Shifts in leaf litter breakdown along a forest–pasture–urban gradient in Andean streams

Tropical montane ecosystems of the Andes are critically threatened by a rapid land‐use change which can potentially affect stream variables, aquatic communities, and ecosystem processes such as leaf litter breakdown. However, these effects have not been sufficiently investigated in the Andean region and at high altitude locations in general. Here, we studied the influence of land use (forest–pasture–urban) on stream physico‐chemical variables (e.g., water temperature, nutrient concentration, and pH), aquatic communities (macroinvertebrates and aquatic fungi) and leaf litter breakdown rates in Andean streams (southern Ecuador), and how variation in those stream physico‐chemical variables affect macroinvertebrates and fungi related to leaf litter breakdown. We found that pH, water temperature, and nutrient concentration increased along the land‐use gradient. Macroinvertebrate communities were significantly different between land uses. Shredder richness and abundance were lower in pasture than forest sites and totally absent in urban sites, and fungal richness and biomass were higher in forest sites than in pasture and urban sites. Leaf litter breakdown rates became slower as riparian land use changed from natural to anthropogenically disturbed conditions and were largely determined by pH, water temperature, phosphate concentration, fungal activity, and single species of leaf‐shredding invertebrates. Our findings provide evidence that leaf litter breakdown in Andean streams is sensitive to riparian land‐use change, with urban streams being the most affected. In addition, this study highlights the role of fungal biomass and shredder species (Phylloicus; Trichoptera and Anchytarsus; Coleoptera) on leaf litter breakdown in Andean streams and the contribution of aquatic fungi in supporting this ecosystem process when shredders are absent or present low abundance in streams affected by urbanization. Finally, we summarize important implications in terms of managing of native vegetation and riparian buffers to promote ecological integrity and functioning of tropical Andean stream ecosystems.     

Effects of chronic chlorine exposure on litter processing in outdoor experimental streams

SUMMARY 1. The effects of chlorine on litter ( Potamogeton crispus L.) processing were examined using six outdoor experimental streams. Downstream portions of two streams were dosed at c . 10 μg l⁻¹ Total Residual Chlorine (TRC), one stream at 64 μgl¹, and one stream at 230μg l⁻¹. Two control streams were not dosed; upstream riffles of each stream served as instream controls.
2. Two 35 day litter breakdown (per cent AFDW remaining) experiments indicated significantly lower decay rates in the high dose riffle. No other concentration of chlorine significantly affected decay rate.
3. A third experiment, conducted in medium and high dose streams, indicated that high dose chlorine exposure reduced litter decomposition rates significantly, and reduced microbial colonization, microbial electron transport system activity, and microbial litter decomposition after 4 days but not after 11 days of exposure. The number of amphipod shredders colonizing litter bags was also reduced significantly with high chlorine dose.
4. A fourth experiment, after dosing was terminated, provided direct evidence that amphipod shredders were important in facilitating litter decomposition: litter bags stocked with amphipods had significantly higher decomposition rates than bags which excluded shredders.
5. Overall results indicate that the high dose (c. 230 μgl⁻¹ TRC) of chlorine reduced litter processing rates partly by reducing initial microbial conditioning, but primarily by reducing the colonization of amphipod shredders.     

Tadpoles enhance microbial activity and leaf decomposition in a neotropical headwater stream

1. Relationships between biodiversity and ecosystem function are of increasing interest, particularly in freshwater ecosystems where species losses are occurring at unprecedented rates. Amphibian declines have been associated with a loss of ecosystem function in neotropical streams, but little is known of the potential roles of stream‐dwelling tadpoles in leaf decomposition. Leaf litter is an important energy source to streams, and the breakdown of this material to fine particulate organic matter (FPOM) is a key ecosystem function. 2. We used mesocosms in a natural stream setting to quantify the effects of grazing tadpoles, shredding macroinvertebrates and a combination of the two on leaf decomposition and associated microbial activity. We measured respiration rates of decomposing leaves, particulate organic matter (POM) and leaf biofilm biomass and C : N : P ratios, and leaf area loss in 4 treatments: Control, tadpole only (TP), tadpole and shredding macroinvertebrates (TP + INV) and shredding macroinvertebrates only (INV). We hypothesised that tadpoles would enhance leaf decomposition by changing nutrient availability and stimulating microbial activity. 3. Respiration rates ranged from 3.1 to 6.0 mg O₂ dry mass^?1 h^?1 and were significantly higher in the TP and TP + INV treatments than in the control. The TP + INV treatment had significantly higher POM in chambers than the control and INV treatments. The TP treatment had significantly lower leaf biofilm biomass than the control and INV treatments. 4. Tadpoles influenced the elemental balance of C and N in POM and leaf biofilm. In contrast to our prediction, molar C : N ratios were higher in the TP + INV treatment than in the control. Mean molar N : P ratios in POM were higher in the TP + INV treatment than in any other treatment. Leaf biofilm followed a similar pattern, but both TP and TP + INV had significantly higher N : P ratios than the control and INV treatments. Leaf area loss was greatest when tadpoles and invertebrates were together (TP + INV = 0.6% leaf area loss per mg organism) than separate (TP = 0.1%, INV = 3%), indicating facilitation. 5. Tadpoles indirectly affected leaf decomposition by influencing microbial communities and macroinvertebrate feeding. As such, ongoing amphibian declines may adversely affect a critical ecosystem function in freshwater habitats.     

Litter decomposition in a temperate and a tropical stream: the effects of species mixing,litter quality and shredders

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Organic Matter Dynamics in a Tropical Gallery Forest in a Grassland Landscape

The high biodiversity of tropical forest streams depends on the strong input of organic matter, yet the leaf litter decomposition dynamics in these streams are not well understood. We assessed how seasonal litterfall affects leaf litter breakdown, density and biomass of aquatic invertebrates, and the microbial biomass and sporulation of aquatic hyphomycetes in a South American grassland ‘vereda’ landscape. Although litter production in the riparian area was low, leaf litter breakdown was high compared with other South American systems, with maximum values coinciding with the rainy season. Fungal biomass in decomposing leaves was high, but spore densities in water and sporulation rates were very low. Invertebrates were not abundant in litter bags, suggesting they play a minor role in leaf litter decomposition. Chironomids accounted for ~70 percent of all invertebrates; only 10 percent of non‐Chironomidae invertebrates were shredders. Therefore, fungi appear to be the drivers of leaf litter decomposition. Our results show that despite low productivity and relatively fast litter decomposition, organic matter accumulated in the stream and riparian area. This pattern was attributed to the wet/dry cycles in which leaves falling in the flat riparian zone remain undecomposed (during the dry period) and are massively transported to the riverbed (rainy season).     

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.     

Growth of an invertebrate shredder on native (Populus) and non-native (Tamarix, Elaeagnus) leaf litter

1. Large-scale invasions of riparian trees can alter the quantity and quality of allochthonous inputs of leaf litter to streams and thus have the potential to alter stream organic matter dynamics. Non-native saltcedar ( Tamarix sp.) and Russian olive ( Elaeagnus angustifolia ) are now among the most common trees in riparian zones in western North America, yet their impacts on energy flow in streams are virtually unknown.
2. We conducted a laboratory feeding experiment to compare the growth of the aquatic crane fly Tipula (Diptera: Tipulidae) on leaf litter from native cottonwood ( Populus ) and non-native Tamarix and Elaeagnus . Tipula showed positive growth on leaf litter of all three species; however, after 7 weeks, larvae fed Tamarix leaves averaged 1.7 and 2.5 times the mass of those fed Elaeagnus and Populus , respectively. Tipula survival was highest on Populus , intermediate on Tamarix and lowest on Elaeagnus .
3. High Tipula growth on Tamarix probably reflects a combination of leaf chemistry and morphology. Conditioned Tamarix leaf litter had intermediate carbon : nitrogen values (33 : 1) compared to Populus (40 : 1) and Elaeagnus (26 : 1), and it had intermediate proportions of structural carbon (42%) compared to Elaeagnus (57%) and Populus (35%). Tamarix leaves are also relatively small and possibly more easily ingested by Tipula than either Elaeagnus or Populus .
4. Field surveys of streams in the western U.S.A. revealed that Tamarix and Elaeagnus leaf packs were rare compared to native Populus , probably due to the elongate shape and small size of the non-native leaves. Thus we conclude that, in general, the impact of non-native riparian invasion on aquatic shredders will depend not only on leaf decomposition rate and palatability but also on rates of leaf litter input to the stream coupled with streambed retention and subsequent availability to consumers.     

Small dams decrease leaf litter breakdown rates in Mediterranean mountain streams

The damming of rivers and streams alters downstream habitat characteristics and biotic assemblages, and might thus alter stream functioning, although there is not much direct evidence of this impact. In this study we compared breakdown of alder leaves upstream and downstream from 4 small (<1 hm³) dams in 4 Mediterranean mountain streams with no appreciable impact on water temperature and nutrient concentrations. Despite no effect on water characteristics, dams decreased leaf litter breakdown rates. Abundance and biomass of invertebrates and shredders and hyphomycete sporulation rates did not differ between upstream and downstream bags. However, the structure of invertebrate and hyphomycete assemblages did. Especially evident was a drop in limnephilids, which might explain the slower breakdown of leaf litter below dams. These results may help to explain some of the variability found in the literature on the effects of dams on decomposition rates. If dams increase water temperature and nutrient concentrations they may promote faster decomposition, but if dams do not change water characteristics, their impact on detritivore communities may cause slower decomposition rates.     

Hot spots of leaf breakdown within a headwater stream reach: comparing breakdown rates among litter patch types with different macroinvertebrate assemblages

1. Detecting hot spots of litter decomposition will promote understanding of litter processing in a heterogeneous system. To identify hot spots of leaf breakdown within a headwater stream reach, we examined the difference in leaf breakdown rate among four types of litter patches, one that formed in riffles and three that formed in pools (middle, alcove, edge), in different seasons. 2. Middle patches showed the highest breakdown rate in some seasons; the rate in middle patches was 1.5–4 times higher than in the other patches. Thus middle patches can be regarded as hot spots of leaf breakdown in the study reach. This result contrasted with other studies showing higher breakdown rate in riffles than in pools. 3. Significant relationships between abundance of caddisfly shredders and breakdown rate were observed in seasons when the rate differed among patch types. Greater abundance of Lepidostoma seems to be responsible for middle patches being hot spots of leaf breakdown. 4. It is expected that when the proportion of leaves retained in middle patches within a reach is higher, the breakdown rate of the entire reach will be increased. Clarifying how the proportion of leaves retained on middle patches within a reach varies temporally and spatially would improve our understanding of leaf breakdown in headwater streams.     

Effects of diversity loss on ecosystem function across trophic levels and ecosystems: A test in a detritus‐based tropical food web

Abstract We investigated the effects of biodiversity loss across trophic levels and across ecosystems (terrestrial to aquatic) on ecosystem function, in a detritus‐based tropical food web. Diversities of consumers (stream shredders) and resources (leaf litter) were experimentally manipulated by varying the number of species from 3 to 1, using different species combinations, and the effects on leaf breakdown rates were examined. In single‐species shredder treatments, leaf diversity loss affected breakdown rates, but the effect depended on the identity of the leaves remaining in the system: they increased when the most preferred leaf species remained, but decreased when this species was lost (leaf preferences were the same for all shredders). In multi‐species shredder assemblages, breakdown rates were lower than expected from single‐species treatments, suggesting an important role of interspecific competition. This pattern was also evident when oneleaf species was available but not with higher leaf diversity, suggesting that lowered leaf diversity promotes competitive interactions among shredders. The influence of diversity and identity of species across trophic levels and ecosystems on stream functioning points to complex interactions that may well be reflected in other types of ecosystem.     

Understanding of colonization and breakdown of leaves by invertebrates in a tropical stream is enhanced by using biomass as well as count data

We hypothesized that (i) the importance of shredders for leaf breakdown is more evident in terms of their biomass than their abundance, due to the large bodies and high-feeding efficiencies of some typical shredders; (ii) non-shredder invertebrates select more refractory leaves because these are a more stable substrate for colonization or to obtain other forms of food. To test these hypotheses, we performed a decomposition experiment with leaves of contrasting chemical composition in a tropical stream, and determined the changes in the ash-free dry mass (AFDM) of the litter, and the invertebrate abundance and biomass during a 44-day period. The biomass of shredders showed a positive relationship with AFDM remaining, whereas their abundance was unrelated to AFDM. While shredder abundance represented only 4–12% of total invertebrate abundance, shredder biomass constituted 19–36% of total invertebrate biomass. We conclude that (i) shredder biomass expresses better than abundance the role of this guild in the decomposition of leaf detritus, demonstrating that they are important for the functioning of tropical streams; (ii) incubation time rather than stability of leaf litter as a substrate influences colonization by non-shredder invertebrates.     

Stimulation of leaf litter decomposition and associated fungi and invertebrates by moderate eutrophication: implications for stream assessment

1. We investigated the effect of moderate eutrophication on leaf litter decomposition and associated invertebrates in five reference and five eutrophied streams in central Portugal. Fungal parameters and litter N and P dynamics were followed in one pair of streams. Benthic invertebrate parameters that are considered useful in bioassessment were estimated in all streams. Finally, we evaluated the utility of decomposition as a tool to assess stream ecosystem functional integrity. 2. Decomposition of alder and oak leaves in coarse mesh bags was on average 2.3–2.7× faster in eutrophied than in reference streams. This was attributed to stimulation of fungal activity (fungal biomass accrual and sporulation of aquatic hyphomycetes) by dissolved nutrients. These effects were more pronounced for oak litter (lower quality substrate) than alder. N content of leaf litter did not differ between stream types, while P accrual was higher in the eutrophied than in the reference stream. Total invertebrate abundances and richness associated with oak litter, but not with alder, were higher in eutrophied streams. 3. We found only positive correlations between stream nutrients (DIN and SRP) and leaf litter decomposition rates in both fine and coarse mesh bags, associated sporulation rates of aquatic hyphomycetes and, in some cases, total invertebrate abundances and richness. 4. Some metrics based on benthic invertebrate community data (e.g. % shredders, % shredder taxa) were significantly lower in eutrophied than in reference streams, whereas the IBMWP index that is specifically designed for the Iberian peninsula classified all 10 streams in the highest possible class as having ‘very good’ ecological conditions. 5. Leaf litter decomposition was sufficiently sensitive to respond to low levels of eutrophication and could be a useful functional measure to complement assessment programmes based on structural parameters.     

Leaf litter breakdown, microbial respiration and shredder production in metal-polluted streams

1. If species disproportionately influence ecosystem functioning and also differ in their sensitivities to environmental conditions, the selective removal of species by anthropogenic stressors may lead to strong effects on ecosystem processes. We evaluated whether these circumstances held for several Colorado, U.S.A. streams stressed by Zn. 2. Benthic invertebrates and chemistry were sampled in five second–third order streams for 1 year. Study streams differed in dissolved metal concentrations, but were otherwise similar in chemical and physical characteristics. Secondary production of leaf‐shredding insects was estimated using the increment summation and size‐frequency methods. Leaf litter breakdown rates were estimated by retrieving litter‐bags over a 171 day period. Microbial activity on leaf litter was measured in the laboratory using changes in oxygen concentration over a 48 h incubation period. 3. Dissolved Zn concentrations varied eightfold among two reference and three polluted streams. Total secondary production of shredders was negatively associated with metal contamination. Secondary production in reference streams was dominated by Taenionema pallidum. Results of previous studies and the current investigation demonstrate that this shredder is highly sensitive to metals in Colorado headwater streams. Leaf litter breakdown rates were similar between reference streams and declined significantly in the polluted streams. Microbial respiration at the most contaminated site was significantly lower than at reference sites. 4. Our results supported the hypothesis that some shredder species contribute disproportionately to leaf litter breakdown. Furthermore, the functionally dominant taxon was also the most sensitive to metal contamination. We conclude that leaf litter breakdown in our study streams lacked functional redundancy and was therefore highly sensitive to contaminant‐induced alterations in community structure. We argue for the necessity of simultaneously measuring community structure and ecosystem function in anthropogenically stressed ecosystems.     

Differences in processing dynamics of fresh and dried leaf litter in a stream ecosystem

SUMMARY. 1. Although the bulk of litter input to stream ecosystems is in the form of fresh leaves, current understanding of organic matter processing is largely founded on experimental studies made with pre-dried leaves. This paradox points to the critical need for evaluating to what extent those experiments with dried leaves reflect natural litter decomposition.
2. The mass loss rates, patterns of mass loss, and chemical changes during processing of fresh leaf litter were compared with air-dried leaf litter in a stream ecosystem.
3. Although overall mass loss rates were similar between treatments ( k = 0.0213 day⁻¹ and 0.0206 day⁻¹), fresh leaves lost mass at a constant rate, whereas the decay of dried leaves proceeded in two distinct phases. Soluble organic carbon, phosphorus, and potassium were rapidly leached from dried litter, but were largely retained in fresh material for more than a week. Kinetics of concentrations of cellulose and changes in amounts of lignin remaining per leaf pack revealed further differences in decomposition dynamics between treatments, apparently related, either directly or indirectly, to differences in leaching behaviour.
4. Dynamics of nitrogen and protein contents were similar between treatments, indicating that microbial colonization was not greatly delayed on fresh leaves.
5. It is concluded that the retention of labile carbon and nutrients in fresh leaf litter facilitates their utilization by leaf-associated micro-organisms and invertebrates, resulting in an increased importance of biotic processes relative to physical processes such as leaching.
6. At the ecosystem level, retention of carbon and nutrients in streams would be increased, allowing greater overall productivity. Conversely, the availability of labile organic carbon would be reduced in compartments such as the epilithon, fine sediments, and the water column.     

Local Variation in Shredder Distribution can Explain their Oversight in Tropical Streams

Stream shredders play an important role in the breakdown of allochthonous leaf litter—a well-known, key process in temperate headwater streams. In contrast, it has been suggested that litter breakdown in tropical streams is driven by microorganisms, shredders being scarce or absent. We propose that shredders have been overlooked in some tropical streams for two reasons: (1) assuming that tropical shredders belong to the same taxa as temperate ones, without determining the diet of tropical litter fauna; and (2) the small spatial scale of most tropical stream studies, which do not account for intra- and inter-site comparisons. We explored shredder abundance and species richness in six streams in each of two tropical regions, the Australian wet tropics (AWT) and Panama (PAN), finding 734 individuals of 12 shredder species in AWT and 391 individuals of 16 species in PAN. Shredder species richness was positively related to altitude in AWT, but not in PAN. Shredder contribution to total leaf breakdown in the field was 24±3 SE percent in AWT and negligible in PAN, but this was probably due to the unsuccessful colonization of experimental cages by PAN shredders. In the laboratory, shredder contribution to total leaf breakdown was higher than in the field (35%±2 SE in AWT and 64%±3 SE in PAN) and varied with leaf decomposability. Our results support earlier indications that shredders are not scarce or functionally unimportant in the tropics, and suggest that their contribution to litter processing should be determined along altitudinal gradients.
     

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.

     

Effects of nutrient enrichment on boreal streams: invertebrates, fungi and leaf-litter breakdown

1. The effect of nutrient enrichment on structural (invertebrate indices) and functional (leaf‐litter breakdown rates) characteristics of stream integrity was studied in nine boreal streams. 2. The results showed predicted changes in biotic indices and leaf‐litter breakdown along a complex (principal component) nutrient gradient. Biotic indices were better correlated with nutrient effects than leaf‐litter breakdown. 3. Fungal biomass and invertebrate densities in the litter bags were positively correlated with leaf‐litter breakdown, and both were also positively related to the nutrient gradient. 4. Invertebrate community composition influenced breakdown rate. High breakdown rates at one site were associated with the high abundance of the detritivore Asellus aquaticus. 5. This study lends support to the importance of invertebrate and fungi as mediators of leaf‐litter decomposition. However, our study also shows that study design (length of incubation) can confound the interpretation of nutrient‐induced effects on decomposition.     

Structural and functional measures of leaf-associated invertebrates and fungi as predictors of stream eutrophication

Eutrophication is a major threat to freshwater ecosystems worldwide that affects aquatic biota and compromises ecosystem functioning. In this study, we assessed the potential use of leaf decomposition and associated decomposer communities to predict stream eutrophication. Because leaf quality is expected to affect leaf decomposition, we used five leaf species, differing in their initial nitrogen concentration. Leaves of alder, chestnut, plane, oak and eucalyptus were placed in coarse-mesh bags and immersed in six streams along an eutrophication gradient to assess leaf decomposition and the structure of associated decomposer communities. A hump-shaped relationship was established between leaf decomposition and the eutrophication gradient for all leaf species, except for eucalyptus. Invertebrate biomass and density as well as fungal biomass and sporulation were lowest at the extremes of the gradient. Leaf-associated invertebrate and fungal assemblages were mainly structured by stream eutrophication. The percentage of shredders on leaves decreased, whereas the percentage of oligochaeta increased along the eutrophication gradient. The Iberian Biological Monitoring Working Party Index (IBMWP) applied to benthic invertebrates increased from oligotrophic to moderately eutrophic streams and then dropped sharply at highly and hypertrophic streams. Overall, leaf decomposition was a valuable tool to assess changes in stream water quality, and it allowed the discrimination of sites classified by the IBMWP within class I and class IV. Moreover, decomposition of most leaf species responded in a similar way to eutrophication when decomposition was normalized by the quality of leaves.     

Leaf diversity influences in-stream litter decomposition through effects on shredders

1. The functioning of many aquatic ecosystems is controlled by surrounding terrestrial ecosystems. In a view of growing interest in linking biodiversity to ecosystem‐level processes, we examined whether and how leaf diversity influences litter decomposition and consumers in streams. 2. We tested experimentally the hypothesis that the effects of leaf diversity on decomposition are determined by the responses of leaf consumers to resource–habitat heterogeneity. Leaves from three common riparian trees, beech (Fagus sylvatica), hazel (Corylus avellana) and ash (Fraxinus excelsior), were exposed alone and in all possible mixtures of two and three species in a stream. We analysed individual leaf species for decomposition rate, microbial respiration and mycelial biomass, and we determined the species composition, abundance and biomass of shredders in leaf bags. 3. We found that the decomposition of the fastest decomposing leaves (hazel and ash) was substantially stimulated (up to twofold higher than single species leaf packs) in mixtures containing beech leaves, which are refractory. In contrast, the decomposition of beech leaves was not affected by leaf mixing. Such species‐specific behaviour of leaves in species mixtures has been overlooked in previous studies that examined the overall decomposition of litter mixtures. 4. The effects of leaf diversity on decomposition varied with the abundance and biomass of shredders but not with microbial parameters. Beech leaves alone were less attractive to shredders than leaf packs made of hazel, ash or any mixture of species. Moreover, the presence of beech leaves in mixtures led to higher shredder abundance and biomass than we had expected from data from single species exposed alone. Lastly, we found that early instars of the caddisfly Potamophylax (the dominant shredder in terms of biomass) almost exclusively used the toughest material (i.e. beech leaves) to construct their cases. 5. Leaf pack heterogeneity may have altered shredder‐mediated decomposition. Shredders colonising diverse leaf packs benefited from the stable substratum provided by beech leaves, whereas ash and hazel leaves were primarily used as food. Thus, our findings provide strong evidence for an intimate linkage between the diversity of riparian vegetation and aquatic communities.     

A meta‐analysis of the effects of nutrient enrichment on litter decomposition in streams

The trophic state of many streams is likely to deteriorate in the future due to the continuing increase in human‐induced nutrient availability. Therefore, it is of fundamental importance to understand how nutrient enrichment affects plant litter decomposition, a key ecosystem‐level process in forest streams. Here, we present a meta‐analysis of 99 studies published between 1970 and 2012 that reported the effects of nutrient enrichment on litter decomposition in running waters. When considering the entire database, which consisted of 840 case studies, nutrient enrichment stimulated litter decomposition rate by approximately 50%. The stimulation was higher when the background nutrient concentrations were low and the magnitude of the nutrient enrichment was high, suggesting that oligotrophic streams are most vulnerable to nutrient enrichment. The magnitude of the nutrient‐enrichment effect on litter decomposition was higher in the laboratory than in the field experiments, suggesting that laboratory experiments overestimate the effect and their results should be interpreted with caution. Among field experiments, effects of nutrient enrichment were smaller in the correlative than in the manipulative experiments since in the former the effects of nutrient enrichment on litter decomposition were likely confounded by other environmental factors, e.g. pollutants other than nutrients commonly found in streams impacted by human activity. However, primary studies addressing the effect of multiple stressors on litter decomposition are still few and thus it was not possible to consider the interaction between factors in this review. In field manipulative experiments, the effect of nutrient enrichment on litter decomposition depended on the scale at which the nutrients were added: stream reach > streamside channel > litter bag. This may have resulted from a more uniform and continuous exposure of microbes and detritivores to nutrient enrichment at the stream‐reach scale. By contrast, nutrient enrichment at the litter‐bag scale, often by using diffusing substrates, does not provide uniform controllable nutrient release at either temporal or spatial scales, suggesting that this approach should be abandoned. In field manipulative experiments, the addition of both nitrogen (N) and phosphorus (P) resulted in stronger stimulation of litter decomposition than the addition of N or P alone, suggesting that there might be nutrient co‐limitation of decomposition in streams. The magnitude of the nutrient‐enrichment effect on litter decomposition was higher for wood than for leaves, and for low‐quality than for high‐quality leaves. The effect of nutrient enrichment on litter decomposition may also depend on climate. The tendency for larger effect size in colder regions suggests that patterns of biogeography of invertebrate decomposers may be modulating the effect of nutrient enrichment on litter decomposition. Although studies in temperate environments were overrepresented in our database, our meta‐analysis suggests that the effect of nutrient enrichment might be strongest in cold oligotrophic streams that depend on low‐quality plant litter inputs.