Two microcrustaceans affect microbial and macroinvertebrate‐driven litter breakdown

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Experimental shading alters leaf litter breakdown in streams of contrasting riparian canopy cover

1. Headwater stream ecosystems are primarily heterotrophic, with allochthonous organic matter being the dominant energy. However, sunlight indirectly influences ecosystem structure and functioning, affecting microbial and invertebrate consumers and, ultimately, leaf litter breakdown. We tested the effects of artificial shading on litter breakdown rates in an open‐canopy stream (high ambient light) and a closed‐canopy stream (low ambient light). We further examined the responses of invertebrate shredders and aquatic hyphomycetes to shading to disentangle the underlying effects of light availability on litter breakdown. 2. Litter breakdown was substantially slower for both fast‐decomposing (alder, Alnus glutinosa) and slow‐decomposing (beech, Fagus sylvatica) leaf litters in artificially shaded stream reaches relative to control (no artificial shading) reaches, regardless of stream type (open or closed canopy). 3. Shredder densities were higher on A. glutinosa than on F. sylvatica litter, and shading had a greater effect on reducing shredder densities associated with A. glutinosa than those associated with F. sylvatica litter in both stream types. Fungal biomass was also negatively affected by shading. Results suggest that the effects of light availability on litter breakdown rates are mediated by resource quality and consumer density. 4. Results from feeding experiments, where A. glutinosa litter incubated under ambient light or artificial shade was offered to the shredder Gammarus fossarum, suggest that experimental shading and riparian canopy openness influenced litter palatability interactively. Rates of litter consumption by G. fossarum were decreased by experimental shading in the open‐canopy stream only. 5. The results suggest that even small variations in light availability in streams can mediate substantial within‐stream heterogeneity in litter breakdown. This study provides further evidence that changes in riparian vegetation, and thus light availability, influence organic matter processing in heterotrophic stream ecosystems through multiple trophic levels.     

The Role of Early Fungal Colonizers in Leaf‐Litter Decomposition in Portuguese Streams Impacted by Agricultural Runoff

We conducted a transplant experiment between two streams in NW Portugal impacted by agricultural runoff, mainly differing in phosphate concentration, to determine whether fungi on decomposing leaves would adapt to the new environment or would be replaced by fungi of the recipient stream. The most nutrient enriched stream had lower fungal diversity but faster leaf decomposition. Leaf transplantation did not alter fungal activity or species dominance. Multidimensional scaling ordination of fungal communities, from DNA fingerprint or conidial production, revealed that transplanted communities resembled more those of the original stream than the recipient stream. Results suggest that early fungal colonizers will determine the development and activity of fungal communities on decomposing leaves in streams impacted by agricultural practices. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Leaf litter decomposition in remote oceanic island streams is driven by microbes and depends on litter quality and environmental conditions

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Leaf litter breakdown in a Mediterranean stream characterised by travertine precipitation

1. Breakdown of four leaf species ( Platanus orientalis , Populus nigra , Salix atrocinerea , Rubus ulmifolius ) was studied in a Mediterranean second-order stream characterised by abundant travertine precipitation, a history of fire in its catchment, and a recently revegetated alluvial corridor.
2. Compared to breakdown rates reported in the literature for congeneric species, breakdown of the four species was slow (k = 0.0024–0.0069 day⁻¹ for the tree species, and 0.0103 and 0.0111 day⁻¹ for Rubus ), in spite of high water temperatures, indicating that the travertine layer that quickly covered submerged leaves impeded decomposer activity and physical fragmentation losses.
3. Breakdown rates nevertheless differed between leaf species in a predictable manner, suggesting that the observed mass loss was largely due to biological processes.
4. The observed tendency towards increasing leaf nitrogen and phosphorus concentrations during breakdown suggests that microorganisms were actively involved in leaf breakdown; however, this interpretation must be viewed with caution because of potentially confounding effects by nutrients contained in the travertine layer.
5. Leaf breakdown of the three indigenous species was faster than that of the exotic species P. orientalis . Due to the recalcitrance of its leaves, the frequent use of Platanus in revegetation schemes following the destruction of indigenous vegetation by fire, exacerbates the negative effect of travertine precipitation on leaf breakdown and, by extension, energy flow in Mediterranean karst streams.     

Fishpond dams affect leaf‐litter processing and associated detritivore communities along intermittent low‐order streams

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Leaf‐litter breakdown in pasture and deciduous woodland streams: a comparison among three European regions

1. Human land‐use has altered catchments on a large scale in most parts of the world, with one of the most profound changes relevant for streams and rivers being the widespread clearance of woody riparian vegetation to make way for livestock grazing pasture. Increasingly, environmental legislation, such as the EU Water Framework Directive (EU WFD), calls for bioassessment tools that can detect such anthropogenic impacts on ecosystem functioning. 2. We conducted a large‐scale field experiment in 30 European streams to quantify leaf‐litter breakdown, a key ecosystem process, in streams whose riparian zones and catchments had been cleared for pasture compared with those in native deciduous woodland. The study encompassed a west–east gradient, from Ireland to Switzerland to Romania, with each of the three countries representing a distinct region. We used coarse‐mesh and fine‐mesh litter bags (10 and 0.5 mm, respectively) to assess total, microbial and, by difference, macroinvertebrate‐mediated breakdown. 3. Overall, total breakdown rates did not differ between land‐use categories, but in some regions macroinvertebrate‐mediated breakdown was higher in deciduous woodland streams, whereas microbial breakdown was higher in pasture streams. This result suggests that overall ecosystem functioning is maintained by compensatory increases in microbial activity in pasture streams. 4. We suggest that simple coefficients of breakdown rates on their own often might not be powerful enough as a bioassessment tool for detecting differences related to land‐use such as riparian vegetation removal. However, shifts in the relative contributions to breakdown by microbial decomposers versus invertebrate detritivores, as revealed by the ratios of their associated breakdown rate coefficients, showed clear responses to land‐use.     

Habitat structure and fish predation: effects on invertebrate colonisation and breakdown of stream leaf packs

1. We investigated the effects of two features of leaf‐pack habitat structure (i.e. mass of a leaf pack and surface area of leaves comprising a leaf pack) and fish predation on colonisation of shredders and leaf breakdown rates in a coldwater stream. Packs were constructed of red maple (Acer rubrum) leaves. 2. A 2 × 3 × 3 factorial experiment was used to manipulate fish predation (exclusion and control cage), leaf‐pack mass (1, 3 and 5 g dry mass) and leaf surface area (small: approx. 17.9 cm², medium: approx. 34.6 cm², large: approx. 65.6 cm²). Exclusion cages had mesh on all sides, whereas control cages lacked mesh on two sides to provide access to fish. 3. Common shredders were Gammarus pseudolimnaeus, Pycnopsyche and Lepidostoma. Shredder biomass per leaf pack increased with the mass of a leaf pack (P < 0.001), but biomass per unit mass of leaf pack did not differ with leaf‐pack mass (P = 0.506). Shredder densities did not respond to the exclusion of fish (P > 0.7) or leaf surface area (P > 0.7), and interactions among treatment factors were not significant (P > 0.2). 4. Breakdown rates were lower for leaf packs comprised of small leaves (P < 0.001) and leaf packs with high mass (P = 0.001). Excluding fish did not significantly affect leaf breakdown rates (P = 0.293), and interactions among treatment factors were not significant (P > 0.3). Breakdown rates were highest when packs consisted of few leaves (i.e. leaf packs with large leaves and low mass) and were colonised by many shredders. 5. Fish predation was not an important factor controlling shredder densities in leaf packs over the spatiotemporal scale of our experiment. Nevertheless, we found shredder colonisation was proportional to leaf‐pack mass and breakdown rates were affected by leaf‐pack size (i.e. number of leaves in a pack). We suspect that fragmentation is the primary mechanism causing the breakdown rates to be dependent on leaf‐pack size.     

Effects of litter size and quality on processing by decomposers in a tropical savannah stream

Litter fragment size and quality can have profound effects on ecosystem functioning and global biogeochemical cycling due to differential utilization by decomposers. Here we study the influence of these factors on decomposers from two guilds found in a tropical savannah stream: invertebrate shredders of the genus Phylloicus and microorganisms. Containers (16 × 16 × 12 cm, ~ 3L) with either Phylloicus (cases removed; N = 16) or stream water containing microorganisms (N = 16) were supplied with litter from the species Inga laurina, Maprounea guianensis, and Richeria grandis, and cut into disks of 18.7, 13.2, and 8.1 mm in diameter (3 sizes × 3 species = 9 disks per container). Relative decomposition was greater for smaller leaf disks and disks of higher quality in microbial‐only cultures. Phylloicus preferentially harvested large fragments for case building, also preferring the leaves of M. guianensis and R. grandis, likely due increased robustness for case formation. Microbial decomposition resulted in ~20% litter mass loss compared to 30% in Phylloicus (of which 8% was used for case building and 24% for food). Thus, changes to input litter size, such as a decrease in leaf size after drought, may alter microbial decomposition and potentially affect shredder populations by limiting the availability of casing material.     

Freshwater crabs (Decapoda: Pseudothelphusidae) increase rates of leaf breakdown in a neotropical headwater stream

1. Freshwater crabs are the largest macroconsumers in many neotropical headwater streams, but few studies have examined their roles in ecosystem processes such as leaf litter breakdown. As omnivorous macroconsumers, freshwater crabs affect multiple trophic levels. They may directly increase leaf breakdown through fragmentation and consumption or indirectly decrease breakdown by consuming other macroinvertebrates, including shredders and detritivores.
2. In a headwater stream in Monteverde, Costa Rica, we conducted an in-stream experiment with 40 enclosures to quantify the effects of pseudothelphusid crabs on both leaf breakdown and macroinvertebrate colonisation of leaves. Half of the enclosures were randomly selected to contain two crabs (mean carapace width = 30 mm) and half were controls without crabs. We sampled mixed leaf packs from the enclosures on days 11, 19, 28, 34, and 42. We found the leaves of one species (Koanophyllon pittieri) almost completely decomposed by day 28 in both treatments (crab versus no crab). The other two leaf species (Meliosma idiopoda, Quercus brenesii) composed the remaining leaf mass at the end of the experiment.
3. At 42 days, enclosures with crabs had faster rates of leaf breakdown than those without crabs (with crabs: k = −0.020; without crabs: k = −0.016; p = 0.034). This suggests that the magnitude of direct leaf breakdown by crabs, due to fragmentation, consumption, or manipulation of leaves, was greater than any indirect effects on leaf breakdown via crab consumption of other leaf-consuming species.
4. Macroinvertebrate composition based on taxa abundances or biomasses did not significantly differ between treatments (ANOSIM; p = 0.73 and p = 0.65, respectively). Shredder and detritivore abundances and biomasses increased significantly through time (ANOVA; p ≤ 0.001), but there was no evidence of an effect of crab presence (p > 0.2), nor were there significant interactions between crab presence and time (p > 0.3).
5. This is one of the first studies to quantify the effects of pseudothelphusid freshwater crabs on leaf breakdown rates. Our results suggest that these crabs can play a significant role in detrital processing in neotropical headwater streams. This study has also demonstrated that short-term enclosure experiments are useful in measuring in-stream effects of crab activity on leaf breakdown.

     

Response of Aquatic Leaf Associated Microbial Communities to Elevated Leachate DOC: A Microcosm Study

In Central Europe climate change will increase summer droughts, which cause both, premature leaf fall and fragmentation of small streams during summer and early autumn. As a consequence dissolved organic carbon (DOC) leached from leaves will be dispersed into pools with long water residence time. A microcosm experiment was performed to test the effect of high concentrations of leachate DOC and the relative importance of labile and refractory leachate compounds on leaf associated microbial parameters. In microcosms leaf discs colonized in a stream were exposed to high concentrations of either leaf leachate, glucose or tannic acid. Leaf associated respiration, fungal sporulation, leaf mass loss and fungal biomass (ergosterol) were measured during a 3 weeks experimental period and compared to control without DOC amendment. The results imply that depending on source and composition elevated leachate DOC may have variable effects on microbial mediated litter decomposition. Our findings suggest reduced microbial decomposition rates in pools of fragmented streams receiving premature leaf fall. (© 2007 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Responses in organic matter accumulation and processing to an experimental wood addition in three headwater streams

1. Additions of large wood are being used to restore streams that have been subjected to channelization, wood removal or riparian timber harvest. This added wood potentially increases channel stability, habitat complexity and organic matter retention and improves habitat and productivity of higher trophic levels. However, few stream restorations monitor restoration effectiveness after project completion. 2. We added 25 aspen logs (each 2.5 m length × 0.5 m diameter) to 100‐m reaches of each of three forested headwater streams in the Upper Peninsula of Michigan, U.S.A. These wood‐poor streams drain forests that were completely harvested of timber over a century ago and have been selectively logged for the past 50–60 years. An upstream unmanipulateds 100‐m reach in each stream served as the control. 3. We evaluated responses in organic matter processing by measuring red maple leaf decomposition 1 year before and 2 years after wood addition. We also quantified coarse organic matter standing stocks in the main channel and in debris accumulations associated with large wood. In response to wood addition, we predicted both organic matter standing stocks and leaf decomposition rates would increase, thereby enhancing resource availability to higher trophic levels. 4. Leaf decomposition rates did not change following wood addition. Temporal variation in rates among streams was mostly explained by differences in degree days, water velocity, scour/burial and water column inorganic nitrogen concentrations, but not large wood. Variation within streams across years was explained by differences in degree days, water velocity and shredder biomass. 5. Contrary to our prediction, organic matter standing stocks did not increase significantly at the reach scale. However, the experimentally added wood retained c. 4% of total annual coarse benthic organic matter (CBOM) in the first year and an additional c. 15% in the second year, suggesting accumulation over time in the manipulated reaches. The CBOM held by the new logs may be more biologically available because it is less susceptible to burial and transport than material in the streambed. 6. Some shredding macroinvertebrates responded to changes caused by the wood additions. In particular, the common caddisfly shredder, Lepidostoma sp., increased in abundance in leaf bags following wood addition, whereas the biomass of the winter stoneflies, Capniidae, declined in the first year. 7. Considerable funds are spent to restore in‐stream habitat, but few restorations are monitored, particularly over long periods (>5 years). Our results show that longer‐term monitoring is needed to determine the efficacy of these restorations on ecosystem function; organic matter decomposition in our low‐gradient streams did not respond to a substantial increase in large wood after 2 years.     

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)     

Catchment land-use, macroinvertebrates and detritus processing in headwater streams: taxonomic richness versus function

1. We used a litter bag technique to assess the effect of catchment land-use (forest, wetland, agriculture, urban) on the processing of red maple ( Acer rubrum L.) litter in 17 streams in Maine, U.S.A. Litter processing by fungi was predicted to increase with nutrient concentrations along a gradient of land use, from relatively unmodified to highly modified. Litter processing by litter-shredding macroinvertebrates was predicted to decline along this gradient because of a decline in their taxonomic richness and biomass.
2. Land use was associated with the anticipated gradient in nutrient and macroinvertebrate attributes, and a significant relationship was found between land use and nitrate concentration. There was, however, no significant relationship between land use and soluble reactive phosphorus (SRP) concentration. Similarly, shredder taxonomic richness was significantly related to land use type, whereas shredder biomass showed no significant relationship to land use.
3. Attributes of the shredder assemblage structure and nutrient concentrations were both strong determinants of litter processing. Increasing biomass and taxonomic richness of shredders was significantly related to increasing rates of litter mass loss. Increasing concentrations of nitrate and SRP were significantly related to increasing rates of litter softening below threshold concentrations (approximately 0.20 mg NO₃-N L^–1 and 5 μg SRP L^–1).
4. The potentially additive effects of nitrate and SRP concentrations or shredder richness and biomass on litter processing rates were confounded by the lack of significant correlation between these pairs of variables. Consequently, rates of litter processing (as rates of softening or mass loss) did not vary systematically among different land use regimes.     

Effects of land use on the structure and function of leaf‐litter microbial communities in boreal streams

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Effects of organic matter and season on leaf litter colonisation and breakdown in cave streams

1. Low organic matter availability is thought to be a primary factor influencing evolutionary and ecological processes in cave ecosystems. We examined links among organic matter abundance, macroinvertebrate community structure and breakdown rates of red maple (Acer rubrum) and corn litter (Zea mays) in coarse‐ (10 × 8 mm) and fine‐mesh (500‐μm) litter bags over two seasonal periods in four cave streams in the south‐eastern U.S.A. 2. Organic matter abundance differed among cave streams, averaging from near zero to 850 g ash‐free dry mass m^?2. Each cave system harboured a different macroinvertebrate community. However, trophic structure was similar among caves, with low shredder biomass (2–17% of total biomass). 3. Corn litter breakdown rates (mean k = 0.005 day^?1) were faster than red maple (mean k = 0.003 day^?1). Breakdown rates in coarse‐mesh bags (k = 0.001–0.012 day^?1) were up to three times faster than in fine‐mesh bags (k = 0.001–0.004 day^?1). Neither invertebrate biomass in litter bags nor breakdown rates were correlated with the ambient abundance of organic matter. Litter breakdown rates showed no significant temporal variation. Epigean (surface‐adapted) invertebrates dominated biomass in litter bags, suggesting that their effects on cave ecosystem processes may be greater than hypogean (cave‐adapted) taxa, the traditional focus of cave studies. 4. The functional diversity of our cave communities and litter breakdown rates are comparable to those found in previous litter breakdown studies in cave streams, suggesting that the factors that control organic matter processing (e.g. trophic structure of communities) may be broadly similar across geographically diverse areas.     

Tropical shift in decomposers’ relative contribution to leaf litter breakdown in two Guinean streams

The hypothesis that leaf litter breakdown in Guinean streams is governed by microorganisms was confirmed, supporting the reported latitudinal shift in decomposers’ contribution to this process. The large body size of dominant macroinvertebrate decomposers (shrimps) only partially compensated for their very low densities. In contrast with other tropical regions mostly dominated by insect larvae, the functional consequences of global warming on these stream ecosystems may be less severe due to the lower sensitivity of crustaceans to temperature increase.     

Top‐down and bottom‐up control of litter decomposers in streams

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Changes in mechanisms of summer detritus processing between wooded and agricultural sites in a Michigan headwater stream

The mechanisms of leaf decay, leaf-associated macroinvertebrate community structure, leaf-associated microbial activity and physicochemical stream characteristics were investigated on a mid-Michigan headwater stream in summer. An undisturbed wooded site was compared with two agriculturally perturbed sites. Discharge, total suspended particulates, and nutrients were all higher and more variable throughout the season within the agricultural reaches. Leaf decay rates were higher at the agricultural sites and results suggest discharge abrasion was the major leaf processing mechanism at these sites while microbial decay and macroinvertebrate shredding appear to be the primary mediators of leaf weight loss at the wooded site. Total macroinvertebrate densities on leaf packs at the agricultural sites were 1.9 times the densities at the wooded site. It is suggested that experimentally introduced leaf packs acted as a lure for net-spinning invertebrates limited by stable substratum at the agricultural sites. Species shifts were observed from wooded reaches where Pychnopsyche spp., Gammarus, Ephemeroptera, Bezzia, and Nigronia serricornis were important, to downstream agricultural reaches which were dominated by Cheumatopsyche, Chironomidae, Elmidae, Hydracarina, Hemerodromia, and Caecidotea.