Feeding tests with caddis larvae (Insecta: Trichoptera) and amphipods (Crustacea: Amphipoda) on Platanus orientalis (Platanaceae) and other leaf litter

In the southern parts of the Mediterranean region, as in the island of Crete, there are few species of riparian trees and shrubs among the dominant Platanus orientalis. Feeding tests have shown that leaves of Platanus are not eaten by aquatic shredders of continental and Cretean origin. The large quantities of organic matter are therefore not used as a source of food and energy by the stream communities. In addition, the high winter flow shortly after leaf fall, and the short courses of rivers result in loss of most of the leaf litter to the sea. The River Continuum Concept does therefore not apply to this region. Field observations have shown that amphipods and limnephilid larvae are shifting from shredding to scraping habits if no leaf litter except Platanus was available.     

Litter processing in diffuse and conduit springs

We used bedrock geology and prior water chemistry data to classify and choose 4 conduit and 4 diffuse flow springs in a karst region of southeast Minnesota. Decomposition processes and chemical/physical conditions were compared between spring classes during two seasons (May and September, 1987).Although large storms and conduit run-in events did not occur during our sampling, baseline water chemistry supported our a priori spring classification. Baseline nitrate and atrazine levels were significantly higher in conduit than in diffuse springs. During a minor run-in event, atrazine levels increased significantly to at least 1.2 µg L^–1 in all conduit systems, but remained unchanged in diffuse systems.Decay rates of the two predominant litter sources in the springs (watercress [Nasturtium officinale R.Br.] in May and box elder [Acer negundo L.] in September) were used to assess differences in biological activity between the spring classes. During May, watercress decayed (1 mm coarse mesh bags) at approximately the same rate in conduit and diffuse systems, k = 0.094 and 0.099 respectively. This result was unexpected since shredder colonization was much greater on litter bags in diffuse systems. In September, boxelder decay rate was significantly higher in diffuse (k = 0.018) than in conduit (k = 0.013) springs; and decay rates seemed to reflect significantly higher shredder colonization on bags in diffuse systems. Microbial activity on watercress and boxelder litter, measured as electron transport system (ETS) activity, was seldom significantly different between spring classes. Also, both watercress and boxelder litter decay rates for fine mesh bags (240µm) were similar between conduit and diffuse springs, suggesting that microbial processing did not differ greatly between spring classes.In conduit springs, low shredder colonization was apparently the result of low benthic shredder densities. Shredders and other macroinvertebrates may be adversely affected by discharge fluctuations from conduit run-in events. We suspect that, during times of low flow, watercress standing stock was also an important influence on shredder abundance.     

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

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

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

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

The effects of shredding invertebrates on the transfer of organic carbon from littoral leaf litter to water-column bacteria

Leaf litter can be of great importance for the productivity of small oligotrophic lakes surrounded by deciduous forests. Feeding invertebrate shredders produce particulate organic leftovers, but their feeding also enhances the release of dissolved organic carbon (DOC). We tested whether invertebrate-mediated DOC release affects the production of heterotrophic water-column bacteria. Submersed leaves were incubated in microcosms with and without shredders; and DOC, absorbance, bacterial abundance and bacterial production in the water column were monitored. We also measured dry weight of the organic particles (FPOC, fine particulate organic carbon, leaf residues and shredders). Total leaf-litter carbon decreased by nearly 80% in the presence of shredders, and on average 56% of the initial leaf carbon ended up as FPOC after 126 days of incubation. Without shredders FPOC production was almost zero, and 72% of the added leaf carbon could be retrieved as leaves when the experiment ended. Both these figures include the rapid release of DOC during the first week of leaf incubation in the lake water (equivalent to 16–19% of total added leaf carbon). Although bacterial production in the water was several times higher in treatments with shredders, bacterial consumption of leaf-derived DOC from shredding was obviously of minor importance in the total carbon budget. This result suggests, although shredders have a strong impact on transformation of leaves to FPOC, they do not greatly enhance the initial rate of mineralization of the leaf-derived detritus.     

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

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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.     

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)     

Resource quality and stoichiometric constraints on stream ecosystem functioning

1. Resource quality and stoichiometric imbalances in carbon : nutrient ratios between consumers and resources can influence key ecosystem processes. In many streams, this has important implications for food webs that are based largely upon the utilization of terrestrial leaf‐litter, which varies widely among litter types in its value as a food source for detritivores and as a substrate for microbial decomposers. 2. We measured breakdown rates and macroinvertebrate colonization of leaf‐litter from a range of native and exotic plants of differing resource quality and palatability to consumers [e.g. carbon : nitrogen : phosphorus (C : N : P) ratios, lignin and cellulose content], in a field experiment. We also measured C : N : P ratios of the principal leaf‐shredding invertebrates, which revealed strong stoichiometric imbalances across trophic levels: C : N and C : P ratios typically differed by at least one order of magnitude between consumers and resources, whereas N : P imbalances were less marked. Application of the threshold elemental ratio approach, which integrates animal bioenergetics and body elemental composition in examining nutrient deficiency between consumers and resources, revealed less marked C : P imbalances than those based on the simpler arithmetic differences described above. 3. Litter breakdown rates declined as nutrient imbalances widened and resource quality fell, but they were independent of whether resources were exotic or native. The principal drivers of total, microbial and invertebrate‐mediated breakdown rates were lignin : N, lignin : P and fungal biomass, respectively. However, multiple regression using orthogonal predictors yielded even more efficient models of litter breakdown, as consumers responded to more than one aspect of resource quality. For example, fungal biomass and litter C : N both influenced invertebrate‐mediated breakdown. 4. Large stoichiometric imbalances and changes in resource quality are likely to have serious consequences for stream ecosystem functioning, especially when riparian zones have been invaded by exotic plant species whose chemical composition differs markedly from that of the native flora. Consequently, the magnitude and direction of change in breakdown rates and, thus, resource depletion, will be driven to a large extent by the biochemical traits (rather than taxonomic identity per se) of the resident and invading flora.