Composition of speciose leaf litter alters stream detritivore growth, feeding activity and leaf breakdown

Leaf litter derived from riparian trees can control secondary production of detritivores in forested streams. Species-rich assemblages of leaf litter reflect riparian plant species richness and represent a heterogeneous resource for stream consumers. Such variation in resource quality may alter consumer growth and thus the feedback on leaf breakdown rate via changes in feeding activity. To assess the consequences of this type of resource heterogeneity on both consumer growth and subsequent litter breakdown, we performed a laboratory experiment where we offered a leaf-shredding stream detritivore (the stonefly Tallaperla maria, Peltoperlidae) ten treatments of either single- or mixed-species leaf litter. We measured consumer growth rate, breakdown rate and feeding activity both with and without consumers for each treatment and showed that all three variables responded to speciose leaf litter. However, the number of leaf species was not responsible for these results, but leaf species composition explained the apparent non-additive effects. T. maria growth responded both positively and negatively to litter composition, and growth on mixed-litter could not always be predicted by averaging estimates of growth in single-species treatments. Furthermore, breakdown and feeding rates in mixed litter treatments could not always be predicted from estimates of single-species rates. Given that species richness and composition of senesced leaves in streams reflects riparian plant species richness, in-stream secondary production of detritivores and organic matter dynamics may be related to species loss of trees in the riparian zone. Loss of key species may be more critical to maintaining such processes than species richness per se.     

Detritivore stoichiometric diversity alters litter processing efficiency in a freshwater ecosystem

Many studies have estimated relationships between biodiversity and ecosystem functioning, and observed generally positive effects. Because detritus is a major food resource in stream ecosystems, decomposition of leaf litter is an important ecosystem process and many studies report the full range of positive, negative and no effects of diversity on decomposition. However, the mechanisms underlying decomposition processes in fresh water remain poorly understood. Organism body stoichiometry relates to consumption rates and tendencies, and decomposition processes of litter may therefore be affected by diversity in detritivore body stoichiometry. We predicted that the stoichiometric diversity of detritivores (differences in C: nutrient ratios among species) would increase the litter processing efficiency (litter mass loss per total capita metabolic capacity) in fresh water through complementation regarding different nutrient requirements. To test this prediction, we conducted a microcosm experiment wherein we manipulated the stoichiometric diversity of detritivores and quantified mass loss of leaf litter mixtures. We compared litter processing efficiency among litter species in each microcosm with single species detritivores, and observed detritivores with nutrient‐rich bodies tended to prefer litter with lower C: nutrient ratios over litter with higher C: nutrient ratios. Furthermore, litter processing efficiencies were significantly higher in the microcosms containing species of detritivores with both nutrient‐rich and ‐poor bodies than microcosms containing species of detritivores including only nutrient‐rich or ‐poor bodies. This might mean a higher stoichiometric diversity of detritivores increased litter processing efficiency. Our results suggest that ecological stoichiometry may improve understanding of links between biodiversity and ecosystem function in freshwater ecosystems.     

Interactive responses of grass litter decomposition to warming,nitrogen addition and detritivore access in a temperate old field

Plant litter decomposition has been studied extensively in the context of both climate warming and increased atmospheric N deposition. However, much of this research is based on microbial responses, despite the potential for detritivores to contribute substantially to litter breakdown. We measured litter mass-loss responses to the combined effects of warming, N addition and detritivore access in a grass-dominated old field. We concurrently assessed the roles of litter treatment origin vs. microenvironment (direct warming and N-addition effects) to elucidate the mechanisms through which these factors affect decomposition. After 6 weeks, mass loss increased in N-addition plots, and it increased with detritivore access in the absence of warming. After 1 year, warming, N addition, and detritivore access all increased litter mass loss, although the effects of N addition and warming were non-additive in the detritivore-access plots. For the litter-origin experiment, mass loss after 6 weeks increased in litter from N-addition plots and warmed plots, but unlike the overall decomposition response, the N-addition effect was enhanced by detritivore access. Conversely, for the microenvironment experiment, detritivore access only increased mass loss in unfertilized plots. After 1 year, detritivore access increased mass loss in the litter-origin and microenvironment experiments, but there were no warming or N-addition effects. Overall, our results provide support for a substantial role of detritivores in promoting litter mass loss in our system. Moreover, they reveal important interactions between litter origin, microclimate and detritivores in determining decomposition responses to global change.     

Stoichiometric imbalances between detritus and detritivores are related to shifts in ecosystem functioning

How are resource consumption and growth rates of litter‐consuming detritivores affected by imbalances between consumer and litter C:N:P ratios? To address this question, we offered leaf litter as food to three aquatic detritivore species, which represent a gradient of increasing body N:P ratios: a crustacean, a caddisfly and a stonefly. The detritivores were placed in microcosms and submerged in a natural stream. Four contrasting leaf species were offered, both singly and in two‐species mixtures, to obtain different levels of stoichiometric imbalance between the resources and their consumers. The results suggest that detritivore growth was constrained by N rather than C or P, even though 1) the N:P ratios of the consumers’ body tissue was relatively low and 2) microbial leaf conditioning during the experiment reduced the N:P imbalance between detritivores and leaf litter. This surprisingly consistent N limitation may be a consequence of cumulative N‐demand arising from the production of N‐rich chitin in the exoskeletons of all three consumer species, which is lost during regular moults, in addition to N‐demand for silk production by the caddisfly. These N requirements are not commonly quantified in stoichiometric analyses of arthropod consumers. There was no evidence for compensatory feeding, but when offered mixed‐species litter varying in C:N:P ratios, detritivores consumed more of the litter species showing the highest N:P and lowest C:N ratio, accelerating the mass loss of the preferred leaf species in the litter mixture. These results show that imbalances in consumer–resource stoichiometry can have contrasting effects on coupled processes, highlighting a challenge in developing a mechanistic understanding of the role of stoichiometry in regulating ecosystem processes such as leaf litter decomposition.     

Interactions between detritivores and microfungi during the leaf detritus decomposition in a volcanic lake (Lake Vico – central Italy)

The role of biota in the mass loss of Phragmites australis (Cav.) Trin. ex Steud was studied in the littoral belt of a central Italy volcanic lake. The research focussed on the feeding interactions between detritivores and decomposing fungi as drivers of the leaf litter decomposition. The litterbag technique was used to assess the leaf mass loss, the number of colonizing fungi and the patterns of leaf colonization by detritivores during 40 days of submersion in 16 sampling sites. Cores of bottom sediment were collected to estimate the organic content and ergosterol concentration as measure of fungal mass. The rate of leaf mass loss showed significant variability among the sampling sites and was non-linearly related to the quantity of organic depositions onto the lake bottom, peaking at about 40% of the dry matter. The rate was also positively correlated with the density of detritivore mass relative to the leaf unit mass, which increased with time. On the 20th day of litterbag immersion, when 40% of the initial leaf litter remained, we observed the best accordance between the two measures as well as the lowest difference in the detritivore mass density among sampling sites. In the absence of animals, the decomposition rate was positively related to the number of fungi on the decaying litter. The feeding activity of detritivores changed both the species richness and composition of the fungal community on the litter. The substrate reduction due to intense animal feeding appeared to limit the ability of fungi to regrow after grazing. As a result, an inverse relationship between the number of fungi and the decomposition rate was observed.     

Effects of macro-detritivores density on leaf detritus processing rate: a macrocosm experiment

The effect of macroinvertebrate detritivore density on the mass loss rates of leaf litter of Alnus glutinosa (alder) was assessed. Experimental freshwater macrocosms, with increasing densities of four species of macroinvertebrate detritivores belonging to two functional groups (shredders and scrapers), were set up outdoors. The litter bag technique was used to assess decomposition rates of alder leaves. Indirect effects of increasing density of macroinvertebrates on phytoplankton standing crop in the water column were investigated by analysing Chlorophyll a concentration. Decomposition rate increased as animal density increased, although a continuous increase in detritivores density resulted in a discrete, step-wise increase of the decomposition rates. Animal colonisation followed an exponential pattern in low-medium density treatments versus a typical `bell-shape' curve in high density treatments; animals started to leave the consumed patches when about 60% of the initial leaf mass was lost (35th day in high-density treatments). Diversity (Hs) of the simplified detritivore community decreased as decomposition proceeded, with a dominance of shredders during the last phase of decomposition. Faster decomposition rate of detritus in the benthic compartment lead to a higher microalgae standing crop in the water column emphasising the role of allochthonous detritus as a source of nutrients for algae primary production in coastal freshwater ecotones.     

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.     

Species diversity and decomposition in laboratory aquatic systems: the role of species interactions

1. Interest in the effects of biodiversity on ecosystem processes is increasing, stimulated by the global species decline. Different hypotheses about the biodiversity‐ecosystem functioning (BEF) relationship have been put forward and various underlying mechanisms proposed for different ecosystems. 2. We investigated BEF relationships and the role of species interactions in laboratory experiments focussing on aquatic decomposition. Species richness at three different trophic levels (leaf detritus, detritus‐colonising fungi and invertebrate detritivores) was manipulated, and its effects on leaf mass loss and fungal growth were assessed in two experiments. In the first, monocultures and mixtures of reed (Phragmites australis), alder (Alnus glutinosa) and oak (Quercus cerris) leaf disks were incubated with zero, one or eight fungal species. Leaf mixtures were also incubated with combinations of three and five fungal species. In the second experiment, reed leaf disks were incubated with all eight fungal species and offered to combinations of one, two, three, four or five macroinvertebrate detritivores with different feeding modes. 3. Results from the first experiment showed that leaf mass loss was directly related to fungal mass and varied unimodally with the number of fungi, with a maximum rate attained at intermediate diversity in oak and reed and at maximum diversity in alder (the fastest decomposing leaf). 4. Mixing litter species stimulated fungal growth but interactions between species of fungi slowed down decomposition. In contrast, mixtures of macroinvertebrate detritivores reduced fungal mass and accelerated leaf decomposition. Possible explanations of the positive relationship between detritivore diversity and decomposition are a reduction in fungal dominance and a differentiation in the use of different resource patches promoted by higher fungal diversity. 5. In conclusion, the results show a general increase in decomposition rate with increasing biodiversity that is controlled by within‐ and between‐trophic level interactions, and support the hypothesis of both bottom‐up and top‐down effects of diversity on this process.     

Litter composition rather than plant presence affects decomposition of tropical litter mixtures

Litter decomposition is strongly controlled by litter quality, but the composition of litter mixtures and potential interactions with live plants through root activity may also influence decomposers. In a greenhouse experiment in French Guiana we studied the combined effects of the presence of tropical tree seedlings and of distinct litter composition on mass and nitrogen (N) loss from decomposing litter and on microbial biomass. Different litter mixtures decomposed for 435 days in pots filled with sand and containing an individual seedling from one of four different tree species. We found both additive and negative non-additive effects (NAE) of litter mixing on mass loss, whereas N loss showed negative and positive NAE of litter mixing. If litter from the two tree species, Platonia insignis and Goupia glabra were present, litter mixtures showed more positive and more negative NAE on N loss, respectively. Overall, decomposition, and in particular non-additive effects, were only weakly affected by the presence of tree seedlings. Litter mass loss weakly yet significantly decreased with increasing fine root biomass in presence of Goupia seedlings, but not in the presence of seedlings of any other tree species. Our results showed strong litter composition effects and also clear, mostly negative, non-additive effects on mass loss and N loss. Species identity of tree seedlings can modify litter decomposition, but these live plant effects remain quantitatively inferior to litter composition effects.     

Facilitation and predation structure a grassland detrital food web: the responses of soil nematodes to isopod processing of litter

1. Detritus can support successive consumers, whose interactions may be structured by changes in the condition of their shared resource. One model of such species interactions is a processing chain, in which consumers feeding on the resource in a less processed state change the resource condition for subsequent consumers. 2. In a series of experiments, the hypothesis was tested that a common detritivore, the terrestrial isopod Porcellio scaber, affects soil nematodes through the processing of plant litter. Different detrital resources were added to soil from a California coastal prairie in order to simulate litter processing by the detritivore. Treatments that included only whole grass litter corresponded to detrital food webs lacking detritivores, while treatments that included mixtures of P. scaber faeces and grass litter corresponded to different densities or feeding rates of P. scaber. 3. Simulated litter processing by P. scaber increased the abundance of bacterivorous nematodes by between 32% and 202% after 24-44 days in laboratory experiments, but had no effect on fungivorous or predaceous nematodes. 4. In a subsequent field experiment, however, fungivorous nematodes were suppressed by isopod litter processing while bacterivores showed no response. Instead, P. scaber processing of litter increased the abundance of predaceous nematodes in the field experiment by 176%. 5. When simulated litter processing of litter was crossed in laboratory experiments with predaceous nematode addition (comparable to the response of predators in the field experiment), the abundance of bacterivores was increased by isopod processing of litter (by an average of 122%), but suppressed by elevated densities of predaceous nematodes (by an average of 41%). 6. This suggests that litter processing by P. scaber facilitates the bacterial channel of the soil food web, but that predaceous nematodes suppress the response of bacterivores in the field. Processing chain interactions may, therefore, be important in understanding the relative importance of bacterial and fungal channels in the soil food web, while top-down effects of predators determine the resulting changes in population abundance and biomass.     

Positive indirect effect of tadpoles on a detritivore through nutrient regeneration

In aquatic food webs consumers can affect other members of the web by releasing nutrients as a result of their feeding activity. There is increasing evidence of these positive effects on primary producers, but such nutrient regeneration can also affect detritivores, by favoring the activities of detritus-associated microbes. We examined the effects of nutrient regeneration by tadpoles on leaf-eating detritivores under laboratory conditions. We fed four species of tadpoles three different food items (leaf litter, algae, and sludgeworms). We then conditioned terrestrial dead leaves with water from reared tadpoles (treatments) or food items alone (controls), and compared the C:N ratios of the conditioned leaves and the growth of the isopod Asellus hilgendorfii fed on the conditioned leaves. Tadpole feeding activity reduced the C:N ratio of conditioned leaves, and the effect was greatest when tadpoles were fed algae. Isopod growth rates were often higher when they were fed the litter conditioned with water from reared tadpoles. Thus, nutrient regeneration by tadpoles had a positive indirect effect on detritivores by enhancing leaf quality. Tadpoles often occur in nutrient-limited habitats where leaf litter is the major energy source, and their facilitative effects on leaf-eating detritivores may be of great significance in food webs by enhancing litter decomposition.     

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

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Species interactions between estuarine detritivores: inhibition or facilitation?

Native Hawaiian estuarine detritivores; the prawn Macrobrachium grandimanus, and the neritid gastropod Neritina vespertina, were maintained in flow-through microcosms with conditioned leaves from two riparian tree species, Hau (Hibiscus tiliaceus) and guava (Psidium guajava). Their ability to beak down leaf detritus was determined when alone and when they were together. In single-species treatments, N. vespertina processed leaves from both trees at higher rates than M. grandimanus, but in combined treatments, facilitation occurred when the substrate consisted of Hau leaves, and interference occurred when the substrate consisted of guava leaves. From this, we conclude that whether detritivore species are functionally redundant, facilitating or inhibiting in their processing of detritus depends not only on the detritivore species, but also on the species composition of the detritus food source.     

Top‐down and bottom‐up control of litter decomposers in streams

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Functional dissimilarity across trophic levels as a driver of soil processes in a Mediterranean decomposer system exposed to two moisture levels

The role of biodiversity for soil processes remains poorly understood. Existing evidence suggests that functional diversity rather than species richness is relevant for soil functioning. However, the importance of functional diversity has rarely been assessed simultaneously at more than one trophic level, critically limiting the prediction of consequences of biodiversity loss for soil functioning. In a laboratory microcosm experiment, we tested the hypothesis that increasing functional dissimilarity of both litter‐feeding soil fauna and litter mixtures interactively affects the rates of five different soil processes related to litter decomposition. We created trait‐based functional dissimilarity gradients using five assemblages of two detritivore species and five mixtures of two plant litter species commonly found in Mediterranean shrubland ecosystems of southern France. With increasing drought periods predicted for Mediterranean ecosystems in the future, we additionally included two different watering frequencies to evaluate the impact of drought on soil processes and how drought interacts with functional dissimilarity. The different fauna assemblages and litter mixtures showed strong effects on litter mass loss, soil organic carbon and nitrogen leaching, as well as on soil microbial activities. Up to 20% of the variation in response variables was explained by functional dissimilarity, suggesting an ecologically relevant impact of functional diversity on soil process rates. Detritivore functional dissimilarity tended to have stronger effects when combined with increasingly dissimilar litter mixtures, suggesting that trait dissimilarity interacts across trophic levels. Drought affected several soil processes but did not modify the relationships between functional dissimilarity and process rates. Our results indicate that trait diversity of detritivore assemblages and litter mixtures is an important predictor of soil process rates. The common and easily measurable traits used in our study suggest straightforward application across different types of ecosystems and environmental conditions.     

Decomposition of spruce litter needles of different quality by Setulipes androsaceus and Thysanophora penicillioides

Different components of functional biodiversity, such as functional type richness and composition, have been reported to affect the decomposition of litter mixtures. In spite of the numerous reports of these effects, mechanisms underlying patterns of decomposition in litter mixtures are still unclear. We analyzed whether mixture decomposition was affected by: (a) the number of species in the mixture (mixture richness); and (b) the mixture’s functional composition (% of fast- vs. slow-decomposing species included in the mixture). We then tested if variation between observed and expected values of decomposition in mixtures was associated to: (c) the initial litter characteristics of the component species (initial nitrogen, lignin, cellulose and hemicellulose content of litters); and (d) the chemical heterogeneity of the mixtures (variation in the same chemical compounds between the components in each mixture). When up to 5 species representing different functional types were included, both species richness and functional composition showed statistically significant non-additive, and in general positive, effects on litter mixture decomposition. The positive effect of mixture richness on decomposition did not disappear, but was much less marked, when considering mixture with slow-decomposing species only. Although the main driver of decomposition in a mixture is still the average decomposability of the component species (itself largely determined by litter quality), the species interactions in a mixture add a consistent source of variability that is worth considering when predicting the decomposability of a given mixture. We showed that a greater positive difference between observed decomposition rates and that expected from its component species alone was found in mixtures with higher mean nitrogen content and a higher heterogeneity in non-labile compounds. Our results offer quantitative proof that litter chemical heterogeneity, as well as the mean quality of the mixture, can affect the decomposability in litter mixtures.     

The effects of leaf litter evenness on decomposition depend on which plant functional group is dominant

Background and aims

Climbing plants are increasing in dominance in the subtropical forests of South China and other areas around the world, altering patterns of plant dominance and evenness in community. We investigated how changes in species’ identity and patterns of leaf litter evenness affected decomposition of litter mixtures.

Methods

We used litter-bag method to study the influence of different relative abundance mixtures (75 % : 25 %; 50 % : 50 %; 25 % : 75 %) of plant litter from two functional groups (climbing plants and trees) on decomposition rates in a subtropical forest in Guangdong, China.

Results

We found negative non-additive effects of mixing litter overall and species composition affected decomposition rates the most. In addition, when climbing plants were dominant, even mixtures decomposed slower significantly than uneven mixtures. Evenness did not affect decomposition rates, however, when trees were dominant. The magnitude of antagonistic effects increased with increasing dominance of climbing plants but decreased with time, suggesting a strong negative feedback between litter proportion of climbing plant and decomposition rates at the initial stage.

Conclusion

The evenness in leaf litter composition affects rates of decomposition, but these effects depend on which plant functional group is dominant. Thus, we should pay more attention to shifts in identity of dominant species and patterns of community evenness.     

Effects of nickel hyperaccumulation in Alyssum pintodasilvae on model arthropods representatives of two trophic levels

An experimental assessment of the defence hypothesis of nickel (Ni) hyperaccumulation in Alyssum was lacking. Also, to date no study had investigated the effects of hyperaccumulator litter on a detritivore species. We performed several experiments with model arthropods representatives of two trophic levels: Tribolium castaneum (herbivore) and Porcellio dilatatus (detritivore). In no-choice trials using artificial food disks with different Ni concentrations, T. castaneum fed significantly less as Ni concentration increased and totally rejected disks with the highest Ni concentration. In choice tests, insects preferred disks without Ni. In the no-choice experiment, mortality was low and did not differ significantly among treatments. Hence, this suggested a deterrent effect of high Ni diet. Experiments with P. dilatatus showed that isopods fed A. pintodasilvae litter showed significantly greater mortality (83%) than isopods fed litter from the non-hyperaccumulator species Iberis procumbens (8%), Micromeria juliana (no mortality) or Alnus glutinosa (no mortality). Also, isopods consumed significantly greater amounts of litter from the non-hyperaccumulator plant species. The behaviour of isopods fed A. pintodasilvae litter suggested an antifeedant effect of Ni, possibly due to post-ingestive toxic effects. Our results support the view that Ni defends the Portuguese serpentine hyperaccumulator A. pintodasilvae against herbivores, indicating that Ni can account both for feeding deterrence and toxic effects. The effects of hyperaccumulator litter on the detritivore P. dilatatus suggest that the activity of these important organisms may be significantly impaired with potential consequences on the decomposition processes.     

Influence of Changing Plant Food Sources on the Gut Microbiota of Saltmarsh Detritivores

Changes in agricultural land-use of saltmarshes along the German North Sea coast have favoured the succession of the marsh grass Elytrigia atherica over the long-established Spartina anglica. Consequently, E. atherica represents a potential food source of increasing importance for plant-feeding soil detritivores. Considering the importance of this ecological guild for decomposition processes and nutrient cycling, we focussed on two sympatric saltmarsh soil macrodetritivores and their associated gut microbiota to investigate how the digestive processes of these species may be affected by changing plant food sources. Using genetic fingerprints of partial 16S rRNA gene sequences, we analysed composition and diversity of the bacterial gut community in a diplopod and an amphipod crustacean in relation to different feeding regimes representing the natural vegetation changes. Effects of syntopy on the host-specific gut microbiota were also taken into account by feeding the two detritivore species either independently or on the same plant sample. Bacterial community composition was influenced by both the host species and the available plant food sources, but the latter had a stronger effect on microbial community structure. Furthermore, bacterial diversity was highest after feeding on a mixture of both plant species, regardless of the host species. The gut microbiota of these two detritivores can thus be expected to change along with the on-going succession at the plant community level in this environment. Cloning and sequencing of bacterial 16S rRNA gene fragments further indicated a host-related effect since the two detritivores differed in terms of predominant bacterial taxa: diplopods harboured mainly representatives of the phyla Bacteroidetes and Gammaproteobacteria. In contrast, the genus Vibrio was found for the amphipod host across all feeding conditions.     

Non-additive effects of litter mixtures on decomposition of leaf litters in a Mediterranean maquis

Many studies across a range of ecosystems have shown that decomposition in mixed litter is not predictable from single-species results due to synergistic or antagonistic interactions. Some studies also reveal that species composition and relative abundance may be more important than just richness in driving non-additive effects. Most studies on litter decomposition in Mediterranean maquis, an high-diversity shrubby ecosystem, have dealt exclusively with single species. In this study we investigated, at the individual-litter level, as well as at the litter-mixture level, the effect of litter mixing on decomposition of 3-species litter assemblages with different relative abundance of the component litters; we set up two types of litter assemblages that reflected the heterogeneity of bush cover in the inner maquis and at the edge maquis/gaps, as related to the leaf traits, i.e. sclerophylly vs mesophylly. We measured mass loss, decay of lignin, cellulose and ADSS (acid detergent soluble substances) and fungal mycelium ingrowth. The results show that over a 403-day incubation period, the decomposition of individual litters in mixtures deviated from that of monospecific litters and had different directions. In litter mixtures of the sclerophylls Phillyrea angustifolia and Pistacea lentiscus with the mesophyll Cistus, decomposition was lower than expected (antagonistic effect); in the mixtures of litters with similar physical structure (Ph. angustifolia and P. lentiscus with Quercus ilex) decomposition was faster than expected (synergistic effect). When considering the different decomposition phases, both negative and positive effects occurred in Quercus mixtures depending on the phase of decomposition. In both types of 3-species litter assemblages the greatest effect occurred in uneven mixtures rather than in even mixtures. Our results show that species composition drives the direction whilst the decomposability and the relative abundance drive the magnitude of non-additive effects of litter mixing on decomposition.