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 decomposition and macroinvertebrate communities in headwater streams draining pine and hardwood catchments

Benthic invertebrates, litter decomposition, andlitterbag invertebrates were examined in streamsdraining pine monoculture and undisturbed hardwoodcatchments at the Coweeta Hydrologic Laboratory in thesouthern Appalachian Mountains, USA. Bimonthlybenthic samples were collected from a stream draininga pine catchment at Coweeta during 1992, and comparedto previously collected (1989–1990) benthic data froma stream draining an adjacent hardwood catchment. Litter decomposition and litterbag invertebrates wereexamined by placing litterbags filled with pine ormaple litter in streams draining pine catchments andhardwood catchments during 1992–1993 and 1993–1994. Total benthic invertebrate abundance and biomass inthe pine stream was ca. 57% and 74% that of thehardwood stream, respectively. Shredder biomass wasalso lower in the pine stream but, as a result ofhigher Leuctra spp. abundance, shredderabundance was higher in the pine stream than thehardwood stream. Decomposition rates of both pine andred maple litter were significantly faster in pinestreams than adjacent hardwood streams (p<0.05). Total shredder abundance, biomass, and production weresimilar in maple bags from pine and hardwood streams. However, trichopteran shredder abundance and biomass,and production of some trichopteran taxa such asLepidostoma spp., were significantly higher in maplelitterbags from pine streams than hardwood streams(p<0.05). In contrast, plecopteran shredders(mainly Tallaperla sp.) were more important inmaple litterbags from hardwood streams. Shredderswere well represented in pine litterbags from pinestreams, but low shredder values were obtained frompine litterbags in hardwood streams. Resultssuggest conversion of hardwood forest to pinemonoculture influences taxonomic composition of streaminvertebrates and litter decomposition dynamics. Although the impact of this landscape-leveldisturbance on invertebrate shredder communitiesappeared somewhat subtle, significant differences indecomposition dynamics indicate vital ecosystem-levelprocesses are altered in streams draining pinecatchments.     

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.     

Small stonefly predators affect microbenthic and meiobenthic communities in stream leaf packs

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

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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.
     

Leaf litter degradation in the wave impact zone of a pre-alpine lake

Contrary to streams, decomposition processes of terrestrial leaf litter are still poorly understood in lakes. Here, we examined the decomposition of two leaf species, beech (Fagus sylvatica) and poplar (Populus nigra ‘italica’) in the littoral zone of a large pre-alpine lake at a wave exposed site. We focussed on the shredding impact of benthic invertebrates in a field experiment and on the effects of wave-induced disturbances under field and mesocosm conditions. In contrast to our expectations, benthic shredders did not reveal an important role in leaf processing under the conditions of the field experiment (early spring time, wave impact zone). Strong wave turbulence during storm events significantly reduced leaf mass, FPOM and invertebrate densities at field conditions. Several reasons can explain the low importance of shredders in our field study: (a) phenology of the shredder species, (b) feeding preferences and alternative food sources for gammarids, (c) generally low abundance of the native gammarid species due to the recent occurrence of an invasive predator, (d) disturbance of shredder activity due to high wave impact and (e) relatively low food value of the offered leaves. We suggest that leaf litter decomposition in lakes occurs in specific process domains, which largely depend on the hydraulic characteristics and on water-level fluctuations.     

Leaf litter quality drives the feeding by invertebrate shredders in tropical streams

Amazon and Cerrado‐forested streams show natural fluctuations in leaf litter quantity along the time and space, suggesting a change on litter quality input. These natural fluctuations of leaf litter have repercussion on the organic matter cycling and consequently effects on leaf decomposition in forested streams. The effects of the quantity of leaf litter with contrasting traits on consumption by larvae of shredder insects from biomes with different organic matter dynamics have still been an understudied question. The Trichoptera Phylloicus spp. is a typical shredder in tropical headwater streams and keep an important role in leaf litter decomposition. Here, we assessed the consumption by shredder Phylloicus spp., from Amazonia and Cerrado biomes, on higher (Maprounea guianensis) and lower quality leaves (Inga laurina) in different proportions and quantities. Experiments were performed concomitantly in microcosms approaches, simulating Cerrado and Amazonian streams. Higher leaf consumption occurred in Cerrado microcosms. Litter quantity influenced negatively leaf consumption by shredders in Cerrado, in opposition to Amazonia, where consumption was not affected by leaf quantity. In both sites, we observed higher consumption by shredders in treatment with only M. guianensis and no difference between other treatments with mixture of leaves. In treatment with litter of I. laurina, we noted the use of substrate for case building (due to the higher leaf toughness), affecting the fragmentation process. Therefore, our results indicate that leaf litter quality drives the preference of consumption by Phylloicus larvae in Cerrado and Amazonia streams.     

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).     

Effects of drying regime on microbial colonization and shredder preference in seasonal woodland wetlands

1. Energy budgets of wetlands in temperate deciduous forests are dominated by terrestrially derived leaf litter that decays under different drying conditions depending on autumn precipitation. We compared decay rates and microbial colonization of maple leaves under different inundation schedules in a field experiment, and then conducted a laboratory study on shredder preference. In the field, litter bags either remained submerged (permanent), were moved to a dried part of the basin once and then returned (semi-permanent), or were alternated between wet and dry conditions for 8 weeks (temporary).
2. There was no difference in decay rates among treatments, but leaves incubated under permanent and semi-permanent conditions had higher fungal and bacterial biomass, and lower C : N ratios than those incubated under alternating drying and wetting conditions.
3. To determine the effects of these differences in litter nutritional quality on shredder preference, we conducted a laboratory preference test with larvae of leaf-shredding caddisflies that inhabit the wetland. Caddisflies spent twice as much time foraging on permanent and semi-permanent litter than on litter incubated under temporary conditions.
4. There is considerable variation among previous studies in how basin drying affects litter breakdown in wetlands, and no previous information on shredder preference. We found that frequent drying in a shallow wetland reduces the nutritional quality of leaf litter (lower microbial biomass and nitrogen content), and therefore preference by invertebrate shredders. These results suggest that inter-annual shifts in drying regime should alter detritus processing rates, and hence the mobilization of the energy and nutrients in leaf litter to the wetland food web.     

Effects of burial on leaf litter quality,microbial conditioning and palatability to three shredder taxa

Summary 1. Heterotrophic microorganisms are crucial for mineralising leaf litter and rendering it more palatable to leaf‐shredding invertebrates. A substantial part of leaf litter entering running waters may be buried in the streambed and thus be exposed to the constraining conditions prevailing in the hyporheic zone. The fate of this buried organic matter and particularly the role of microbial conditioning in this habitat remain largely unexplored. 2. The aim of this study was to determine how the location of leaf litter within the streambed (i.e. at the surface or buried), as well as the leaf litter burial history, may affect the leaf‐associated aquatic hyphomycete communities and therefore leaf consumption by invertebrate detritivores. We tested the hypotheses that (i) burial of leaf litter would result in lower decomposition rates associated with changes in microbial assemblages compared with leaf litter at the surface and (ii) altered microbial conditioning of buried leaf litter would lead to decreased quality and palatability to their consumers, translating into lower growth rates of detritivores. 3. These hypotheses were tested experimentally in a second‐order stream where leaf‐associated microbial communities, as well as leaf litter decomposition rates, elemental composition and toughness, were compared across controlled treatments differing by their location within the streambed. We examined the effects of the diverse conditioning treatments on decaying leaf palatability to consumers through feeding trials on three shredder taxa including a freshwater amphipod, of which we also determined the growth rate. 4. Microbial leaf litter decomposition, fungal biomass and sporulation rates were reduced when leaf litter was buried in the hyporheic zone. While the total species richness of fungal assemblages was similar among treatments, the composition of fungal assemblages was affected by leaf litter burial in sediment. 5. Leaf litter burial markedly affected the food quality (especially P content) of leaf material, probably due to the changes in microbial conditioning. Leaf litter palatability to shredders was highest for leaves exposed at the sediment surface and tended to be negatively related to leaf litter toughness and C/P ratio. In addition, burial of leaf litter led to lower amphipod growth rates, which were positively correlated with leaf litter P content. 6. These results emphasise the importance of leaf colonisation by aquatic fungi in the hyporheic zone of headwater streams, where fungal conditioning of leaf litter appears particularly critical for nutrient and energy transfer to higher trophic levels.     

Early stages of leaf decomposition are mediated by aquatic fungi in the hyporheic zone of woodland streams

1. Leaf litter constitutes the major source of organic matter and energy in woodland stream ecosystems. A substantial part of leaf litter entering running waters may be buried in the streambed as a consequence of flooding and sediment movement. While decomposition of leaf litter in surface waters is relatively well understood, its fate when incorporated into river sediments, as well as the involvement of invertebrate and fungal decomposers in such conditions, remain poorly documented. 2. We tested experimentally the hypotheses that the small interstices of the sediment restrict the access of the largest shredders to buried organic matter without compromising that of aquatic hyphomycetes and that fungal decomposers in the hyporheic zone, at least partly, compensate for the role of invertebrate detritivores in the benthic zone. 3. Alder leaves were introduced in a stream either buried in the sediment (hyporheic), buried after 2 weeks of exposure at the sediment surface (benthic‐hyporheic), or exposed at the sediment surface for the entire experiment (benthic). Leaf decomposition was markedly faster on the streambed surface than in the two other treatments (2.1‐ and 2.8‐fold faster than in the benthic‐hyporheic and hyporheic treatments, respectively). 4. Fungal assemblages were generally less diverse in the hyporheic habitat with a few species tending to be relatively favoured by such conditions. Both fungal biomass and sporulation rates were reduced in the hyporheic treatment, with the leaves subject to the benthic‐hyporheic treatment exhibiting an intermediate pattern. The initial 2‐week stage in the benthic habitat shaped the fungal assemblages, even for leaves later subjected to the hyporheic conditions. 5. The abundance and biomass of shredders drastically decreased with burial, except for Leuctra spp., which increased and was by far the most common leaf‐associated taxon in the hyporheic zone. Leuctra spp. was one of the rare shredder taxa displaying morphological characteristics that increased performance within the limited space of sediment interstices. 6. The carbon budgets indicated that the relative contributions of the two main decomposers, shredders and fungi, varied considerably depending on the location within the streambed. While the shredder biomass represented almost 50% of the initial carbon transformed after 80 days in the benthic treatment, its contribution was <0.3% in the hyporheic one and 2.0% in the combined benthic‐hyporheic treatment. In contrast, mycelial and conidial production in the permanently hyporheic environment accounted for 12% of leaf mass loss, i.e. 2–3 times more than in the two other conditions. These results suggest that the role of fungi is particularly important in the hyporheic zone. 7. Our findings indicate that burial within the substratum reduces the litter breakdown rate by limiting the access of both invertebrate and fungal decomposers to leaves. As a consequence, the hyporheic zone may be an important region of organic matter storage in woodland streams and serve as a fungal inoculum reservoir contributing to further dispersal. Through the temporary retention of litter by burial, the hyporheic zone must play a significant role in the carbon metabolism and overall functioning of headwater stream ecosystems.     

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.

     

Role of transported particulate organic matter in the macroinvertebrate colonization of litter bags in streams

1. The exposure of mesh litter bags has been widely used to investigate the role of benthic macroinvertebrates in leaf processing in freshwaters. In this sense, several studies have related litter bag breakdown rates to the presence of colonizing invertebrates. A possible confounding factor in such experiments is that the litter bags trap suspended particulate organic matter that can itself attract invertebrate colonists, irrespective of the intended experimental treatment.
2. We attempted to quantify the accumulation of particulate organic matter (POM) within litter bags and to investigate its possible impact on macroinvertebrate density and richness. In seven headwater forested streams we exposed mesh bags filled either with beech leaves ( Fagus sylvatica ) or with plastic strips of an equal surface area.
3. Principal component analysis (PCA) showed that bag type and stream were the main explanatory variables for invertebrate colonization and POM accumulation within the bags. In contrast, there was little variation among sampling dates (6.4% of the total inertia).
4. The accumulated POM within the bags was substantial (up to 8.83 g ash-free dry mass (AFDM)) but highly variable among sites (mean from 0.32 to 4.58 g AFDM). At each of the seven sites, both richness and abundance of invertebrates were positively correlated with the mass of accumulated POM in bags. Macroinvertebrate colonization (notably taxon richness) was directly linked with the quantity of POM accumulated.
5. Our findings provide evidence of a potential pitfall in linking invertebrates to litter processing in mesh bags, particularly when large amounts of POM, entrained in stream flow, accumulate within the bags. An evaluation of the POM mass trapped in litter bags could account for the erratic patterns sometimes observed in their colonization by invertebrates.     

大型底栖无脊椎动物对溪流中三角枫落叶分解的影响

采用复合网袋法,研究了冬季南京紫金山三角枫落叶在无干扰溪流和生态恢复溪流中的分解过程.结果表明：112 d后,三角枫落叶无灰干质量剩余率为31％～62％,分解速率符合指数衰减模型(P<0.05).在生态恢复溪流和无干扰溪流的流水生境中, 三角枫落叶的分解速率分别为0.0030 d^-1和0.0064 d^-1,静水生境中分别为0.0018 d^-1和0.0016 d^-1.流水生境中,无干扰溪流网袋内的大型底栖动物多度和生物量显著高于恢复溪流(P<0.05),而静水中无显著差异(P>0.05).无干扰溪流中的撕食者多度比例最高(70.4%),以栉水虱为主;生态恢复溪流中滤食者的多度比例最高(37.8%),以长跗摇蚊属为主.流水生境中三角枫落叶的分解速率与撕食者物种丰富度和多度相关性显著(P<0.01),而与生物量的相关性不显著(P>0.05).说明冬季溪流中撕食者的物种丰富度和多度决定三角枫落叶的分解速率.     

Effects of salmon carcasses on experimental stream ecosystems in Hokkaido,Japan

The effects of salmon carcasses on dissolved nutrients, epilithic production, leaf decomposition rates, and aquatic invertebrates were examined using 10-m-long artificial channels fed by an adjacent natural stream in Hokkaido, northern Japan. Bags containing maple leaf litter were placed in nine channels, each of which was assigned to one of three treatments with three replicates, for 6 weeks in fall 2002. The three treatments were: (1) salmon carcasses+invertebrates, (2) invertebrates only, and (3) control (no salmon carcasses or invertebrates added). Nutrient concentrations, biomass of epilithic algae (chlorophyll), leaf weight loss, abundance and biomass of invertebrates in the leaf packs were compared among the three treatments at 14, 27, and 40 days after the beginning of the experiment. The NH₄⁺ in stream water and chlorophyll concentrations of epilithic algae were higher in the salmon treatment than the other treatments, and the maple leaves decomposed faster in the salmon treatment than in the other treatments. Moreover, the N content of the leaves was highest and the C/N ratio was lowest in the salmon treatment, although not significantly so. The abundance and biomass of the dominant leaf-shredding invertebrate Goerodes satoi did not differ between the first two treatments. However, the stable N isotope ratio in G. satoi was nearly 3 higher in the salmon treatment, suggesting that around 20% of salmon-derived N was taken up by this shredder. Our results indicate that salmon carcasses affect stream ecosystems directly by enhancing primary production, indirectly by accelerating woody leaf decomposition, and finally by incorporating into the food web primary consumers that utilize fertilized woody leaves.     

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     

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.     

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.     

Spatial Variability of Plant Litter Decomposition in Stream Networks: from Litter Bags to Watersheds

The decomposition of plant litter plays a fundamental role in the cycling of carbon and nutrients and is driven by complex interactions of biological and physical controls, yet little is known about its variability and controls across spatial scales. Here we address the indirect effects of riparian canopy cover on litter decomposition and decomposers and their variability within a set of hierarchical scales (watershed, stream segments and reaches) controlling for confounding factors that could co-vary with canopy cover (for example, temperature and nutrients), in high-altitude subtropical streams. Total, microbial and invertebrate-driven decomposition rates were approximately 1.4–6.6 times higher in closed-canopy than in open-canopy watersheds. Riparian canopy cover accounted for 62–69% of total variability of decomposition rates and indirectly (via light availability and litter inputs) promoted fungal facilitation of shredders through leaf litter conditioning. In contrast to what we expected, much of the spatial variability in the decomposition occurred at smaller scale (4–20% of total variability among reaches versus <1% among watersheds) and coincided with the greatest variability in shredder abundance and fungal biomass (70 and 17% among reaches, respectively). We conclude that riparian canopy cover may be an important control of natural variability of litter decomposition at the watershed scale through its effects on fungal decomposers and shredder consumption. We also provide evidence of higher reach and minor watershed variability of litter decomposition in stream networks. Our results point to the importance of identifying the sources of natural variability of decomposition and how they interact within and among spatial scales.