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.     

Biodiversity and tallgrass prairie decomposition: the relative importance of species identity, evenness, richness, and micro-topography

Biodiversity has been declining in many areas, and there is great interest in determining whether this decline affects ecosystem functioning. Most biodiversity—ecosystem functioning studies have focused on the effects of species richness on net primary productivity. However, biodiversity encompasses both species richness and evenness, ecosystem functioning includes other important processes such as decomposition, and the effects of richness on ecosystem functioning may change at different levels of evenness. Here, we present two experiments on the effects of litter species evenness and richness on litter decomposition. In the first experiment, we varied the species evenness (three levels), identity of the dominant species (three species), and micro-topographic position (low points [gilgais] or high points between gilgais) of litter in three-species mixtures in a prairie in Texas, USA. In a second experiment, we varied the species evenness (three levels), richness (one, two, or four species per bag), and composition (random draws) of litter in a prairie in Iowa, USA. Greater species evenness significantly increased decomposition, but this effect was dependent on the environmental context. Higher evenness increased decomposition rates only under conditions of higher water availability (in gilgais in the first experiment) or during the earliest stages of decomposition (second experiment). Species richness had no significant effect on decomposition, nor did it interact with evenness. Micro-topographic position and species identity and composition had larger effects on decomposition than species evenness. These results suggest that the effects of litter species diversity on decomposition are more likely to be manifested through the evenness component of diversity than the richness component, and that diversity effects are likely to be environmentally context dependent.     

Does initial litter chemistry explain litter mixture effects on decomposition?

Abstract We tested the hypothesis that interactions in litter mixtures (expressed as the difference between observed and expected decomposition rates) are greater when the component species differ more in their initial litter chemistry. Thereto, we collected freshly senesced leaf litter from a wide range of species from an old field and woodland vegetation, and a fen ecosystem in The Netherlands. Litterbags with either mono-specific litter (20 and 15 species), or litter mixtures (50 and 42 species pairs) of randomly drawn combinations of two representatives from different plant functional types were incubated for 20, 35 and 54 weeks in a purpose-built decomposition bed (woodland/old field) or in the field (fen). Species showed a wide range of decomposition rates. For the woodland/old field species, initial litter C and P concentrations were significantly correlated with litter decomposition rate. For the fen species, litter phenolics concentration was correlated with decomposition rate. If the Sphagnum species were left out of the analyses, initial litter P and phenolics concentration both showed a significant relationship, albeit only with the remaining mass after 1 year. Differences between observed and expected decomposition were often considerable in individual litter mixtures. Regression analysis showed that such differences were not related to the differences in litter chemistry of the component species. Furthermore, litter mixtures containing species with very different initial litter chemistry did not show stronger interaction when tested against litter mixtures containing chemically similar litter types. From these observations we conclude that the difference in initial single litter chemistry parameters of the component is not a useful concept to explain interactions in litter mixtures.Due to an error in the citation line, this revised PDF (published in December 2003) deviates from the printed version, and is the correct and authoritative version of the paper.     

Influence of evenness on the litter-species richness-decomposition relationship in Mediterranean grasslands

Aims Human impacts on natural ecosystems induce changes in their functioning through alterations in species richness, composition and evenness of plant communities. Most litter diversity–decomposition processes studies have only manipulated species richness, ignoring the role of evenness. Here, results from a field litterbag experiment are presented to test whether changes in evenness of species distribution in litter mixtures affected the strength of the litter-species richness–decomposition relationship.Methods Ten herbaceous species abundant in Mediterranean grassland communities and representative of different genera and functional groups were used. Species richness was directly manipulated to produce litter mixtures of three and six plant species, as well as litter of each individual species used. Each level of species richness was replicated several times such that each repeat had a different species composition. Three- and six-species litter mixtures were also treated to vary in evenness (three levels). Decomposition rate was assessed by percentage dry weight loss over the 90 days of the experiment.Important findings Decomposition rate was positively related to the linear increase in litter-species richness and was affected by the composition of the litter-species mixture. Decomposition rates differed significantly between evenness treatments and moreover, the strength of the positive relationship between litter-species richness and decomposition rate decreased notably in the low-evenness treatment. The effects of evenness on decomposition rate, at different richness levels, were partially explained by the differences in the initial litter mixture's carbon-to-nitrogen ratio within them. This study reveals that short-term decomposition rate is positively affected by both components of Mediterranean grassland litter-species diversity.     

Long-term presence of tree species but not chemical diversity affect litter mixture effects on decomposition in a neotropical rainforest

Plant litter diversity effects on decomposition rates are frequently reported, but with a strong bias towards temperate ecosystems. Altered decomposition and nutrient recycling with changing litter diversity may be particularly important in tree species-rich tropical rainforests on nutrient-poor soils. Using 28 different mixtures of leaf litter from 16 Amazonian rainforest tree species, we tested the hypothesis that litter mixture effects on decomposition increase with increasing functional litter diversity. Litter mixtures and all single litter species were exposed in the field for 9 months using custom-made microcosms with soil fauna access. In order to test the hypothesis that the long-term presence of tree species contributing to the litter mixtures increases mixture effects on decomposition, microcosms were installed in a plantation at sites including the respective tree species composition and in a nearby natural forest where these tree species are absent. We found that mixture decomposition deviated from predictions based on single species, with predominantly synergistic effects. Functional litter diversity, defined as either richness, evenness, or divergence based on a wide range of chemical traits, did not explain the observed litter mixture effects. However, synergistic effects in litter mixtures increased with the long-term presence of tree species contributing to these mixtures as the home field advantage hypothesis assumes. Our data suggest that complementarity effects on mixed litter decomposition may emerge through long-term interactions between aboveground and belowground biota.     

What type of diversity yields synergy during mixed litter decomposition in a natural forest ecosystem?

Investigating the relationship of biodiversity and ecosystem function in natural forests allows incorporation of established feedbacks between long-lived plants and soil processes. We studied forested stands in northern Arizona that vary in dominant species richness across small areas. We examined the effects of natural variation in dominant tree biodiversity on ecosystem parameters, particularly litter decomposition. We determined not only whether plant species decompose in mixture as predicted by their individual decomposition rates but also: (1) how particular species affect the decomposition rate of each other in mixture; and (2) whether litter decomposes more rapidly at its site of origin; i.e. is there a “home field advantage” to decomposition? Over a 2-year period, litter mixtures of functionally similar tree species decomposed more rapidly than expected from rates of the individual species alone. Mixtures of conifer species litter decomposed up to 50% faster than expected, with individual conifer members of those mixtures decomposing up to 85% faster than expected. In contrast, more functionally diverse mixtures of litter, which included a deciduous species, did not show synergistic effects during decomposition. We found no significant “home-field advantage” to decomposition. Our study is the first to demonstrate that litter mixtures from more closely related plant species give rise to the most synergistic effects of biodiversity on litter dynamics, indicating that more taxonomically and functionally diverse plant assemblages do not always drive greater emergent effects on ecosystem function.     

Browsing-induced Effects on Leaf Litter Quality and Decomposition in a Southern African Savanna

We investigated the linkages between leaf litter quality and decomposability in a savanna plant community dominated by palatable-spinescent tree species. We measured: (1) leaf litter decomposability across five woody species that differ in leaf chemistry; (2) mass decomposition, nitrogen (N); and carbon (C) dynamics in leaf litter of a staple browse species (Acacia nigrescens) as well as (3) variation in litter composition across six sites that experienced very different histories of attack from large herbivores. All decomposition trials included litter bags filled with chopped straw to control for variation in site effects. We found a positive relationship between litter quality and decomposability, but we also found that Acacia and straw litter mass remaining did not significantly vary between heavily and lightly browsed sites. This is despite the fact that both the quality and composition of litter returned to the soil were significantly different across sites. We observed greater N resorption from senescing Acacia leaves at heavily browsed sites, which in turn contributed to increase the C:N ratio of leaf litter and caused greater litter N immobilization over time. This, together with the significantly lower tree- and herb-leaf litter mass beneath heavily browsed trees, should negatively affect decomposition rates. However, estimated dung and urine N deposition from both browsers and grazers was significantly greater at high- than at low-herbivory sites. We hypothesize that N inputs from dung and urine boost litter N mineralization and decomposition (especially following seasonal rainfall events), and thereby offset the effects of poor leaf litter quality at chronically browsed sites. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Mixed leaf litter effects on decomposition rates and soil microarthropod communities in an oak–pine stand in Japan

Rates of decomposition, and soil faunal abundance and diversity associated with single-species and mixed-species litters were studied in a litter bag experiment in an oak–pine forest. We used two canopy species of leaf litter, pine and oak, and one shrub species, Sasa, and compared decomposition rates, and soil microarthropod abundance and community structure of oribatid mites in the litter bags. Mass loss of single species decreased in the order: oak > pine > Sasa. While the total mass loss rates of mixed litter were intermediate between those of the constituent species, enhancement of mass loss from the three-species mixture and from mixed slow-decomposing litters (pine and Sasa) was observed. Faunal abundance in litter bags was higher in mixed-species litter than in those with single-species litter, and species richness of oribatid mites was also higher in the three-species mixed litter. Faunal abundance in single-species litter bags was not correlated with mass loss, although enhancement of mass loss in mixed litter bags corresponded with higher microarthropod abundance. Habitat heterogeneity in mixed litter bags seemed to be responsible for the more abundant soil microarthropod community.     

Effects of fertilisation and irrigation on ‘foliar afterlife’ in alpine tundra

Question: How do increases in soil nutrient and water availability alter the nutrient fluxes through the resorption and litter decomposition pathways and how do they affect litter nutrient pools in a low‐productive alpine tundra ecosystem? Location: An alpine lichen‐rich tundra on Mt. Malaya Khati‐para in the NW Caucasus, Russia (43°27’ N, 41°42’ E; altitude 2800 m a.s.l.). Methods: We conducted a 4‐year fertilisation (N, P, N+P, lime) and irrigation experiment, and analysed the responses of nutrient resorption from senescing leaves, leaf litter quality and decomposability of six pre‐dominant vascular plant species, total plant community litter production and litter (nutrient) accumulation. Results: Vascular plant litter [N] and [P] increased 1.5 and 10 fold in response to N and P additions, due to increased concentrations of the nutrients in fresh leaves and unchanged or reduced resorption efficiency. Litter decomposability was not affected by nutrient amendments. Fertilisation enhanced litter production (180%; N+P treatment) and litter accumulation (80%; N+P), owing to tremendously increased production and low decomposability of graminoids. Together with increased litter [N] and [P] this led to great increases in total litter nutrient pools. Conclusions: Due to increased production of graminoids, nutrients added to the alpine tundra soil were mostly immobilised in recalcitrant, nutrient‐rich litter. This suggests that changing species composition in low productive ecosystems may act as an internal buffer mechanism, which under increased soil nutrient availability prevents the community from rapidly acquiring features typical of a high productive ecosystem such as high decomposability and high nutrient availability.     

Do non‐additive effects on decomposition in litter‐mix experiments result from differences in resource quality between litters?

Differences in resource quality between litter species have been postulated to explain why litter-mixtures may decompose at a different rate to that which would be predicted from single species litters (termed 'non-additive effects'). In particular, positive, non-additive effects of litter-mixing on decomposition have been explained by differences in initial nitrogen concentration between litter species. This interpretation is confounded because litter species that differ in nitrogen content also differ by a number of other resource quality attributes. Thus, to investigate whether initial nitrogen concentration does account for positive, non-additive effects of litter-mixing, we mixed grass litters that differed in initial nitrogen concentration but not species or structural plant part identity, and then followed mass loss from the litter-mixes over time. We used the litterbag technique and three grass species for which a gradient of four distinct initial nitrogen concentrations had been generated. We produced all no- to four-mix compositions of litter qualities for each species. Litter from different species was never mixed.
Contrary to what would be predicted, we found that when litters of the same species but with different initial nitrogen concentrations were mixed, that negative, non-additive effects on decomposition were generally observed. In addition, we found that once mixed, increasing litter quality richness from two to four mixtures had no significant, non-additive effect on decomposition. Litter quality composition explained little of the experimental variation when compared to litter quality richness, and different compositions generally behaved in the same manner. Our findings challenge the commonly held assumption that differences in nitrogen concentration between plant species are responsible for positive, non-additive effects of litter-mixing on decomposition.     

Litter quality and decomposability of species from a Mediterranean succession depend on leaf traits but not on nitrogen supply

Background and Aims

The rate of plant decomposition depends on both the decomposition environment and the functional traits of the individual species (e.g. leaf and litter quality), but their relative importance in determining interspecific differences in litter decomposition remains unclear. The aims of this study were to: (a) determine if species from different successional stages grown on soils with low and high nitrogen levels produce leaf and litter traits that decompose differently under identical conditions; and (b) assess which trait of living leaves best relates to litter quality and litter decomposability

Methods

The study was conducted on 17 herbaceous species representative of three stages of a Mediterranean successional sere of Southern France. Plants were grown in monocultures in a common garden under two nitrogen levels. To elucidate how different leaf traits affected litter decomposition a microcosm experiment was conducted to determine decomposability under standard conditions. Tests were also carried out to determine how successional stage and nitrogen supply affected functional traits of living leaves and how these traits then modified litter quality and subsequent litter decomposability.

Key Results

The results demonstrated that leaf traits and litter decomposability varied according to species and successional stage. It was also demonstrated that while nitrogen addition affected leaf and litter traits, it had no effect on decomposition rates. Finally, leaf dry matter content stood out as the leaf trait best related to litter quality and litter decomposability

Conclusions

In this study, species litter decomposability was affected by some leaf and litter traits but not by soil nitrogen supply. The results demonstrated the strength of a trait-based approach to predict changes in ecosystem processes as a result of species shifts in ecosystems.Key words: Leaf traits, litter quality, litter decomposability, nitrogen addition, secondary succession     

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

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Consequences of the reduction of plant diversity for litter decomposition: effects through litter quality and microenvironment

Decomposition of plant litter is a key process for the flow of energy and nutrients in ecosystems that may be sensitive to the loss of biodiversity. Two hypothetical mechanisms by which changes in plant diversity could affect litter decomposition are (1) through changes in litter species composition, and (2) by altering the decomposition microenvironment. We tested these ideas in relation to the short-term decomposition of herbaceous plant litter in experimental plant assemblages that differed in the numbers and types of plant species and functional groups that they contained to simulate loss of plant diversity. We used different litterbag experiments to separate the two potential pathways through which diversity could have an effect on decomposition. Our two litterbag trials showed that altering plant diversity affected litter breakdown differently through changes in decomposition microenvironment than through changes in litter composition. In the decomposition microenvironment experiment there was a significant but weak decline in decomposition rate in relation to decreasing plant diversity but no significant effect of plant composition. The litter composition experiment showed no effect of richness but significant effects of composition, including large differences between plant species and functional groups in litter chemistry and decomposition rate. However, for a nested subset of our litter mixtures decomposition was not accurately predicted from single-species bags; there were positive, non-additive effects of litter mixing which enhanced decomposition. We critically assess the strengths and limitations of our short-term litterbag trials in predicting the longer-term effects of changes in plant diversity on litter decomposition rates.     

Litter mixing significantly affects decomposition in the Hulun Buir meadow steppe of Inner Mongolia,China

Aims We explored the decomposition rates of single- and mixed-species litter, the litter-mixing effect and the effect of component litters in a mixture on decomposition.Methods In a litter bag experiment, shoot litters from two dominant grasses (Leymus chinensis and Stipa baicalensis) and one legume (Melissitus ruthenica) were decomposed separately and as a mixture from May 2010 to September 2011 in the Hulun Buir meadow steppe of Inner Mongolia, China. We separated the litter mixture into its individual component litters (i.e. the different single-species litters) and analyzed the changes in litter mass remaining and litter nitrogen (N) remaining during single- and mixed-species litter decomposition.Important findings (i) Litter mixing had significant positive effects on litter decomposition. The litter-mixing effect was strongest for the mixture of S. baicalensis and L. chinensis litters, followed by the mixture of S. baicalensis and M. ruthenica litters. (ii) Single-species component litters decomposed faster in the mixtures than separately (positive effect), but these effects were not significant for legume species M. ruthenica litter. Relative to single-species litter decomposition, the decomposition rates of the two grass (S. baicalensis and L. chinensis) litters significantly increased when they were mixed with each other or with M. ruthenica litter. (iii) For each species litter type, the percentage of litter N remaining during decomposition (N R) differed between the single-species litter and mixed litter treatments. The N R of S. baicalensis litter was higher when it was decomposed in the mixture than in isolation. However, the N R of L. chinensis litter was lowest in its mixture with M. ruthenica among the treatments. Regardless of its decomposition in the mixture or in isolation, the N R of M. ruthenica litter varied little among treatments. There was a significant positive relationship between the N R and percentage of initial litter mass remaining in both the single litter and mixed litter treatments. These results suggest that N transfer may happen among component litters in mixture and further affect the decomposition.     

Chemical diversity – highlighting a species richness and ecosystem function disconnect

The lack of predictability in litter-mix studies may result from the low correlation between species number and the traits that drive the processes under observation. From the standpoint of litter-quality-dependent ecological processes, we propose that litter chemical qualities are functional traits and introduce a multivariate index of chemical diversity (CD_Q) based on Rao's quadratic entropy to describe the compositional heterogeneity of litter and foliar mixtures. Using published data from temperate and tropical forest systems to illustrate the relationship between species richness and chemical diversity, we show the variation of chemical diversity based on profiles of total nutrient concentrations (N, P, K, Ca and Mg) with species richness. We discuss how this behavior may explain the idiosyncratic responses exhibited in litter-mix experiments and how it may contribute to the observed dominance of species identity over species diversity. As a summary of resource heterogeneity relevant to detritivore and microbial processes, the chemical diversity index is potentially a better predictor of diversity effects on nutrient dynamics than species richness. Finally, we propose the use of infrared spectroscopy techniques for a rapid and more comprehensive determination of foliar and litter chemical composition to provide a more information-rich index.     

Impacts of tree species diversity on litter decomposition in northern temperate forests of Wisconsin, USA: a multi-site experiment along a latitudinal gradient

Over the past decade an increasing amount of research has sought to understand how the diversity of species in an ecosystem can influence fluxes of biologically important materials, such as the decomposition of organic matter and recycling of nutrients. Generalities among studies have remained elusive, perhaps because experimental manipulations have been performed at relatively small spatial scales where site-specific variation generates patterns that appear idiosyncratic. One approach for seeking generality is to perform parallel experiments at different sites using an identical species pool. Here we report results from a study where we manipulated the diversity of leaf litter from the same six dominant tree species in the litter layer of three forested ecosystems. These ecosystems spanned a 300 km latitudinal transect in Wisconsin, USA, and were characterized by a large gradient in temperature and moisture, and thus, rates of decomposition. After allowing combinations of one, two, four, and six species of leaf litter to decompose for 1 year, we found that increasing leaf litter richness led to slower rates of decomposition and higher fractions of nitrogen lost from litter. Across all sites, climate and initial litter chemistry explained more of the variation in decomposition rates than did litter richness. Effects of leaf litter diversity were non-additive, meaning they were greater than expected from the impacts of individual species, and appeared to be strongly influenced by the presence/absence of just 1–2 species (Tilia americana and Acer saccharum). The rate of decomposition of these two species was highly site-specific, which led to strong negative effects of litter richness only being observed at the southernmost sites where T. americana and A. saccharum decomposed more quickly. In contrast, litter diversity increased nitrogen loss at the northernmost sites where decomposition of T. americana was notably slowed. Our study shows that species diversity affected at least one of the two litter processes at each site along this 300-km gradient, but the exact nature of these effects were spatially variable because the performance of individual species changed across the heterogeneous landscape.     

Edge and herbivory effects on leaf litter decomposability in a subtropical dry forest

It is increasingly recognized that understanding the functional consequences of landscape change requires knowledge of aboveground and belowground processes and their interactions. For this reason, we provide novel information addressing insect herbivory and edge effects on litter quality and decomposition in fragmented subtropical dry forests in central Argentina. Using litter from Croton lachnostachyus (a common shrub species in the region) in a decomposition bed experiment, we evaluated whether litter quality (carbon and nitrogen content; carbon: nitrogen ratio) and decomposability (percentage of remaining dry weight) differ between litter from forest interiors or edges (origin) and with or without herbivory (damaged/undamaged leaves). We found that edge/interior origin had a strong effect on leaf litter quality (mainly on carbon content), while herbivory was associated with a smaller increase in nitrogen content. Herbivore damage increased leaf litter decomposability, but this effect was related to origin during the initial period of litter incubation. Overall, undamaged leaf litter from the forest edge showed the lowest decomposability, whereas damaged leaf litter decomposed at rates similar to those observed in litter from the forest interior. The interacting edge and herbivory effects on leaf litter quality and decomposability shown in our study are important because of the increasing dominance of forest edges in human-modified landscapes and the profound effect of leaf litter decomposition on nutrient cycling.     

Changing leaf litter feedbacks on plant production across contrasting sub-arctic peatland species and growth forms

Plant species and growth forms differ widely in litter chemistry, which affects decay and may have important consequences for plant growth via e.g. the release of nutrients and growth-inhibitory compounds. We investigated the overall short-term (9.5 months) and medium-term (21.5 months) feedback effects of leaf litter quality and quantity on plant production, and tested whether growth forms can be used to generalise differences among litter species. Leaf litter effects of 21 sub-arctic vascular peatland species on Poa alpina test plants changed clearly with time. Across all growth forms, litter initially reduced plant biomass compared with untreated plants, particularly litters with a high decomposition rate or low initial lignin/P ratio. In the second year, however, litter effects were neutral or positive, and related to initial litter N concentration (positive), C/N, polyphenol/N and polyphenol/P ratios (all negative), but not to decomposability. Differences in effect size among several litter species were large, while differences in response to increasing litter quantities were not significant or of similar magnitude to differences in response to three contrasting litter species. Growth forms did not differ in initial litter effects, but second-year plant production showed a trend (P < 0.10) for differences in response to litters of different growth forms: evergreen shrubs < graminoids or deciduous shrubs < forbs. While long-persisting negative litter effects were predominant across all growth forms, our data indicate that even within nutrient-constrained ecosystems such as northern peatlands, vascular plant species, and possibly growth forms, differ in litter feedbacks to plant growth. Differences in the composition of undisturbed plant communities or species shifts induced by external disturbance, such as climate change, may therefore feedback strongly to plant biomass production and probably nutrient cycling rates in northern peatlands. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Limited evidence that mixing leaf litter accelerates decomposition or increases diversity of decomposers in streams of eastern Canada

Most studies of terrestrial litter decomposition in streams and rivers have used leaves from a single tree species, but leaf packs in streams in eastern North America are usually mixtures of two or more species. Litter mixtures may decay more quickly than either of the component species. If so, estimates of stream energy and nutrient budgets may be inaccurate. In northern Nova Scotia, Canada, we measured mass loss from binary mixtures (1:1 mass ratio) of leaf litter in mesh bags, using freshly fallen or air-dried litter from five species of canopy trees. We repeated the experiment eight times, in summer and fall, in two streams and a small river, over 3 years. In some trials we enumerated benthic invertebrate and fungal colonization of decaying litter. Although there were marked differences in mass loss rates among litter types, decomposition was accelerated in mixtures relative to the mean of the component species in only three of eight trials, and only in mixtures containing N-rich speckled alder leaves. Mixing yellow birch and red maple leaves inhibited decomposition. Diversity (Shannon–Weaver Index), species richness, and abundance of aquatic hyphomycete fungi, as indexed by conidial production, were never greater (and sometimes less) on litter mixtures than on the component species. Total numbers, taxonomic richness and diversity of benthic invertebrates generally, and litter-feeding species in particular, were not augmented by mixing litter types. Litter mixtures appear to dilute a preferred substrate with patches of a less preferred substrate. Our results provide only weak support for the contention that combining two litter types leads to acceleration of decomposition rates. Handling editor: K. Martens     

Plant palatability to leaf‐cutter ants (Atta laevigata) and litter decomposability in a Neotropical woodland savanna

Although leaf‐cutter ants have been recognized as the dominant herbivore in many Neotropical ecosystems, their role in nutrient cycling remains poorly understood. Here we evaluated the relationship between plant palatability to leaf‐cutter ants and litter decomposability. Our rationale was that if preference and decomposability are related, and if ant consumption changes the abundance of litter with different quality, then ant herbivory could affect litter decomposition by affecting the quality of litter entering the soil. The study was conducted in a woodland savanna (cerrado denso) area in Minas Gerais, Brazil. We compared the decomposition rate of litter produced by trees whose fresh leaves have different degrees of palatability to the leaf‐cutter ant Atta laevigata. Our experiments did not indicate the existence of a significant relationship between leaf palatability to A. laevigata and leaf‐litter decomposability. Although the litter mixture composed of highly palatable plant species showed, initially, a faster decay rate than the mixture of poorly palatable species, this difference was no longer visible after about 6 months. Results were consistent regardless of whether litter invertebrates were excluded or not from litter bags. Similarly, experiments comparing the decomposition rate of litter from pairs of related plant species also showed no association between plant palatability and decomposition. Decomposition rate of the more palatable species was faster, slower or similar to that of the less palatable species depending upon the particular pair of species being compared. We suggest that the traits that mostly influence the decomposition rate of litter produced by cerrado trees may not be the same as those that influence plant palatability to leaf‐cutter ants. Atta laevigata select leaves of different species based – at least in part – on their nitrogen content, but N content was a poor predictor of the decomposition rates of the species we studied.