, Mg, Fe, Cu and Zn, but scarcely affects the dynamics of K and Na. The comparison of observed to expected values for changes of nutrients in litterbags indicates the occurrence of non-additive effects of litter mixing on movements of N, Fe, Cu, and Zn to or from the litterbags containing the mixtures. The effects depend on the composition in species of the mixture, whereas the relative amount of the species component the mixture is not relevant.

Living in the litter: the influence of tree leaf litter on wetland communities

Empirical research in streams has demonstrated that terrestrial subsidies of tree leaf litter influence multiple community factors including composition, diversity and growth of individuals. However, little research has examined the importance of tree litter species on wetlands, which are ubiquitous across the landscape and serve as important habitats for a unique and diverse community of organisms. Using outdoor mesocosms, we assessed the impact of 12 litter monocultures and three litter mixtures (from both broadleaf and conifer trees) on pond communities containing gray tree frog tadpoles Hyla versicolor, periphyton, phytoplankton and zooplankton. We found that leaf litter species had substantial and differential impacts on all trophic groups in the community including effects on algal abundance, zooplankton density and amphibian growth. In many instances, patterns of responses were specific to individual litter species yet some responses, including both pH values and periphyton biomass, were generalizable to broad taxonomic groups. In addition, while most responses of litter mixtures were additive, we found evidence for antagonistic effects of litter mixing among responses of periphyton and amphibian body mass. Our results highlight the potential impact of human and naturally driven changes in forest composition on wetland communities through associated changes in leaf litter.

Snail and millipede complementarity in decomposing Mediterranean forest leaf litter mixtures

Summary 1. The projected increase in loss of biodiversity worldwide has prompted the need to understand the role that diversity plays in key ecosystem functions such as litter decomposition and nutrient cycling. Here we asked how two contrasting species of saprophagous macrofauna and four Mediterranean forest leaf litter species interactively affect decomposition in a 6‐week microcosm study. 2. Litter mass loss and macrofauna relative consumption rates (RCR) were measured on treatments with freshly fallen or partially decomposed leaf litter of Alnus glutinosa, Fraxinus angustifolia, Pistacia terebinthus and Quercus ilex as single‐species or mixtures in absence and presence of the gastropod, Pomatias elegans, the diplopod, Glomeris marginata, or both macrofauna species. 3. Macrofauna consumed all litter substrates except freshly fallen P. terebinthus as a single‐species litter that was fatal to both animals, although its presence in litter mixtures increased overall RCR. Consumption was higher in partially decomposed than freshly fallen litter, and higher in litter mixtures than in single‐species litter. Both litter state and mixing interacted significantly with macrofauna treatment where generally, RCR by P. elegans alone was inferior to that of G. marginata alone or in combination with P. elegans. 4. An overall positive complementarity effect on litter RCR between G. marginata and P. elegans was observed in freshly fallen litter. Particularly strong complementarity was observed in two mixtures of freshly fallen litter and also in one mixture of partially decomposed litter. 5. There were no non‐additive effects of litter mixing on litter mass loss in the absence of animals, indicating no interactions among litter substrates during decomposition. However, in the presence of either G. marginata and/or P. elegans, positive and negative interactions among litter substrates occurred and were enhanced or reversed by the addition of the second macrofauna species. 6. We conclude that saprophagous macrofauna play a critical role in the decomposition dynamics of Mediterranean forest litter by interacting with each other and by driving interactions among litter substrates.

Does mixing of senescent and green leaves result in nonadditive effects on leaf decomposition?

Most studies of the effects of litter mixing have focused on decomposition in mixtures of senescent leaves of different plant species. In temperate forests, substantial quantities of green leaves fall in spring and summer, and previous research has shown that stream detritivores can benefit from the combined use of senescent and green leaves. We conducted a litterbag experiment to test the hypothesis that mixtures of senescent and green leaves of different species decomposed more rapidly, mediated by increased detritivore colonization, than leaves of each type in isolation. The observed abundances of 2 detritivore species, the amphipod Sternomoera rhyaca and the caddisfly Lepidostoma complicatum, were 3 to 6× higher in mixtures of senescent oak and green alder leaves than expected based on litter patches consisting of each species alone. Lepidostoma complicatum also responded positively to mixtures of senescent oak and green maple leaves, but S. rhyaca did not. These results verify that caddisfly behavior can reflect benefits gained from the combined use of senescent and green leaves. In contrast, the extent of the advantage to amphipods depends on the specific combination of senescent and green leaves. Despite the synergistic effect of mixed senescent oak and green alder leaves on detritivore colonization, observed leaf mass loss in mixtures was not higher than expected. Changes in the chemical properties of senescent oak and green alder leaves in mixtures might decrease feeding rates of detritivores because of preferential or compensatory feeding. Mixtures of senescent and green leaves did not alter leaf decomposition of each component in spring and summer, but might enhance leaf decomposition in autumn and winter by augmenting detritivore populations.

Species shift drives decomposition rates following invasion by hemlock woolly adelgid

Insect and disease outbreak is an important cause of selective species removal and accompanying functional change in North American forests. Outbreak of hemlock woolly adelgid, Adelgies tsugae– HWA, is causing selective removal of eastern hemlock Tsuga canadensis at a regional scale. Impacts of outbreak-caused canopy mortality and shifts in dominant species on litter decay were compared across sites that range in HWA-caused canopy damage and subsequent canopy dominance by black birch Betula lenta. Senescent litter from eastern hemlock, black birch, and equal litter mixes were decomposed in the field for 36 months within nine sites in Connecticut and Massachusetts USA. Mass loss and % N accumulation of black birch was 65% and 52% greater compared to eastern hemlock. In contrast, outbreak related canopy damage increased litter mass loss by 11.5% in high mortality stands relative to uninfested stands but canopy damage had no impact on % N dynamics. Non-additive effects of litter mixing influenced chemical dynamics of decaying litter; black birch accumulated less N and eastern hemlock accumulated more N compared to each species decaying alone. However, these changes offset and mixed litter bags overall showed no differences in N dynamics compared to values from each species decaying alone. In eastern hemlock stands invaded by hemlock woolly adelgid, canopy damage influences the rates and dynamics of decay but species differences between hemlock and black birch leaf litter are the dominant mechanisms of decomposition changes and a long-lasting driver of increased N cycling rates. Species shifts may be the dominant driver of altered ecosystem processes for other insect outbreaks, particularly when replacement species have very different characteristics regulating decomposition and N cycling.

Non‐additive effects of litter mixing are suppressed in a nutrient‐enriched stream

Investigations of how species compositional changes interact with other aspects of global change, such as nutrient mobilization, to affect ecosystem processes are currently lacking. Many studies have shown that mixed species plant litters exhibit non‐additive effects on ecosystem functions in terrestrial and aquatic systems. Using a full‐factorial design of three leaf litter species with distinct initial chemistries (carbon:nitrogen; C:N) and breakdown rates (Liriodendron tulipifera, Acer rubrum and Rhododendron maximum), we tested for additive and non‐additive effects of litter species mixing on breakdown in southeastern US streams with and without added nutrients (N and phosphorus). We found a non‐additive (antagonistic) effect of litter mixing on breakdown rates under reference conditions but not when nutrient levels were elevated. Differential responses among single‐species litters to nutrient enrichment contributed to this result. Antagonistic litter mixing effects on breakdown were consistent with trends in litter C:N, which were higher for mixtures than for single species, suggesting lower microbial colonization on mixtures. Nutrient enrichment lowered C:N and had the greatest effect on the lowest‐ (R. maximum) and the least effect on the highest‐quality litter species (L. tulipifera), resulting in lower interspecific variation in C:N. Detritivore abundance was correlated with litter C:N in the reference stream, potentially contributing to variation in breakdown rates. In the nutrient‐enriched stream, detritivore abundance was higher for all litter and was unrelated to C:N. Thus, non‐additive effects of litter mixing were suppressed by elevated streamwater nutrients, which increased nutrient content of all litter, reduced variation in C:N among litter species and increased detritivore abundance. Nutrients reduced interspecific variation among plant litters, the base of important food web pathways in aquatic ecosystems, affecting predicted mixed‐species breakdown rates. More generally, world‐wide mobilization of nutrients may similarly modify other effects of biodiversity on ecosystem processes.

Litter Mixture Interactions at the Level of Plant Functional Types are Additive

It is very difficult to estimate litter decomposition rates in natural ecosystems because litters of many species are mixed and idiosyncratic interactions occur among those litters. A way to tackle this problem is to investigate litter mixing effects not at the species level but at the level of Plant Functional Types (PFTs). We tested the hypothesis that at the PFT level positive and negative interactions balance each other, causing an overall additive effect (no significant interactions among PFTs). Thereto, we used litter of four PFTs from a temperate peatland in which random draws were taken from the litter species pool of each PFT for every combination of 2, 3, and 4 PFTs. Decomposition rates clearly differed among the 4 PFTs (Sphagnum spp. < graminoids = N-fixing tree < forbs) and showed little variation within the PFTs (notably for the Sphagnum mosses and the graminoids). Significant positive interactions (4 out of 11) in the PFT mixtures were only found after 20 weeks and in all these combinations Sphagnum was involved. After 36 and 56 weeks of incubation interactions were not significantly different from zero. However, standard deviations were larger than the means, indicating that positive and negative interactions balanced each other. Thus, when litter mixture interactions are considered at the PFT level the interactions are additive. From this we conclude that for estimating litter decomposition rates at the ecosystem level, it is sufficient to use the weighted (by litter production) average decomposition rates of the contributing PFTs.

Litter decomposition and nutrient release as affected by soil nitrogen availability and litter quality in a semiarid grassland ecosystem

Nitrogen availability is critically important to litter decomposition, especially in arid and semiarid areas where N is limiting. We studied the relative contributions of litter quality and soil N to litter decomposition of two dominant grassland species, Stipa krylovii and Artemisia frigida, in a semiarid typical steppe ecosystem in Inner Mongolia, China. The study had four different rates of N addition (0, 8, 32, and 64 g N m(-2) year(-1)), and litter samples were decomposed under varying site conditions and by litter types. Litter-mixing effects of the two species were also examined. We found that N addition increased litter N concentration and thus enhanced litter decomposition by improving substrate quality. This increase, however, was offset by the negative effect of increased soil N, resulting in a diminished effect of increased soil N availability on in situ litter decomposition. The positive effects of improved litter quality slightly out-performed the negative effects of increased soil N. Our further analysis revealed that the negative effect of increasing soil N on litter decomposition could be partially explained by reduced soil microbial biomass and activity. Decomposition was significantly faster for litters of a two-species mixture than litters of the single species, but the rate of litter decomposition did not differ much between the two species, suggesting that compositional balance, rather than changes in the dominance between Stipa and Artemisia, is more critical for litter decomposition, hence nutrient cycling in this ecosystem. This semiarid steppe ecosystem may become more conservative in nutrient use with switching of dominance from Artemisia to Stipa with increasing soil N, because Stipa has a slower decomposition rate and a higher nutrient retention rate than Artemisia.

Neighbour identity hardly affects litter-mixture effects on decomposition rates of New Zealand forest species

The mass loss of litter mixtures is often different than expected based on the mass loss of the component species. We investigated if the identity of neighbour species affects these litter-mixing effects. To achieve this, we compared decomposition rates in monoculture and in all possible two-species combinations of eight tree species, widely differing in litter chemistry, set out in two contrasting New Zealand forest types. Litter from the mixed-species litter bags was separated into its component species, which allowed us to quantify the importance of litter-mixing effects and neighbour identity, relative to the effects of species identity, litter chemistry and litter incubation environment. Controlling factors on litter decomposition rate decreased in importance in the order: species identity (litter quality) >> forest type >> neighbour species. Species identity had the strongest influence on decomposition rate. Interspecific differences in initial litter lignin concentration explained a large proportion of the interspecific differences in litter decomposition rate. Litter mass loss was higher and litter-mixture effects were stronger on the younger, more fertile alluvial soils than on the older, less-fertile marine terrace soils. Litter-mixture effects only shifted percentage mass loss within the range of 1.5%. There was no evidence that certain litter mixtures consistently showed interactive effects. Contrary to common theory, adding a relatively fast-decomposing species generally slowed down the decomposition of the slower decomposing species in the mixture. This study shows that: (1) species identity, litter chemistry and forest type are quantitatively the most important drivers of litter decomposition in a New Zealand rain forest; (2) litter-mixture effects—although statistically significant—are far less important and hardly depend on the identity and the chemical characteristics of the neighbour species; (3) additive effects predominate in this ecosystem, so that mass dynamics of the mixtures can be predicted from the monocultures.

Resource–consumer diversity: testing the effects of leaf litter species diversity on stream macroinvertebrate communities

Summary1. Understanding relationships between resource and consumer diversity is essential to predicting how changes in resource diversity might affect several trophic levels and overall ecosystem functioning.2. We tested for the effects of leaf litter species diversity (i.e. litter mixing) on litter mass remaining and macroinvertebrate communities (taxon diversity, abundance and biomass) during breakdown in a detritus‐based headwater stream (North Carolina, U.S.A.). We used full‐factorial analyses of single‐ and mixed‐species litter from dominant riparian tree species with distinct leaf chemistries [red maple (Acer rubrum), tulip poplar (Liriodendron tulipifera), chestnut oak (Quercus prinus) and rhododendron (Rhododendron maximum)] to test for additivity (single‐species litter presence/absence effects) and non‐additivity (emergent effects of litter species interactions).3. Significant non‐additive effects of litter mixing on litter mass remaining were explained by species composition, but not richness, and litter‐mixing effects were variable throughout breakdown. Specifically, small differences in observed versus expected litter mass remaining were measured on day 14; whereas observed litter mass remaining in mixed‐species leaf packs was significantly higher on day 70 and lower on day 118 than expected from data for single‐species leaf packs.4. Litter mixing had non‐additive effects on macroinvertebrate community structure. The number of species in litter mixtures (two to four), but not litter species composition, was a significant predictor of the dominance of particular macroinvertebrates (i.e. indicator taxa) within mixed‐species packs.5. In addition, the presence/absence of high‐ (L. tulipifera) and low‐quality (R. maximum) litter had additive effects on macroinvertebrate taxon richness, abundance and biomass. The presence ofL. tulipiferalitter had both positive (synergistic) and negative (antagonistic) effects on invertebrate taxon richness, that varied during breakdown but were not related to litter chemistry. In contrast, the presence/absence ofL. tulipiferahad a negative relationship with total macroinvertebrate biomass (due to low leaf mass remaining whenL. tulipiferawas present and higher condensed and hydrolysable tannins associated with leaf packs lackingL. tulipifera). Macroinvertebrate abundance was consistently lower whenR. maximumwas present, which was partially explained by litter chemistry [e.g., high concentrations of lignin, condensed tannins, hydrolysable tannins and total phenolics and high carbon to nutrient (N and P) ratios].6. The bottom‐up effects of litter species diversity on stream macroinvertebrates and litter breakdown are different, which suggests that structural attributes of macroinvertebrate communities may only partially explain the effects of litter‐mixing on organic matter processing in streams. In addition, stream macroinvertebrates colonising decomposing litter are influenced by resource diversity as well as resource availability. Broad‐scale shifts in riparian tree species composition will alter litter inputs to streams, and our results suggest that changes in the diversity and availability of terrestrial litter may alter stream food webs and organic matter processing.

Linkages between below and aboveground communities: Decomposer responses to simulated tree species loss are largely additive

Inputs of aboveground plant litter influence the abundance and activities of belowground decomposer biota. Litter-mixing studies have examined whether the diversity and heterogeneity of litter inputs affect decomposer communities in ways that can be predicted from monocultures. They have mainly attempted to detect non-additive effects of litter mixing, although individual species effects (additivity) as well as species interactions (non-additivity) may alter decomposition rates. To determine potential impacts of plant species loss on aboveground-decomposer linkages, we assessed both additive and non-additive effects of litter mixing on decomposer communities. A full-factorial litterbag experiment with leaves from four deciduous tree species was conducted, to assess responses of bacteria, fungi, nematodes, and microarthropods. Data were analyzed using a statistical method that first looked for additive effects based on the presence or absence of species and then any significant species interactions. We observed almost exclusively additive effects of all four litter species on decomposer biota, with each species exerting effects on different aspects of the community. These results imply that the consequences of species loss for the decomposer community will be largely predictable from knowledge of single species litter dynamics. The two species at opposite ends of the quality spectrum exerted the most effects. High-quality Liriodendron tulipifera supported a more diverse arthropod community and drove bottom-up effects on the decomposer food web. Low-quality Rhododendron maximum had negative effects on most groups of biota. Litter of mid-quality species exerted fewer effects. The influence of litter species richness on the Tylenchidae (nematodes) was the only non-additive effect of litter mixing. Together, these data demonstrate an effect of plant community composition on decomposer biomass, abundance, and diversity, confirming a link between above and belowground communities. We were able to identify the species to which the decomposer community is most sensitive, aiding predictions of the consequences of the loss of these dominant species on the decomposer community, with potential feedbacks for organic matter and nutrient turnover.

Among‐ and within‐species variation in plant litter decomposition in contrasting long‐term chronosequences

1. Following major disturbances ecosystem development occurs but in the prolonged absence of disturbance a decline (retrogressive) phase follows in which productivity and nutrient availability diminishes. Although it is recognized that litter quality and decomposition rates decrease as retrogression proceeds, little is known about the extent to which this is driven among- vs. within-species variation across these sequences. 2. We selected six long-term chronosequences that each included retrogressive stages, in New Zealand, Hawaii, Sweden, Alaska and Australia. Two involve significant species turnover across the sequence so that different species dominate at different stages, two involve low species turnover so that the same dominant species occur at all stages, and two involve some turnover of species but with certain species persisting throughout most of the sequence. 3. For each chronosequence, we collected litter from each dominant plant species at each stage of that sequence. For each litter collection we measured concentrations of N and P, and performed laboratory decomposition bioassays to measure mass loss, N and P loss, and the response of mass loss to mixture with litters of coexisting species. 4. We found that litter N and P concentrations often declined with increasing ecosystem age, both among- and within-species. However, the relative importance of among- and within-species effects varied across the six chronosequences. Rates of litter mass, N, and P loss during decomposition sometimes decreased with increasing ecosystem age, but most often at the among-species rather than the within-species level. 5. Litter mixing effects often varied across chronosequence stages, but the magnitude and direction of these effects was inconsistent among sequences. Variation in litter mixing effects across chronosequence stages was driven mainly by among- rather than within-species variation. 6. Although several recent studies have emphasized the role of within-species variation on ecosystem properties, our results point to among-species variation as a consistently important ecological driver, with within-species variation being important only for some variables and in some instances. As such they highlight that decomposition processes are most likely to be highly responsive to gradients of soil fertility (such as across chronosequences) when significant species turnover occurs across the gradient.

Context dependency of litter‐mixing effects on decomposition and nutrient release across a long‐term chronosequence

Litter decomposition is an important driver of terrestrial systems, and factors that determine decomposition rate for individual litter species have been widely studied. Fewer studies have explored the factors that regulate how mixing litters of multiple species affects litter decomposition and nutrient dynamics, and only a handful of studies have investigated how litter‐mixing effects may differ among different habitats or ecosystems, or how they respond to environmental gradients. We used a well‐established retrogressive chronosequence involving thirty lake islands in northern Sweden in which time since fire disturbance increases with decreasing island size; smaller islands therefore have reduced rates of aboveground and belowground ecosystem processes. On each of these islands we utilized plots with and without the long‐term experimental removal of shrubs. Litters from the six most common plant species on the islands were prepared in single‐, three‐ and six‐species litterbags, and placed on both the shrub‐removal and non‐removal plots on each island to decompose for one year. We found significant non‐additive effects of litter mixing on litter decomposition rates, on final litter N and P concentrations, and on litter N loss, but these non‐additive effects varied both in direction and magnitude with changed number of species, and even among litter mixtures with the same number of species. Further, the magnitude of non‐additive effects of litter mixing on both litter decomposition and nutrient dynamics was significantly influenced by both island size and the interaction between island size and shrub‐removal treatment. When shrubs were present, there was a U‐shaped relationship between these non‐additive effects and island size, while the relationship was positive when shrubs were removed. Hence, our results support previous findings that litter mixing may produce non‐additive effects on litter decomposition and nutrient dynamics, and that these effects tend to be idiosyncratic due to the importance of effects of individual species in the mixture. Most importantly, our results show that non‐additive litter‐mixing effects change greatly across environmental gradients, meaning that the biotic and abiotic characteristics of an ecosystem can be a powerful driver of the magnitude and even the direction of litter‐mixing effects on ecosystem processes.

Nitrogen and Phosphorus Release from Mixed Litter Layers is Lower than Predicted from Single Species Decay

Ecosystem-level nutrient dynamics during decomposition are often estimated from litter monocultures. If species effects are additive, we can statistically predict nutrient dynamics in multi-species systems from monoculture work, and potential consequences of species loss. However, if species effects are dependent on interactions with other litter species (that is, non-additive), predictions based on monoculture data will likely be inaccurate. We conducted a 3-year, full-factorial, mixed-litter decomposition study of four dominant tree species in a temperate forest and measured nitrogen and phosphorus dynamics to explore whether nutrient dynamics in mixtures were additive or non-additive. Following common approaches, we used litterfall data to predict nutrient dynamics at the ecosystem-level. In mixtures, we observed non-additive effects of litter mixing on nutrient dynamics: the presence of nutrient-rich species in mixture facilitated nutrient release, whereas nutrient-poor species facilitated nutrient retention. Fewer nutrients were released from mixtures containing high-quality litter, and more immobilized from mixtures containing low-quality litter, than predicted from monocultures, creating a difference in overall nutrient release between predicted and actual dynamics in litter mixtures. Nutrient release at the ecosystem-level was greatly overestimated when based on monocultures because the effect of species interactions on nutrient immobilization was not accounted for. Our data illustrate that the identity of species in mixtures is key to their role in non-additive interactions, with repercussions for mineral nutrient availability and storage. These results suggest that predictions of ecosystem-level nutrient dynamics using litter monoculture data likely do not accurately represent actual dynamics because the effects of litter species interactions are not incorporated.

Context dependency of litter-mixing effects on decomposition and nutrient release across a long-term chronosequence

Context dependency of litter-mixing effects on decomposition and nutrient release across a long-term chronosequence

Effect of litter quality on its decomposition in broadleaf and coniferous forest

Recently there has been much interest in the effect of litter mixing as well as the effect of different forest habitats on the decomposition process. Our aim was to test two hypotheses: high quality litter promotes decomposition of poor quality litter, and litter decomposes faster in broadleaf than in coniferous forest. We conducted a litter mixing experiment using litterbags placed in two forest floors, in which treatments consisted of litter monocultures of each of two campy species ( Castanopsis eyrei and Pinus massoniana), as well as mixtures of these two species. The results showed that C. eyrei leaves decomposed significantly faster in the coniferous habitat than in their native habitat. On the other hand, P. massoniana needles decomposed significantly faster in their native coniferous habitat than in the broadleaf habitat. In our experiment we found that the mixture had different effect on different quality litter. P. massoniana needles (poor quality) had a positive effect on the decomposition of C. eyrei leaves (high quality), while C. eyrei leaves had a negative effect on the decomposition of P. massoniana needles in the mixture case in both broadleaf and coniferous habitats. The diversity of the fungi identified from different litters varied among treatments and the mass loss was positively correlated with the Shannon–Weaver diversity index of fungi. It is suggested that fungi may be one of the major drivers to control the decomposition process.

Functional diversity affects decomposition processes in experimental grasslands

Summary 1. A number of biodiversity experiments have shown that plant diversity plays a significant role for ecosystem functioning. However, diversity effects on processes involving multi-trophic interactions such as litter decomposition are rather rare. In these experiments, plant diversity is usually categorized into taxonomic units or functional groups. Continuous measures of functional diversity that are based on measurable traits, in contrast, may be a more flexible way to analyse the functional significance of biodiversity. 2. Litter decomposition is a key process in ecosystem biogeochemistry. To understand the consequences of altered biodiversity for ecosystem functioning, it is thus crucial to quantify any potential diversity effects on decomposition processes. 3. I performed several complementary decomposition trials within the BIODiversity and Ecological Processes in Terrestrial Herbaceous ecosystems (BIODEPTH) experiment, which established a gradient of plant species richness and number of functional groups. I hypothesized that decomposi tion rates increase with increasing plant diversity due to non-additive litter mixing effects and more favourable microenvironmental conditions. 4. Decomposition rates of both standard materials and community-specific litter increased with the number of functional groups and with a continuous measure of functional diversity. Species richness, in contrast, had no or rather small positive effects on decomposition. Presence of nitrogen fixing legumes strongly enhanced decomposition, via effects on both litter quality and on the decomposition microenvironment. 5. The predominance of functional diversity rather than of species richness effects suggests that profound knowledge on functional attributes of plant species is the key to understand and to predict biodiversity-decomposition relationships.

Consequences of non‐random species loss for decomposition dynamics: experimental evidence for additive and non‐additive effects

Summary Although litter decomposition is a fundamental ecological process, most of our understanding comes from studies of single‐species decay. Recently, litter‐mixing studies have tested whether monoculture data can be applied to mixed‐litter systems. These studies have mainly attempted to detect non‐additive effects of litter mixing, which address potential consequences of random species loss – the focus is not on which species are lost, but the decline in diversity per se. Under global change, species loss is likely to be non‐random, with some species more vulnerable to extinction than others. Under such scenarios, the effects of individual species (additivity) as well as of species interactions (non‐additivity) on decomposition rates are of interest. To examine potential impacts of non‐random species loss on ecosystems, we studied additive and non‐additive effects of litter mixing on decomposition. A full‐factorial litterbag experiment was conducted using four deciduous leaf species, from which mass loss and nitrogen content were measured. Data were analysed using a statistical approach that first looks for additive identity effects based on the presence or absence of species and then significant species interactions occurring beyond those. It partitions non‐additive effects into those caused by richness and/or composition. This approach addresses questions key to understanding the potential effects of species loss on ecosystem processes. If additive effects dominate, the consequences for decomposition dynamics will be predictable based on our knowledge of individual species, but not statistically predictable if non‐additive effects dominate. We found additive (identity) effects on mass loss and non‐additive (composition) effects on litter nitrogen dynamics, suggesting that non‐random species loss could significantly affect this system. We were able to identify the species responsible for effects that would otherwise have been considered idiosyncratic or absent when analysed by the methods used in previous work. Synthesis. We observed both additive and non‐additive effects of litter‐mixing on decomposition, indicating consequences of non‐random species loss. To predict the consequences of global change for ecosystem functioning, studies should examine the effects of both random and non‐random species loss, which will help identify the mechanisms that influence the response of ecosystems to environmental change.