Impacts of invasive riparian knotweed on litter decomposition,aquatic fungi,and macroinvertebrates

Bohemian knotweed (Polygonum × bohemicum), the hybrid between Japanese and giant knotweed, is the most common invasive knotweed species in western North America and the most difficult to control. Invasive knotweed congeners spread aggressively along streams and establish dense monotypic stands, reducing riparian plant species diversity. Allochthonous organic matter inputs from riparian plants are an important source of energy and nutrients for organisms in small streams. However, little information exists concerning the influence of knotweed on stream processes. This study examines the quality of Bohemian knotweed leaves compared to native red alder and black cottonwood leaves, along with leaf-associated fungal biomass accumulation, macroinvertebrate communities, and decay rates from three forested streams in western Washington State. Senesced knotweed leaves were lower in nitrogen and phosphorus, and higher in cellulose, fiber, and lignin content than alder leaves, but were more similar to cottonwood leaves. Fungal biomass differed among species and changed over time. Macroinvertebrate shredders collected from leaf packs after 31 days were proportionately more abundant on alder leaves than knotweed and cottonwood. Decay rates were not significantly different among leaf species, but during the first 31 days alder broke down faster than knotweed. After 56 days, all of the leaf packs were mostly decomposed. Overall, these findings do not show major discrepancies between leaf species except those related to initial litter structural and chemical quality. However, changes in the timing and quantity of litter inputs are also important factors to be considered in understanding the impact of invasive knotweed on stream ecosystem processes.     

Over-winter decomposition and associated macroinvertebrate communities of three deciduous leaf species in forest streams on the Canadian Boreal Shield

The decomposition of deciduous leaf material provides a critical source of energy to aquatic food webs. Changes to riparian forests through harvesting practices may alter the species composition of deciduous leaf material entering streams. We compared over-winter decomposition of three different riparian leaf species (speckled alder (Alnus incana ssp. rugosa (Du Roi) J. Clausen), white birch (Betula papyrifera Marsh.), and trembling aspen (Populus tremuloides Michx.)) to determine their importance as a food resource for macroinvertebrate communities within Boreal Shield streams in northeastern Ontario, Canada. Leaf pack decomposition of the three leaf species formed a processing continuum throughout winter, where alder and birch leaf packs decomposed at a medium rate (k = 0.0065/day and 0.0053/day, respectively) and aspen leaf packs decomposed more slowly (k = 0.0035/day). Macroinvertebrate community colonization on leaf packs changed through time regardless of leaf species. Alder leaf packs supported higher abundances of macroinvertebrates in the fall while aspen leaf packs supported greater shredder abundances in the following spring. The study shows that leaf diversity may be important for providing a sustained food resource for aquatic macroinvertebrates throughout the relatively long over-winter period in Canadian Boreal Shield streams. Riparian forest management strategies should ensure that deciduous plant species richness is sustained in riparian areas.     

Effects of invasive European bird cherry (Prunus padus) on leaf litter processing by aquatic invertebrate shredder communities in urban Alaskan streams

European bird cherry (Prunus padus) (EBC) is an invasive ornamental tree that is spreading rapidly in riparian forests of urban Alaska. To determine how the spread of EBC affects leaf litter processing by aquatic invertebrate shredders, we conducted complementary leaf pack experiments in two streams located in Anchorage, Alaska. The first experiment contrasted invasive EBC with three native tree species—thin-leaf alder (Alnus tenuifolia), paper birch (Betula neoalaskana), and black cottonwood (Populus trichocarpa)—in one reach of Chester Creek; finding that EBC leaf litter broke down significantly faster than birch and cottonwood, but at a similar rate to alder. The second experiment contrasted EBC with alder in four reaches of Campbell and Chester creeks; finding that while EBC leaf litter broke down significantly faster than alder in Chester Creek, EBC broke down at a similar rate to alder in Campbell Creek. Although EBC sometimes supported fewer shredders by both count and mass, shredder communities did not differ significantly between EBC and native plants. Collectively, these data suggest that invasive EBC is not currently exhibiting strong negative impacts on leaf litter processing in these streams, but could if it continues to spread and further displaces native species over time.     

Assessing the influence of wildfire on leaf decomposition and macroinvertebrate communities in boreal streams using mixed-species leaf packs

1. We investigated how compositional differences in riparian leaf litter derived from burned and undisturbed forests influenced leaf breakdown and macroinvertebrate communities using experimental mixed-species leaf packs in boreal headwater streams. Leaf pack mixtures simulating leaf litter from dominant riparian woody-stem species in burned and undisturbed riparian zones were incubated in two references and two fire-disturbed streams for 5 weeks prior to measuring temperature-corrected breakdown rates and macroinvertebrate community composition, richness, and functional metrics associated with decomposers such as shredder abundance and % shredders.
2. Leaf litter breakdown rates were higher and had greater variability in streams bordered by reference riparian forests than in streams where riparian forests had been burned during a wildfire. Streams bordered by fire disturbance showed significant effects of litter mixture on decomposition rates, observed as significantly higher decomposition rates of a fire-simulated leaf mixture compared to all other mixtures.
3. Variation among sites was higher than variation among litter mixtures, especially for macroinvertebrate community composition. In general, fire-simulated leaf mixtures had greater shredder abundances and proportions, but lower overall macroinvertebrate abundance; however, the shredder abundance trend was not consistent across all leaf mixtures at each stream.
4. These results show that disturbance-driven riparian forest condition and resulting composition of leaf subsidies to streams can influence aquatic invertebrate community composition and their function as decomposers. Therefore, if one of the primary goals of modern forest management is to emulate natural disturbance patterns, boreal forest managers should adapt silvicultural practices to promote leaf litter input that would arise post-fire, thereby supporting stream invertebrate communities and their function.

     

The Importance of Detritivore Species Diversity for Maintaining Stream Ecosystem Functioning Following the Invasion of a Riparian Plant

The strength of linkages between riparian plants and stream communities can be expected to be influenced by invading plants. While most studies so far have been focussed on the effects of the leaf litter quality of the invader, this study addresses the impact of detritivores on the pool of detritus. In a natural setting, we found that species richness of shredding macroinvertebrates significantly influenced the breakdown rate of an invasive weed species, the Japanese knotweed (Fallopia japonica), which has become a major plant invader along streams and rivers in Europe and North America. Our findings imply that a reduction of the diversity of shredder species, which may be the result of disturbances, could negatively influence stream ecosystems' capacity of processing knotweed leaves. Although the knotweed showed breakdown rates similar to those of common native tree and shrub species, other exotic leaf species might show considerably slower rates and hence have greater consequences for the ecosystems. We have, in this study, indicated a technique by which the effects of other non-indigenous plants on ecosystem functioning might be considered.     

Do non-native Platanus hybrida riparian plantations affect leaf litter decomposition in streams?

In forest headwater streams where the riparian canopy limits autochthonous primary production, leaf litter decomposition is a key process controlling nutrient and carbon cycling. Any alteration of the riparian vegetation may influence litter decomposition and detrital food webs. We evaluated the effect of non-native Platanus hybrida riparian plantations on leaf litter decomposition in Mediterranean streams. The experiment was conducted in six headwater streams; three lined by native riparian vegetation and three crossing P. hybrida plantations. We have characterized the processing rates of alder leaves and the assemblages of shredder macroinvertebrates and fungi. Litter decomposition was significantly faster in the P. hybrida than in the reference streams. Although the dissolved inorganic nitrogen concentration was higher in P. hybrida, no significant effect was observed in decomposition rates. Differences in decomposition rates reflected the macroinvertebrate and shredder colonization in alder litter, with higher abundance and richness in the P. hybrida streams. However, aquatic hyphomycete sporulation rate was higher in reference streams, suggesting that the variation in decomposition rates is a direct consequence of shredder abundance. Our findings support part of the substrate quality-matrix quality (SMI) hypothesis, which expects that high-quality litter will show increased decomposition rates in a low-quality litter matrix.     

Composition of riparian litter input regulates organic matter decomposition: Implications for headwater stream functioning in a managed forest landscape

Although the importance of stream condition for leaf litter decomposition has been extensively studied, little is known about how processing rates change in response to altered riparian vegetation community composition. We investigated patterns of plant litter input and decomposition across 20 boreal headwater streams that varied in proportions of riparian deciduous and coniferous trees. We measured a suite of in‐stream physical and chemical characteristics, as well as the amount and type of litter inputs from riparian vegetation, and related these to decomposition rates of native (alder, birch, and spruce) and introduced (lodgepole pine) litter species incubated in coarse‐ and fine‐mesh bags. Total litter inputs ranged more than fivefold among sites and increased with the proportion of deciduous vegetation in the riparian zone. In line with differences in initial litter quality, mean decomposition rate was highest for alder, followed by birch, spruce, and lodgepole pine (12, 55, and 68% lower rates, respectively). Further, these rates were greater in coarse‐mesh bags that allow colonization by macroinvertebrates. Variance in decomposition rate among sites for different species was best explained by different sets of environmental conditions, but litter‐input composition (i.e., quality) was overall highly important. On average, native litter decomposed faster in sites with higher‐quality litter input and (with the exception of spruce) higher concentrations of dissolved nutrients and open canopies. By contrast, lodgepole pine decomposed more rapidly in sites receiving lower‐quality litter inputs. Birch litter decomposition rate in coarse‐mesh bags was best predicted by the same environmental variables as in fine‐mesh bags, with additional positive influences of macroinvertebrate species richness. Hence, to facilitate energy turnover in boreal headwaters, forest management with focus on conifer production should aim at increasing the presence of native deciduous trees along streams, as they promote conditions that favor higher decomposition rates of terrestrial plant litter.     

Effect of native and exotic leaf litter on macroinvertebrate communities and decomposition in a western Montana stream

Invasion by exotic trees into riparian areas has the potential to impact terrestrial and aquatic systems. To test the effect of different streamside tree species on the aquatic food web in a stream in Montana, we compared decomposition rates of leaf litter and invertebrate assemblages associated with the leaf litter of the exotic Acer platanoides and the dominant native Populus trichocarpa trees. Macroinvertebrate family richness, evenness, and diversity increased with days of aquatic processing; however, there was no effect of leaf species. Leaves of the A. platanoides were associated with 70% greater density of macroinvertebrates of the family Nemouridae. This family consists primarily of detritivores and had the greatest overall density and frequency of occurrence relative to other macroinvertebrate families. The density of a family of predatory macroinvertebrates (Rhyacophilidae) was also generally (73%) greater in association with A. platanoides than P. trichocarpa leaves. The density of Ephemerellidae and Rhyacophilidae increased over time. In contrast to studies comparing leaves of exotic vs. native trees, we observed no difference in leaf decomposition rates; however, the amount of leaf inputs are likely to differ between native and invaded forests. The results indicate that replacement of native riparian trees with exotics affected the most common family of macroinvertebrates and possibly a common family of predatory macroinvertebrates (Rhyacophilidae), which may affect the detrital food web.     

Higher temperature reduces the effects of litter quality on decomposition by aquatic fungi

1. We investigated the effects of riparian plant diversity (species number and identity) and temperature on microbially mediated leaf decomposition by assessing fungal biodiversity, fungal reproduction and leaf mass loss. 2. Leaves of five riparian plant species were first immersed in a stream to allow microbial colonisation and were then exposed, alone or in all possible combinations, at 16 or 24 °C in laboratory microcosms. 3. Fungal biodiversity was reduced by temperature but was not affected by litter diversity. Temperature altered fungal community composition with species of warmer climate, such as Lunulospora curvula, becoming dominant. 4. Fungal reproduction was affected by litter diversity, but not by temperature. Fungal reproduction in leaf mixtures did not differ or was lower than that expected from the weighted sum of fungal sporulation on individual leaf species. At the higher temperature, the negative effect of litter diversity on fungal reproduction decreased with the number of leaf species. 5. Leaf mass loss was affected by the identity of leaf mixtures (i.e. litter quality), but not by leaf species number. This was mainly explained by the negative correlation between leaf decomposition and initial lignin concentration of leaves. 6. At 24 °C, the negative effects of lignin on microbially mediated leaf decomposition diminished, suggesting that higher temperatures may weaken the effects of litter quality on plant litter decomposition in streams. 7. The reduction in the negative effects of lignin at the higher temperature resulted in an increased microbially mediated litter decomposition, which may favour invertebrate‐mediated litter decomposition leading to a depletion of litter stocks in streams.     

Litter quality, stream characteristics and litter diversity influence decomposition rates and macroinvertebrates

1. We examined the relative importance of litter quality and stream characteristics in determining decomposition rate and the macroinvertebrate assemblage living on autumn‐shed leaves. 2. We compared the decomposition rates of five native riparian tree species (Populus fremontii, Alnus oblongifolia, Platanus wrightii, Fraxinus velutina and Quercus gambelii) across three south‐western streams in the Verde River catchment (Arizona, U.S.A.). We also compared the decomposition of three‐ and five‐species mixtures to that of single species to test whether plant species diversity affects rate. 3. Decomposition rate was affected by both litter quality and stream. However, litter quality accounted for most of the variation in decomposition rates. The relative importance of litter quality decreased through time, explaining 97% of the variation in the first week but only 45% by week 8. We also found that leaf mixtures decomposed more quickly than expected, when all the species included were highly labile or when the stream environment led to relatively fast decomposition. 4. In contrast to decomposition rate, differences in the invertebrate assemblage were more pronounced across streams than across leaf litter species within a stream. We also found significant differences between the invertebrate assemblage colonising leaf mixtures compared with that colonising pure species litter, indicating non‐additive properties of litter diversity on stream invertebrates. 5. This study shows that leaf litter diversity has the capacity to affect in‐stream decomposition rates and stream invertebrates, but that these effects depend on both litter quality and stream characteristics.     

Implications of water column ammonium uptake and regeneration for the nitrogen budget in temperate, eutrophic Missisquoi Bay, Lake Champlain (Canada/USA)

Invasion by exotic trees into riparian areas has the potential to impact aquatic systems. We examined the effects of the exotic Salix fragilis (crack willow) on the structure and functioning of small streams in northern Patagonian Andes via a field survey of benthic invertebrates and leaf litter and an in situ experiment. We compared leaf decomposition of the native Ochetophila trinervis (chacay) and S. fragilis in reaches dominated by native vegetation versus reaches dominated by crack willow. We hypothesized that S. fragilis affects the quality of leaf litter entering the streams, changing the aquatic biota composition and litter decomposition. Our study showed that crack willow leaves decomposed slower than chacay, likely related to leaf properties (i.e., leaf toughness). Benthic leaf litter mass was similar between the two riparian vegetation types, though in stream reaches dominated by crack willow, leaves of this species represented 82% of the total leaf litter. Benthic invertebrate abundance and diversity were similar between reaches but species composition differed. Our study found little evidence for strong impacts of crack willow on those small streams. Further studies on other aspects of ecosystem functioning, such as primary production, would enhance our understanding of the impacts of crack willow on Patagonian streams.     

Leaf‐Litter Mixtures Affect Breakdown and Macroinvertebrate Colonization Rates in a Stream Ecosystem

Previous work in terrestrial and aquatic ecosystems has suggested that the relationship between breakdown rates of leaf litter and plant species richness may change unpredictability due to non‐additive effects mediated by the presence of key‐species. By using single‐ and mixed‐species leaf bags (7 possible combinations of three litter species differing in toughness; common alder [Alnus glutinosa ], sweet chestnut [Castanea sativa ], and Spanish oak [Quercus ilex ilex ]), I tested whether leaf species diversity, measured as richness and composition, affects breakdown dynamics and macroinvertebrate colonization (abundance, richness and composition) during 90 days incubation in a stream. Decomposition rates were additive, i.e., observed decomposition rates were not different from expected ones. However, decomposition rates of individual leaf species were affected by the mixture, i.e., there were species‐specific responses to mixing litter. The invertebrate communities colonizing the mixtures were not richer and more diverse in mixtures than in single‐species leaf bags. On the opposite, mixing leaf species had a negative, non‐additive effect on rates of shredder and taxa colonization and on macroinvertebrate diversity. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Decomposition of Eucalyptus globulus leaves and three native leaf species (Alnus glutinosa, Castanea sativa and Quercus faginea) in a Portuguese low order stream

Leaf decomposition of the exotic evergreen Eucalyptus globulus (eucalyptus), and three native deciduous tree species, Alnus glutinosa (alder), Castanea sativa (chestnut) and Quercus faginea (oak), was compared in a second order stream in Central Portugal. Changes in dry weight, nitrogen and polyphenolic compounds and microbial colonization were periodically assessed for three months.Negative exponential curves fit the leaf weight loss with time for all leaf species. Mass loss rate was in the order alder (K = 0.0161) > chestnut (K = 0.0079) > eucalyptus (K = 0.0068) > oak (K = 0.0037). Microbial colonization followed the same pattern as breakdown rates. Evidence of fungal colonization was observed in alder after 3 days in the stream, whereas it took 21 days in oak leaves to have fungal colonization. Fungal diversity was leaf species-dependent and increased with time. In all cases, percent nitrogen per unit leaf weight increased, at least, at the initial stages of decay while soluble polyphenolics (expressed as percentage per unit leaf weight) decreased rapidly in the first month of leaves immersion.Intrinsic factors such as nitrogen and polyphenolic content may explain differences in leaf decomposition. The possible incorporation of eucalyptus litter into secondary production in a reasonable time span is suggested, although community balance and structure might be affected by differences in allochthonous patterns determined by eucalyptus monocultures.     

外来植物凋落物分解对底栖动物多样性及其摄食功能群的影响

在亚热带地区一条2级溪流中比较了本地植物(红锥和薏米)和外来植物(托里桉和马缨丹)凋落物的分解速率及底栖动物定殖及其摄食功能群的组成。结果显示,本地植物凋落物的分解速率为:0.020 d-1(红锥)和0.056 d-1(薏米),外来树叶的分解速率则为:0.038 d-1(托里桉)和0.041 d-1(马缨丹),它们均属于快速分解组。定殖在外来和本地树叶凋落物上的大型底栖动物种类分别为21种和24种。在外来和本地树叶上定殖的底栖动物平均密度差异不显著(P0.05)。从大型底栖动物摄食功能群的比例来看,定殖在本地凋落物上的集食者的比例最高(73.5%),其次是撕食者(10.9%)和捕食者(8.7%),刮食者(6.8%)的比例最低,在外来凋落物上,摄食功能群的比例分别为集食者(67.6%)、撕食者(13.9%)、刮食者(12.1%)和捕食者(6.1%),两者相比,底栖动物的摄食功能群均以集食者和撕食者为主,在组成比例上并无显著差异。结果表明2种外来植物凋落物分解对底栖动物多样性及其摄食功能群的影响不明显。     

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.     

Effects of Riparian Plant Diversity Loss on Aquatic Microbial Decomposers Become More Pronounced with Increasing Time

We examined the potential long-term impacts of riparian plant diversity loss on diversity and activity of aquatic microbial decomposers. Microbial assemblages were obtained in a mixed-forest stream by immersion of mesh bags containing three leaf species (alder, oak and eucalyptus), commonly found in riparian corridors of Iberian streams. Simulation of species loss was done in microcosms by including a set of all leaf species, retrieved from the stream, and non-colonized leaves of three, two or one leaf species. Leaves were renewed every month throughout six months, and microbial inoculum was ensured by a set of colonized leaves from the previous month. Microbial diversity, leaf mass loss and fungal biomass were assessed at the second and sixth months after plant species loss. Molecular diversity of fungi and bacteria, as the total number of operational taxonomic units per leaf diversity treatment, decreased with leaf diversity loss. Fungal biomass tended to decrease linearly with leaf species loss on oak and eucalyptus, suggesting more pronounced effects of leaf diversity on lower quality leaves. Decomposition of alder and eucalyptus leaves was affected by leaf species identity, mainly after longer times following diversity loss. Leaf decomposition of alder decreased when mixed with eucalyptus, while decomposition of eucalyptus decreased in mixtures with oak. Results suggest that the effects of leaf diversity on microbial decomposers depended on leaf species number and also on which species were lost from the system, especially after longer times. This may have implications for the management of riparian forests to maintain stream ecosystem functioning.     

A global meta‐analysis of exotic versus native leaf decay in stream ecosystems

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

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

Riparian plant species loss alters trophic dynamics in detritus-based stream ecosystems

Riparian vegetation is closely connected to stream food webs through input of leaf detritus as a primary energy supply, and therefore, any alteration of plant diversity may influence aquatic ecosystem functioning. We measured leaf litter breakdown rate and associated biological parameters in mesh bags in eight headwater streams bordered either with mixed deciduous forest or with beech forest. The variety of leaf litter types in mixed forest results in higher food quality for large-particle invertebrate detritivores (‘shredders’) than in beech forest, which is dominated by a single leaf species of low quality. Breakdown rate of low quality (oak) leaf litter in coarse mesh bags was lower in beech forest streams than in mixed forest streams, a consequence of lower shredder biomass. In contrast, high quality (alder) leaf litter broke down at similar rates in both stream categories as a result of similar shredder biomass in coarse mesh bags. Microbial breakdown rate of oak and alder leaves, determined in fine mesh bags, did not differ between the stream categories. We found however aquatic hyphomycete species richness on leaf litter to positively co-vary with riparian plant species richness. Fungal species richness may enhance leaf litter breakdown rate through positive effects on resource quality for shredders. A feeding experiment established a positive relationship between fungal species richness per se and leaf litter consumption rate by an amphipod shredder (Gammarus fossarum). Our results show therefore that plant species richness may indirectly govern ecosystem functioning through complex trophic interactions. Integrating microbial diversity and trophic dynamics would considerably improve the prediction of the consequences of species loss.     

Effect of leaf litter characteristics on leaf conditioning and on consumption by Gammarus pulex

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