Forest canopy cover determines invertebrate diversity and ecosystem process rates in depositional zones of headwater streams

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Managing and Rehabilitating Ecosystem Processes in Regional Urban Streams in Australia

Urbanization is acknowledged as one of the most severe threats to stream health, spawning recent research efforts into methods to ameliorate these negative impacts. Attention has focused on streams in densely-populated cities but less populous regional urban centres can be equally prone to some of the same threats yet might not meet the conventional definitions of urban. Several recent reviews have identified the changes to streams that occur during urbanization but they note that few ecological studies have explored ecosystem-level responses, typically focusing instead on state variables such as invertebrate abundance. In many regional urban streams, changes to the extent of impervious drainage have implications for their hydrology and channel morphology but the influence of these changes on fundamental ecosystem processes of leaf litter breakdown and transport compared with those in nearby rural streams are poorly known. The widespread practice of planting exotic trees along riparian zones and street margins draining into urban streams further exacerbates the disruption of natural organic matter dynamics. The combination of seasonal leaf fall by exotic species and the altered drainage patterns through urbanization in Armidale, a regional town in New South Wales, Australia, resulted in contrasting patterns of benthic organic matter storage over 18 months compared to nearby reference and rural streams. Macroinvertebrate detritivore densities were low in the urban stream, implying disruption of the usual biological pathways of leaf breakdown. Understanding the interactions of hydrology, drainage pattern, leaf input and biological attributes of a stream is crucial for managers trying to restore stream ecosystem services without incurring public concern about the appearance of regional urban streams.     

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

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

     

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

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

Ecosystem Linkages between Southern Appalachian Headwater Streams and Their Banks: Leaf Litter Breakdown and Invertebrate Assemblages

We examined red maple (Acer rubrum L.) leaf litter breakdown in streams and riparian zones at two sites in the southern Appalachian Mountains to understand how differences in abiotic and biotic factors influence leaf breakdown rates. Litterbags were placed in three riparian habitats differing in litter layer moisture: stream > bank > upland. Invertebrates colonizing litterbags at one site were also examined to determine how variations in community and functional structure affect breakdown rates. Leaves broke down fastest in streams and slowest in upland habitats, whereas bank habitats were intermediate and characterized by high variability. Faster leaf breakdown rates in streams appeared to be a function of greater moisture availability, a more stable thermal regime, and a higher biomass of leaf-shredding invertebrates, especially the stonefly Tallaperla. In addition, patterns of leaf breakdown and invertebrate community structure provided evidence for a stronger than expected ecological connection between the stream and the bank. Overall, detritus processing within this narrow riparian ecosystem varied considerably depending on the availability of moisture. Results from this study show that stream channel–floodplain interactions in riparian ecosystems of steep forested mountains are analogous to ones in larger downstream or low-gradient systems. Riparian zones throughout a river network display a remarkable heterogeneity in their ability to process organic matter, which is ultimately driven by changes in hydrological conditions. Received 6 March 2001; accepted 3 July 2001.     

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.     

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.     

Effects of agricultural development on processing of tussock leaf litter in high country New Zealand streams

1. We compare the rates and mechanisms of processing of tussock (Chionochloa spp.) leaf litter in six New Zealand streams draining grassland catchments that contrast in the extent to which they have been developed for pasture. 2. Rates of processing, measured as rate of weight loss of leaf packs and rate of leaf softening, were at the slow end of the spectrum for vascular plant processing. Processing was faster at developed sites, mediated mainly through the influence of oxidized nitrogen concentration on microbial activity. 3. Few invertebrate shredders colonized leaf packs and it is unlikely that invertebrates had an appreciable effect on leaf processing in our study streams, which do not effectively retain leaf litter. Very small headwater tributaries appear to retain leaf litter and possess a more abundant shredder community. 4. Measures of leaf processing in our six streams were significantly correlated with Petersen's (1992) RCE score of stream condition. We discuss the potential for using rate of leaf litter processing as a method of bioassessment. 5. Even the most degraded stream in our study is classed as ‘good’ using the RCE inventory system. Human impact in the Taieri River is relatively small compared with the degradation observed in some parts of the world.     

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.     

An invasive species may be better than none: invasive signal and native noble crayfish have similar community effects

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Do leaf-litter decomposers control biofilm primary production and benthic algal community structure in forest streams? Insights from an outdoor mesocosm experiment

• 1. Forested headwater streams are generally considered to be light-limited ecosystems where primary production is reduced, and the main source of energy and nutrients is composed of allochthonous detritus. We hypothesised that in these ecosystems, the development of primary producers might also be limited by (1) competition for nutrients with leaf-litter decomposers (e.g. bacteria and fungi), and (2) leaf-litter leachates or allelopathic compounds produced by aquatic fungi.
• 2. To test these hypotheses, a 48-day mesocosm experiment was performed in 12 artificial streams containing stream water inoculated with epilithic biofilm suspensions collected from a forested headwater stream. Three different treatments were applied: control without leaf litter (C), microbially conditioned leaf litter added at the beginning of the experiment and left to decompose throughout the experiment (L), or leaf litter renewed three times during the experiment (RL).
• 3. We predicted that (1) the presence of litter, through microbial nutrient immobilisation and allelopathy, would reduce primary production and that (2) this effect would be amplified by litter renewal. We also predicted that nutrient competition would mean that (3) leaf-litter decomposers will alter primary producer community composition and physiology. These predictions were tested by analysing biofilm development, physiology, stoichiometry, and benthic algal community structure. To distinguish between the effects of nutrient immobilisation and allelopathy, the biofilm responses to leaf-litter leachates collected after different microbial conditioning durations were also measured in a parallel laboratory experiment.
• 4. Contrary to our expectations, by day 28, primary producer growth was higher in the mesocosms containing leaf litter (L and RL) despite the rapid decrease in dissolved nutrients when leaf litter was present. After 48 days, the lowest phototrophic biofilm development was observed when leaf litter was renewed (RL), whereas phototrophic biofilm development was similar in the C and L treatments. Biofilm stoichiometry indicated that this effect was most probably related to greater nitrogen limitation in the RL treatment. The presence of leaf litter also affected primary producers' photophysiology, which could be attributed to changes in taxonomic composition and to physiological adjustments of primary producers.
• 5. Laboratory measurements showed that despite a strong inhibition of primary producer growth by unconditioned leaf-litter leachates, microbially conditioned leaf litter had either low or no effects on the development of primary producers.
• 6. These results reveal that leaf-litter decomposers can have both positive and negative effects on primary producers underlining the need to consider microbial interactions when investigating the functioning of forested headwater streams.

     

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.     

Resource–consumer diversity: testing the effects of leaf litter species diversity on stream macroinvertebrate communities

1. Understanding relationships between resource and consumer diversity is essential to predicting how changes in resource diversity might affect several trophic levels and overall ecosystem functioning. 2. We tested for the effects of leaf litter species diversity (i.e. litter mixing) on litter mass remaining and macroinvertebrate communities (taxon diversity, abundance and biomass) during breakdown in a detritus‐based headwater stream (North Carolina, U.S.A.). We used full‐factorial analyses of single‐ and mixed‐species litter from dominant riparian tree species with distinct leaf chemistries [red maple (Acer rubrum), tulip poplar (Liriodendron tulipifera), chestnut oak (Quercus prinus) and rhododendron (Rhododendron maximum)] to test for additivity (single‐species litter presence/absence effects) and non‐additivity (emergent effects of litter species interactions). 3. Significant non‐additive effects of litter mixing on litter mass remaining were explained by species composition, but not richness, and litter‐mixing effects were variable throughout breakdown. Specifically, small differences in observed versus expected litter mass remaining were measured on day 14; whereas observed litter mass remaining in mixed‐species leaf packs was significantly higher on day 70 and lower on day 118 than expected from data for single‐species leaf packs. 4. Litter mixing had non‐additive effects on macroinvertebrate community structure. The number of species in litter mixtures (two to four), but not litter species composition, was a significant predictor of the dominance of particular macroinvertebrates (i.e. indicator taxa) within mixed‐species packs. 5. In addition, the presence/absence of high‐ (L. tulipifera) and low‐quality (R. maximum) litter had additive effects on macroinvertebrate taxon richness, abundance and biomass. The presence of L. tulipifera litter had both positive (synergistic) and negative (antagonistic) effects on invertebrate taxon richness, that varied during breakdown but were not related to litter chemistry. In contrast, the presence/absence of L. tulipifera had a negative relationship with total macroinvertebrate biomass (due to low leaf mass remaining when L. tulipifera was present and higher condensed and hydrolysable tannins associated with leaf packs lacking L. tulipifera). Macroinvertebrate abundance was consistently lower when R. maximum was present, which was partially explained by litter chemistry [e.g., high concentrations of lignin, condensed tannins, hydrolysable tannins and total phenolics and high carbon to nutrient (N and P) ratios]. 6. The bottom‐up effects of litter species diversity on stream macroinvertebrates and litter breakdown are different, which suggests that structural attributes of macroinvertebrate communities may only partially explain the effects of litter‐mixing on organic matter processing in streams. In addition, stream macroinvertebrates colonising decomposing litter are influenced by resource diversity as well as resource availability. Broad‐scale shifts in riparian tree species composition will alter litter inputs to streams, and our results suggest that changes in the diversity and availability of terrestrial litter may alter stream food webs and organic matter processing.     

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

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

Leaf‐litter colonisation and breakdown in relation to stream typology: insights from Mediterranean low‐order streams

1. Scant information is available on leaf breakdown in streams of arid and semiarid regions, including the Mediterranean, where environmental heterogeneity can be high and the relationship between stream characteristics and leaf breakdown is poorly known. We tested the hypotheses that differences in leaf breakdown metrics would be substantially higher between mountain and lowland Mediterranean streams than among streams within each subregion and that variability among streams would be substantially higher in the lowlands, because permanent reaches in the semiarid lowland streams are rare and isolated. 2. We compared leaf breakdown and associated dynamics of nutrients, fungi and invertebrates in low‐order Mediterranean streams draining sub‐humid forests in the Sierra Nevada Mountains and nearby semiarid lowlands of south‐eastern Spain. Streams differed between the two subregions mainly in water ion content, temperature and riparian tree cover. We detected higher environmental heterogeneity among streams within the lowlands compared to the Sierra Nevada mountain range. In the lowlands, breakdown coefficients (k) of alder leaves spanned almost the entire range reported for this species from temperate streams, overlapping with less variable breakdown coefficients in the Sierra Nevada. 3. The high variability of k values among the lowland sites appeared to be caused primarily by variability in the composition and abundance of a few leaf‐consuming invertebrate taxa, particularly the snail Melanopsis praemorsa. Fungal and nutrient dynamics were less variable among sites within each subregion. 4. These results indicate that the critical condition for stream functional assessment of well‐constrained breakdown rates, or related metrics, could be met at reference sites within homogenous bio‐geo‐climatic regions such as the Sierra Nevada. By contrast, in heterogeneous areas such as the semiarid lowland streams, natural variability of breakdown rates can greatly exceed the magnitude of effects expected in response to anthropogenic disturbances.     

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).     

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.     

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

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

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.     

The influence of water chemistry on the enzymatic degradation of leaves in streams

1. Aquatic hyphomycetes degrade leaf litter in both softwater and hardwater streams. During growth on leaves, these fungi secrete an array of extracellular polysaccharidases that are differentially affected by pH. Hydrolytic enzymes exhibit acidic pH optima, whereas pectin lyases have neutral to alkaline pH optima. 2. Enzyme activities associated with microbial communities colonizing yellow poplar (Liriodendron tulipifera) leaves submerged in an acidic (pH 6.3), softwater stream were compared with those occurring in an alkaline (pH 8.2), hardwater stream. In addition to pH differences, the hardwater stream had higher nutrient concentrations and higher temperatures than the softwater stream. Conditions in the hardwater stream favoured greater microbial growth, fungal activity, rates of leaf breakdown and softening. However, activities of hydrolytic enzymes (xylanase, endocellulase, galacturonanase) were lower in the hardwater stream than in the softwater stream. Consequently, activities of these hydrolytic enzymes were not good indicators of leaf breakdown in these streams. 3. In contrast, the activities of pectin lyase were higher in the hardwater stream than in the softwater stream, corresponding to the greater rates of leaf breakdown and softening that occurred in the hardwater stream. These results support previous findings that pectin lyase is closely associated with the softening and maceration of leaf detritus and suggest that pectin degradation is a key process in the initial stages of leaf breakdown.