Non‐additive effects of litter mixing are suppressed in a nutrient‐enriched stream

Investigations of how species compositional changes interact with other aspects of global change, such as nutrient mobilization, to affect ecosystem processes are currently lacking. Many studies have shown that mixed species plant litters exhibit non‐additive effects on ecosystem functions in terrestrial and aquatic systems. Using a full‐factorial design of three leaf litter species with distinct initial chemistries (carbon:nitrogen; C:N) and breakdown rates (Liriodendron tulipifera, Acer rubrum and Rhododendron maximum), we tested for additive and non‐additive effects of litter species mixing on breakdown in southeastern US streams with and without added nutrients (N and phosphorus). We found a non‐additive (antagonistic) effect of litter mixing on breakdown rates under reference conditions but not when nutrient levels were elevated. Differential responses among single‐species litters to nutrient enrichment contributed to this result. Antagonistic litter mixing effects on breakdown were consistent with trends in litter C:N, which were higher for mixtures than for single species, suggesting lower microbial colonization on mixtures. Nutrient enrichment lowered C:N and had the greatest effect on the lowest‐ (R. maximum) and the least effect on the highest‐quality litter species (L. tulipifera), resulting in lower interspecific variation in C:N. Detritivore abundance was correlated with litter C:N in the reference stream, potentially contributing to variation in breakdown rates. In the nutrient‐enriched stream, detritivore abundance was higher for all litter and was unrelated to C:N. Thus, non‐additive effects of litter mixing were suppressed by elevated streamwater nutrients, which increased nutrient content of all litter, reduced variation in C:N among litter species and increased detritivore abundance. Nutrients reduced interspecific variation among plant litters, the base of important food web pathways in aquatic ecosystems, affecting predicted mixed‐species breakdown rates. More generally, world‐wide mobilization of nutrients may similarly modify other effects of biodiversity on ecosystem processes.     

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)     

Salamander diversity alters stream macroinvertebrate community structure

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Fungi are involved in the effects of litter mixtures on consumption by shredders

1. Decomposition of litter mixtures in both terrestrial and aquatic ecosystems often shows non‐additive diversity effects on decomposition rate, generally interpreted in streams as a result of the feeding activity of macroinvertebrates. The extent to which fungal assemblages on mixed litter may influence consumption by macroinvertebrates remains unknown. 2. We assessed the effect of litter mixing on all possible three‐species combinations drawn from four tree species (Alnus glutinosa, Betula pendula, Juglans regia and Quercus robur) on both fungal assemblages and the rate of litter consumption by a common shredder, Gammarus fossarum. After a 9‐week inoculation in a stream, batches of leaf discs were taken from all leaf species within litter mixture combinations. Ergosterol, an indicator of fungal biomass, and the composition of fungal assemblages, assessed from the conidia released, were determined, and incubated litter offered to G. fossarum in a laboratory‐feeding experiment. 3. Mixing leaf litter species enhanced both the Simpson’s index of the fungal assemblage and the consumption of litter by G. fossarum, but had no clear effect on mycelial biomass. Specifically, consumption rates of J. regia were consistently higher for mixed‐species litter packs than for single‐species litter. In contrast, the consumption rates of B. pendula were not affected by litter mixing, because of the occurrence of both positive and negative litter‐mixing effects in different litter species combinations that counteracted each other. 4. In some litter combinations, the greater development of some fungal species (e.g. Clavariopsis aquatica) as shown by higher sporulation rates coincided with increased leaf consumption, which may have resulted from feeding preferences by G. fossarum for these fungi. 5. Where litter mixture effects on decomposition rate are mediated via shredder feeding, this could be due to indirect effects of the fungal assemblage.     

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.     

Community composition and diversity of riparian forests regulate decomposition of leaf litter in stream ecosystems

Riparian forest plantings are a well‐established restoration technique commonly used to stabilize banks and intercept nutrient flow from adjacent agricultural fields. Tree species planted for these efforts may not reflect mature forest communities within the same region. Given contemporary research on links between biodiversity and ecosystem functioning, we conducted a leaf‐litter decomposition study to investigate how mixing of detrital resources that reflect forest community composition would regulate in‐stream leaf litter. Leaf litter bags containing material from a mature forest (Liriodendron tulipifera, Acer rubrum, Quercus rubra, full factorial treatments = 7) and a restored riparian forest (Cornus sericea, Fraxinus pennsylvanica, Platanus occidentalis, full factorial treatments = 7) were deployed in a stream reach that experienced riparian reforestation in 2004. Litter from the restored riparian community had less mass remaining (45.28 ± 2.27%) than that from the mature riparian community (54.95 ± 2.19%) after 5 weeks. In addition, mixed litter treatments in the restored riparian community had less mass remaining (40.54 ± 2.37%) than single‐species treatments (51.80 ± 4.05%), a pattern not observed in the mature forest community. Results highlight the importance of planting mixed‐species assemblages as this structure may regulate processes such as decomposition and food‐web structure, processes often not targeted in the restoration plans.     

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.     

Leaf decomposition in an experimentally acidified stream channel

Decomposition of Alnus rugosa and Myrica Gale leaves immersed in artificial stream channels fed by a small headwater creek was followed over a three month period. At the end of experiment, remaining weights of both leaf types confined in litter bags were significantly higher after immersion in experimentally acidified water (pH 4.0) than when immersed in control water (pH 6.2–7.0). For both types of leaves and for all sampling times, there was generally no difference in the C:N ratios between leaves in acidified and those in control water. In control water, oxygen uptake by microorganism on A. rugosa leaves was significantly higher after 46 days of immersion, whereas differences between treatments appeared only after 69 days for M. Gale leaves. Transfer of A. rugosa leaves from acid to control water led to a rapid increase in microbial activity; this increased activity was reflected in a fast weight loss of the leaves. For both leaf types, total numbers of macroinvertebrates were usually higher in litter bags immersed in control water. Macroinvertebrates colonizing the litter bags were mainly collector-gatherers: Chironomidae were numerically dominant in control leaf packs whereas Oligochaeta dominated in acid leaf packs. Macroinvertebrate biomass in M. Gale litter was higher in control than in acidified water, which contrasted with macroinvertebrate biomass in A. rugosa leaf packs which was not significantly different between treatments. Macroinvertebrate contribution to the breakdown of leaf litter was thus considered less important than the microbial contribution. This study demonstrated that decomposition of leaf litter in acidic headwater streams can be seriously reduced, mainly as a result of a lower microbial activity.     

Similar breakdown rates and benthic macroinvertebrate assemblages on native and Eucalyptus globulus leaf litter in Californian streams

1. Eucalyptus globulus, a tree species planted worldwide in many riparian zones, has been reported to affect benthic macroinvertebrates negatively. Although there is no consensus about the effects of Eucalyptus on aquatic macrobenthos, its removal is sometimes proposed as a means of ecological restoration. 2. We combined the sampling of macroinvertebrates with measurement of the colonisation of leaf packs in mesh bags, to examine the effects of riparian Eucalyptus and its litter on benthic macroinvertebrates in three small streams in California, U.S.A. Each stream included one reach bordered by Eucalyptus (E‐site) and a second bordered by native vegetation (N‐site). 3. The macrobenthos was sampled and two sets of litter bags were deployed at each site: one set with Eucalyptus litter (Euc‐bags) and one with mixed native tree litter (Nat‐bags) containing Quercus, Umbellularia, Acer and Alnus. Bags were exposed for 28, 56 and 90 days and this experiment was repeated in the autumn, winter and spring to account for effects of changing stream flow and insect phenology. 4. Litter input (average dry mass: 950 g m^?2 year^?1 in E‐sites versus 669 g m^?2 year^?1 in N‐sites) was similar, although in‐stream litter composition differed between E‐ and N‐sites. Litter broke down at similar rates in Euc‐bags and Nat‐bags (0.0193 day^?1 versus 0.0134 day^?1), perhaps reflecting the refractory nature of some of the leaves of the native trees (Quercus agrifolia). 5. Summary metrics for macroinvertebrates (taxon richness, Shannon diversity, pollution tolerance index) did not differ significantly between the E and N sites, or between Euc‐bags and Nat‐bags. No effect of exposure time or site was detected by ordination of the taxa sampled. However, distinct seasonal ordination clusters were observed in winter, spring and autumn, and one of the three streams formed a separate cluster. 6. The presence of Eucalyptus was less important in explaining the taxonomic composition of the macrobenthos than either ‘season’ or ‘stream’. Similarly, these same two factors (but not litter species) also helped explain the variation in leaf breakdown. We conclude that patches of riparian Eucalyptus and its litter have little effect on stream macrobenthos in this region.     

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.     

Leaf Decomposition and Stream Macroinvertebrate Colonisation of Japanese Knotweed,an Invasive Plant Species

Japanese knotweed (Fallopia japonica Houtt. Ronse Decrane ) is a highly invasive exotic plant that forms monocultures in riparian areas, effectively reducing plant diversity. This change in riparian plant composition alters the allocthonous input of leaf litter into adjacent streams. A field experiment was completed to understand how leaf decomposition and macroinvertebrate colonisation associated with the incorporation of exotic leaf litter. Leaf packs of Japanese knotweed, native alder (Alnus incana L.), native cottonwood (Populus trichocarpa Torr . and Gray ), and two additional mixed pack types (alder and cottonwood; alder, cottonwood, and Japanese knotweed) were placed into a 50 m stream reach in Clear Creek, Idaho, and removed over a three‐month period. Leaf decomposition and macroinvertebrate assemblages were similar between leaf types, despite differences in nitrogen and phosphorus content. The diversity of leaf types within a given leaf pack also had no effect on leaf decomposition or macroinvertebrate dynamics. These findings suggest that allochthonous inputs of Japanese knotweed fulfill a detrital function similar to that of native leaf litter. (© 2008 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Riparian land use and the relationship between the benthos and litter decomposition in tropical montane streams

1. Although stream–catchment interactions have been analysed in some detail in temperate environments, little is known about the effects of land‐use changes in the tropics. Here, we analyse differences in benthic communities (macroinvertebrates and fungi) under two contrasting land uses (mature secondary forest and pasture) in montane streams in north‐western Ecuador and their influence on the rates of litter processing. 2. Between 2005 and 2006, we used a combination of coarse and fine mesh bags to study the relative contribution of macroinvertebrates and fungi to processing of two types of litter, Alnus acuminata and Inga spectabilis, in three‐first‐order streams running through mature secondary montane forests and adjacent downstream reaches running through pastures. At the same time, we characterised the assemblages of shreddering macroinvertebrates and fungi communities and the litter processing rates in stream reaches under both vegetation types. 3. Litter processing rates attributable to invertebrate feeding (coarse mesh bags) were significantly slower in streams running through pastures. Nevertheless, shredder diversity and richness were similar between pasture and forest sections, while shredder abundance was significantly higher in forest streams (mainly Phylloicus sp. :Trichoptera). Fungal reproductive activity and litter processing rates were low (fine mesh bags) and did not differ significantly between pasture and forest stream reaches. 4. Phylloicus sp. abundance was the best predictor of the percentage of litter remaining in coarse mesh bags across pasture and forest sites. Neither shredder diversity nor their species richness was a significant predictor of mass loss, as most of the decomposition was performed by a single keystone species. Although litter decomposition by microbial decomposers was low, fungal biomass (but not diversity) was the best variable explaining the percentage of litter remaining in fine mesh bags. 5. Our data suggest that, in these Neotropical montane streams, land use can have a significant impact on the rates of critical ecosystem processes, such as litter decomposition. In this study, this effect was not mediated by a major shift in the structure of the benthos, but by a decrease in the abundance and relative representation of a single species whose life history makes it critical to litter processing. 6. This study highlights the significant role that macroinvertebrate fauna can have in the processing of litter in Neotropical streams and the predominant role that single species can have in terms of controlling stream ecosystem‐level processes. Understanding the extent to which these patterns affect the long‐term and large‐scale functioning of stream ecosystems still needs further research and will become increasingly important in terms of managing lotic ecosystems in the context of rapid land‐use change.     

The Effect of Macroinvertebrate Exclusion on Leaf Breakdown Rates in a Tropical Headwater Stream1

The role of macroinvertebrates in the process of leaf breakdown is well studied in temperate streams, but less is known about their role in the tropics. We investigated the effect of reducing macroinvertebrate access to leaf material on leaf breakdown rates in a forested headwater stream in the Luquillo Experimental Forest, Puerto Rico. We measured leaf mass loss using fine and coarse mesh bags over 12 weeks for two common riparian species: Cecropia schreberiana (Moraceae) and Dacryodes excelsa (Burseraceae). Coarse mesh allowed freshwater shrimp and other macroinvertebrates to access leaf material, while fine mesh did not. Leaf breakdown rates did not differ between C. schreberiana and D. excelsa in coarse mesh bags (?0.0375/day vs. ?0.0395/day, respectively), but C. schreberiana breakdown was significantly slower than D. excelsa in fine mesh bags (?0.0159/day vs. ?0.0266/day). Additionally, breakdown in fine mesh bags was significantly slower compared to coarse mesh bags for C. schreberiana, but less so for D. excelsa. Breakdown rates for all treatments were fast relative to those in temperate‐zone streams indicating that both macroinvertebrates and macroinvertebrate‐independent processing can strongly influence leaf decomposition in tropical streams. The difference between C. schreberiana and D. excelsa indicates that the effect of macroinvertebrate exclusion can change with leaf type.     

Annelid sperm and fertilization biology

This paper provides data on fine particulate organic matter (FPOM) and macroinvertebrates associated to natural and artificial leaf packs in a small woodland stream (Schlaube, Brandenburg). Macroinvertebrate colonisation and the dynamics of FPOM were studied in oven-dried alder leaf packs, air-dried alder leaf packs and packs with artificial leafshaped substrate exposed in the stream during a 68-day period. The importance of FPOM as a potential food source for macroinvertebrates especially in artificial leaf packs was evaluated. Changes in the quantity as well as in the chemical composition of the accumulating FPOM (>63 and <63 μm) was determined using soluble carbohydrates, proteins and chlorophyll a as parameters of the nutritional quality. Mass loss and the chemical changes of alder leaves during the decompositional process were also described. The loss of soluble carbohydrates due to leaching was more rapid in oven-dried alder leaf packs than in air-dried ones. After 3 days of leaf pack exposure weight loss of oven-dried and air-dried leaf packs was nearly comparable, as the similar decay coefficients, k = 0.0228 (oven-dried leaf packs) and k = 0.0214 (air-dried leaf packs), respectively, show. The amount of FPOM per unit leaf area constantly increased in artificial packs, although it remained below that of alder leaf packs at all sampling dates. The nutritional quality of FPOM <63 μm was constantly greater than that of FPOM >63 μm and decreased in both size-fractions with length of exposure. Referring to leaf area the abundance of macroinvertebrates continually increased in all packs till the end of exposure, whereas the numbers in artificial packs remained below that in alder leaf packs. The taxonomic composition of all treatments was very similar with Gammarus pulex being the most abundant taxon in all packs until day 42, while afterwards the caddis fly genus Hydropsyche gained in importance. The amphipod Gammarus pulex in general did not show a preference for air-dried alder leaf packs compared to oven-dried alder leaf and artificial packs. Corresponding dynamics of macroinvertebrate colonisation and FPOM content in artificial packs support the hypothesis that FPOM functions not only as an important food source for macroinvertebrates including gammarideans but also as a control mechanism of macroinvertebrate abundance in stream habitats. Even if the accumulation of FPOM and drifting macroinvertebrates might be influenced by the same abiotic factor (e.g. by reduction in stream velocity inside the packs) it is quite unlikely that only physical properties caused the invertebrates to stay.     

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.

     

An assessment of the sources of error in estimations of bulk sedimentary pigment concentrations and its implications for trophic status assessment

Rates of leaf litter processing and densities of macroinvertebrates in leaf packs were compared at two sites that differed in catchment logging history. The processing rate of leaves of sugar maple (Acer saccharum Marsh.) was significantly faster in a stream draining a catchment that had been logged about 20 years ago than in one that had been undisturbed for 80 years. The faster processing rate was accompanied by significantly higher leaf pack densities of total macroinvertebrates, shredders, and collector-gatherers. The higher densities of leaf pack macroinvertebrates were apparently a result of differences in tree species between the two catchments. These differences resulted in greater inputs of fast-decomposing leaf litter to the stream draining the disturbed catchment and in smaller amounts of leaf litter remaining in the stream draining the disturbed catchment by spring when this study was conducted. The Unit is jointly sponsored by the West Virginia Department of Natural Resources, West Virginia University, the U.S. Fish and Wildlife Service, and the Wildlife Management Institute. The Unit is jointly sponsored by the West Virginia Department of Natural Resources, West Virginia University, the U.S. Fish and Wildlife Service, and the Wildlife Management Institute.     

A Comparative Study of Leaf Breakdown of Three Native Tree Species in a Slowly‐Flowing Headwater Stream in the Colombian Andes

The dynamics of leaf breakdown in a headwater Colombian stream were evaluated for the native tree species Myrsine guianensis, Cupania latifolia and Nectandra lineatifolia using coarse and fine mesh litter bags. Ten bags of each species (five of each mesh size) were retrieved from the stream at 1, 8, 15, 30, 60 and 120 days. k values ranged from 0.0008 to 0.0058 day^–1 and density of macroinvertebrates from 35 to 55 individuals per leaf bag, peaking at day 8. Myrsine guianensis degraded more rapidly than the other species for both coarse and fine mesh bags. This species and Nectandra lineatifolia presented differences in k values between coarse and fine mesh bags, suggesting that macroinvertebrates influenced the decay rate. Despite the low densities of macroinvertebrates found, shredders represented 12.7% of individuals and 50 to 68% of the invertebrate biomass in bags, indicating that this functional feeding group was an important component of fauna associated with litter breakdown in this first order tropical stream. (© 2007 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Nutrients stimulate leaf breakdown rates and detritivore biomass: bottom-up effects via heterotrophic pathways

Most nutrient enrichment studies in aquatic systems have focused on autotrophic food webs in systems where primary producers dominate the resource base. We tested the heterotrophic response to long-term nutrient enrichment in a forested, headwater stream. Our study design consisted of 2 years of pretreatment data in a reference and treatment stream and 2 years of continuous nitrogen (N) + phosphorus addition to the treatment stream. Studies were conducted with two leaf species that differed in initial C:N, Rhododendron maximum (rhododendron) and Acer rubrum (red maple). We determined the effects of nutrient addition on detrital resources (leaf breakdown rates, litter C:N and microbial activity) and tested whether nutrient enrichment affected macroinvertebrate consumers via increased biomass. Leaf breakdown rates were ca. 1.5 and 3× faster during the first and second years of enrichment, respectively, in the treatment stream for both leaf types. Microbial respiration rates of both leaf types were 3× higher with enrichment, and macroinvertebrate biomass associated with leaves increased ca. 2–3× with enrichment. The mass of N in macroinvertebrate biomass relative to leaves tended to increase with enrichment up to 6× for red maple and up to 44× for rhododendron leaves. Lower quality (higher C:N) rhododendron leaves exhibited greater changes in leaf nutrient content and macroinvertebrate response to nutrient enrichment than red maple leaves, suggesting a unique response by different leaf species to nutrient enrichment. Nutrient concentrations used in this study were moderate and equivalent to those in streams draining watersheds with altered land use. Thus, our results suggest that similarly moderate levels of enrichment may affect detrital resource quality and subsequently lead to altered energy and nutrient flow in detrital food webs. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

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.     

Habitat Use of Juvenile Fish in the Lower Danube and the Danube Delta: Implications for Ecotone Connectivity

Functional processes in freshwater ecosystems are highly influenced by acidic conditions. Foodwebs are affected and macroinvertebrate species diversity is decreased. This study aims to investigate leaf decomposition at very low pH in the acidic Banyupahit–Banyuputih river originating from the acidic crater lake Kawah Ijen in Indonesia. Leaf decomposition experiments were carried out for 200 days in the acidic river at pHs of approximately 0.7, 2.3 and 3.0 and in the neutral Kali Sengon river, using leaves from teak, Tectona grandis, and bamboo, Bambusa sp. Two different types of leaf packs were used: fine mesh size packs were used to exclude macroinvertebrates and coarse mesh size packs allowed macroinvertebrate colonization. Clear differences in decomposition rate were observed between the neutral Kali Sengon and the acidic Banyupahit–Banyuputih river with decomposition in the Kali Sengon river proceeding significantly faster for both leaf types. In the Kali Sengon k values (d⁻¹) over 46 days were 0.0202 for fine teak, 0.0236 for coarse teak, 0.0114 for fine bamboo and 0.0151 for coarse bamboo. No significant differences were observed between the three sites in the acidic Banyupahit–Banyuputih river with k values of 0.0034–0.0066 for fine teak, 0.0002–0.0057 for coarse teak, 0.0029–0.0054 for fine bamboo and 0.0000–0.0068 for coarse bamboo. Moreover, no clear adaptation of macroinvertebrates or microbes to low pH conditions could be detected. The coarse mesh leaf packs in the neutral Kali Sengon river revealed that macroinvertebrates are important in the breakdown process. Fine mesh packs revealed that microbial activity is depressed under acidic conditions. Based on this evidence, we conclude that the toxicity at low pH conditions, and probably also the precipitation of metals on the leaf material, seriously affects leaf decomposition.