The importance of crayfish in the breakdown of rhododendron leaf litter

1. Rhododendron (Rhododendron maximum) is a common evergreen shrub in riparian areas of the southern Appalachians, where its leaves can comprise a large proportion of leaf litter in streams. However, they are relatively refractory and generally considered a low quality food resource for detritivores. 2. Our objective was to assess whether macroconsumers [primarily crayfish (Cambarus bartonii)] influence rhododendron leaf breakdown in a forested southern Appalachian stream in both summer (when leaves other than rhododendron are relatively scarce) and autumn (when other leaves are relatively abundant). We conducted two leaf decay experiments, one in summer and one in autumn, using pre‐conditioned leaves. Macroconsumers were excluded from the benthos of a fourth‐order stream using electric ‘fences’; we predicted that excluding macroconsumers would reduce the decay rate of rhododendron leaves in both summer and autumn. 3. In both experiments, breakdown rate was lower in exclusion treatments. Macroconsumers accounted for approximately 33 and 54% of rhododendron decay in summer and autumn, respectively. We attribute this effect to direct shredding of rhododendron by crayfish. Biomass of insect shredders, insect predators and fungi did not differ between control and exclusion treatments, indicating that insectivorous sculpins (Cottus bairdi) had no effect on rhododendron decay and that omnivorous crayfish did not exert an indirect effect via alteration of insect or fungal biomass. 4. The influence of shredding insects varied between summer and autumn. In summer, when other, more palatable leaf types were not available, rhododendron leaf packs appeared to provide ‘resource islands’ for insect shredders. There was a significant inverse relationship between insect shredders and leaf pack mass in the summer exclusion treatment: insects were the only organisms eating leaves in this treatment and, as shredder biomass increased, remaining leaf pack mass decreased. In the control treatment, however, we did not see this relationship; here, the effect of insect shredders was presumably swamped by the impact of crayfish. In autumn, when other leaves were abundant, insect shredder biomass in rhododendron leaf packs was less than one‐third of summer values. 5. Even at low density (approximately 2 m^–2) crayfish were able to influence an ecosystem process such as leaf decay in both summer and autumn. Given the threatened status of many crayfish species in the United States, this finding is especially relevant. Even small alterations in crayfish assemblages, whether via loss of native species and/or introduction of exotic species, may have significant repercussions for ecosystem function.     

Litter retention in Tasmanian headwater streams after clear-fell logging

Clear-fell logging around small headwater streams in Tasmanian wet eucalypt forests was predicted to affect both the retention of leaf litter and the composition and size of leaf packs. Retention structures were surveyed in six natural streams and six streams in forest regenerated 3–5 years after clear-fell and burn logging. Logged streams had more wood, but retained less leaves than natural streams, and consequently had fewer and smaller leaf packs. Leaf packs from natural streams contained 200% more leaves, bark and twigs than packs from logged streams. The effect of buoyancy on leaf retention was assessed with release and recapture of marked Eucalyptus obliqua and Nothofagus cunninghamii leaves. Eucalypt leaves were more likely to be trapped by retention structures on the bed of the stream, while smaller, more buoyant N. cunninghamii leaves were mainly trapped by leaf packs. Leaf packs in natural streams were formed on a matrix of small twigs and long strips of bark, shed from the upper branches of mature stringybark eucalypts, while leaf retention was reduced in logged streams because there are no mature trees to provide effective retention structures. Changes to the channel form increase both discharge and sedimentation. These factors have strong implications for downstream nutrient processing and riverine food webs.     

Urban infrastructure influences dissolved organic matter quality and bacterial metabolism in an urban stream network

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Elevated temperature may intensify the positive effects of nutrients on microbial decomposition in streams

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Factors affecting litter processing by Anisocentropus kirramus (Trichoptera: Calamoceratidae) from an Australian tropical rainforest stream

• 1 Larvae of the caddisfly Anisocentropus kirramus are common leaf shredders in rainforest streams in tropical Queensland. Laboratory experiments were undertaken to examine the effects of (a) differences in leaf species, age and degree of conditioning, (b) leaf preferences, (c) temperature and (d) larval stage on processing of leaf litter by A. kirramus.
• 2 Leaf species (five tested), age (fresh v senescent) and condition (conditioned in stream v unconditioned) each had a significant effect on the amount of leaf material that was processed. For most species, conditioned leaves were processed faster but for one species unconditioned leaves were processed faster. Senescent leaves were processed more rapidly than green leaves in three species but not in the other two. Given a choice of leaf types A. kirramus actively selected leaves that were processed faster; no preference was shown between two different ‘fast’ leaves.
• 3 Processing occurred at all temperatures tested (10, 18 and 25°C), with the processing rate increasing with temperature. Younger instar larvae processed leaves at a greater rate per unit body weight (up to 343% day^?1) than older instars. Final instar (5) larvae were capable of processing some tough leaves that younger instars did not process.
• 4 In rainforest streams, processing of leaves by A. kirramus takes place throughout the year. Its ability to process green leaves is important because of the high input of fresh green leaves into tropical streams, and because of the severe depletion of the supply of conditioned leaves and fine detritus after floods.

     

Spatial heterogeneity in water velocity drives leaf litter dynamics in streams

1. Stream hydro-morphology refers to the heterogeneous distribution of hydrologic conditions that occur above a complex benthic surface such as a streambed.
2. We hypothesised that hydro-morphological conditions will influence the retention, re-distribution, and microbial-driven decomposition of leaf litter inputs in stream ecosystems because each process varies with overlying water velocity.
3. We tested this hypothesis using: (1) the spatial distribution of water velocity within a stream reach as a surrogate of stream hydro-morphology; (2) leaf tracer (i.e. Ginkgo biloba L.) additions with serial recovery to examine the relationship between benthic retention and overlying velocity; and (3) measurements of leaf litter decomposition (i.e. Alnus glutinosa [L.] Gaertn.) under different water velocity conditions.
4. Results demonstrate that water velocity exerts a significant influence on the retention and re-distribution of leaf litter inputs within the reach. The observed range of water velocity (from c. 0 to 0.92 cm/s) also strongly influences the range of leaf litter decomposition rates (0.0076–0.0222/day).
5. Our findings illustrate that water velocity influences leaf litter dynamics in streams by controlling leaf litter transport, retention and re-distribution as well as how leaves decompose within recipient stream reaches. Ultimately, the results show that the efficiency of leaf litter inputs in supporting stream ecosystem function is dependent on the hydro-morphological characteristics of the receiving stream ecosystems.

     

Seasonal dynamics of macroinvertebrate assemblages in the benthos and associated with detritus packs in two low-order streams with different riparian vegetation

• 1 The seasonal dynamics of the benthic macroinvertebrate assemblage, and the subset of this assemblage colonising naturally formed detritus accumulations, was investigated in two streams in south‐west Ireland, one draining a conifer plantation (Streamhill West) and the other with deciduous riparian vegetation (Glenfinish). The streams differed in the quantity, quality and diversity of allochthonous detritus and in hydrochemistry, the conifer stream being more acid at high discharge. We expected the macroinvertebrate assemblage colonising detritus to differ in the two streams, due to differences in the diversity and quantity of detrital inputs.
• 2 Benthic density and taxon richness did not differ between the two streams, but the density of shredders was greater in the conifer stream, where there was a greater mass of benthic detritus. There was a significant positive correlation between shredder density and detritus biomass in both streams over the study period.
• 3 Detritus packs in the deciduous stream were colonised by a greater number of macroinvertebrates and taxa than in the conifer stream, but packs in both streams had a similar abundance of shredders. The relative abundance of taxa colonising detritus packs was almost always significantly different to that found in the source pool of the benthos.
• 4 Correspondence analysis illustrated that there were distinct faunal differences between the two streams overall and seasonally within each stream. Differences between the streams were related to species tolerances to acid episodes in the conifer stream. Canonical correspondence analysis demonstrated a distinct seasonal pattern in the detrital composition of the packs and a corresponding seasonal pattern in the structure of the detritus pack macroinvertebrate assemblage.
• 5 Within‐stream seasonal variation both in benthic and detritus pack assemblages and in detrital inputs was of similar magnitude to the between‐stream variation. The conifer stream received less and poorer quality detritus than the deciduous stream, yet it retained more detritus and had more shredders in the benthos. This apparent contradiction may be explained by the influence of hydrochemistry (during spate events) on the shredder assemblage, by differences in riparian vegetation between the two streams, and possibly by the ability of some taxa to exhibit more generalist feeding habits and thus supplement their diets in the absence of high quality detritus.

     

The Influence of Pack Size and Position,Leaf Type,and Shredder Access on the Processing Rate of Atherosperma moschatum Leaves in an Australian Cool Temperate Rainforest Stream

Factors affecting the processing rate of packs of southern sassafras (Atherosperma moschatum) leaves were investigated in an Australian cool temperate rainforest stream pool. Processing rate was strongly influenced by pack weight fitting a linear inverse relationship. Processing was not significantly (p>0.05) affected by whether the packs were placed on bricks or free on the stream bed. Free packs, but not packs on bricks, were processed more rapidly near the bank than in midstream (p = 0.050). Packs placed in 300 μm mesh bags, with one side unsealed were processed at the same rate as packs attached to bricks or free on the stream bed, but packs in sealed 300 μm mesh bags were processed significantly more slowly (p<10⁻⁴). Abscissed leaves were processed significantly more slowly than fresh leaves (p<10⁻⁵) and at a rate not significantly different to leaves sealed in mesh bags (p = 0.197). There was no consistent pattern of difference between processing rates in summer and winter. The results indicate that the size of litter packs may be a more important factor in influencing processing rate estimates than the method of attachment of the packs. The slow rate of processing of abscissed leaves compared with fresh leaves indicates that they were at most, slightly influenced by stream shredders. The absence of a consistent pattern of difference between summer and winter processing rates is consistent with the results of previous litter processing studies from southeastern Australia although no clear causal factor can be identified.     

Nitrogen processing by grazers in a headwater stream: riparian connections

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Colonization and processing of leaf litter by macroinvertebrate shredders in streams of contrasting pH

• 1 Leaf litter processing rates and macroinvertebrate shredder assemblages in leaf packs were compared in four streams on the Allegheny plateau in the central Appalachian Mountains, U.S.A.; these streams were characterized by different bedrock geology and streamwater pH.
• 2 Leaf litter processing rates were fastest in the neutral streams, slowest in the acidic stream, and intermediate in the most alkaline stream.
• 3 Slower processing rates in the acidic stream were associated with lower total shredder biomass, made up predominantly by small leuctrid and nemourid stoneflies.
• 4 The differences in processing rates between the more alkaline stream and the neutral streams were not associated with differences in shredder biomass, but appeared to be related to taxonomic differences in the shredder assembiages. Insects were dominant in the neutral streams, and amphipods were dominant in the more alkaline stream.

     

Effects of a forest disturbance on shredder production in southern Appalachian headwater streams

• 1 The effects of a forest disturbance were investigated by comparing production of leaf-shredding aquatic insects in three streams draining a mature hardwood forest and three streams draining an 11-year-old, cable-logged clearcut.
• 2 Reference streams contained significantly greater mean annual standing crop of leaf material and significantly more slow-processing leaf material than disturbed streams. Disturbed streams had a significantly higher mean annual standing crop of fast-processing leaf material than the reference streams.
• 3 Leaf-shredding cranefly (Tipula abdominalis), caddisfly (Pycnopsyche gentilis), and stonefly (Tallaperla maria) larvae comprised over 95% of shredder biomass in all streams. Total shredder production was significantly greater (P<0.05) in disturbed versus reference streams, but individual production rates were not significantly different between stream types.
• 4 Pycnopsyche gentilis larvae were present at higher densities and achieved significantly greater annual biomass in disturbed versus references streams, Biomass of P. gentilis was significantly correlated with the standing crop of fast-processing, early successional leaf material in samples, whereas biomass of other shredders was correlated significantly with medium or slow-processing leaf species characteristic of later stages of forest succession.

     

Preconditioning of leaves by solar radiation and anoxia affects microbial colonisation and rate of leaf mass loss in an intermittent stream

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Seasonal differences in green leaf breakdown and nutrient content of deciduous and evergreen tree species and grass in a granitic headwater stream

Litter processing was examined in autumn–winter and spring–summer in a second order stream in Galicia (NW Spain). We compared decay rate and nutrient dynamics of green leaves of several deciduous (riparian: Alnus glutinosa, Betula alba and Populus×canadensis; terrestrial: Castanea sativa, Quercus robur), and evergreen tree species (terrestrial: Pinus radiata and Eucalyptus nitens), in addition to ray-grass (Lolium perenne). In the autumn–winter period, the decay rates (–k) ranged between 0.0086 degree-days^–1 for poplar, and 0.0019 degree-days^–1 for birch leaves. Alder showed the most rapid breakdown in spring–summer (0.0124 degree-days^–1), and pine the slowest (0.0016 degree-days^–1). Deciduous species exhibited general higher processing rates than evergreen species and ray-grass. The initial nitrogen and phosphorus contents were higher in riparian species leaves and ray-grass, being higher in spring (2.28±0.14% and 0.24±0.04% of nitrogen and phosphorus, respectively) than in autumn (1.88±0.36% of nitrogen and 0.18±0.03% of phosphorus). A significant correlation coefficient was found only between mean nitrogen leaf packs contents during incubation and decay rates (r=0.61; p=0.012).In deciduous species, processing was faster during the spring–summer than in the autumn–winter period, which may be attributed to the greater nutritional value and less consistency of the leaves during this season. Within evergreen species, pine had a significantly faster processing rate in autumn, attributed in this study to greater physical fragmentation of the needles. Ray-grass and eucalyptus did not exhibit any seasonal differences in processing rate.During the spring–summer period, litterfall inputs are quantitatively less important than during the autumn–winter, but due to high retention and fast breakdown during the spring–summer, green inputs should contribute substantially to nutrient incorporation and cycling in benthic communities.     

Macroinvertebrate community patterns in relation to leaf-associated periphyton under contrasting light and nutrient conditions in headwater streams

1. Temperate headwater streams traditionally have been considered heterotrophic and brown food web dominated with little primary production. Recent work, however, suggests algae on leaves in these streams may play a greater role than previously thought through interactions with microbial decomposers like fungi. Algae also may be important for macroinvertebrates colonizing leaves in streams. Algae are a more nutritious food resource for shredders than fungi and bacteria and provide a food resource for non-shredder macroinvertebrates.
2. In a field experiment, we manipulated light in three low-nutrient and three high-nutrient streams using leaf bags filled with red maple leaves in winter and spring. After four weeks we measured algal and fungal biomass, leaf stoichiometry, and macroinvertebrate abundance and biomass associated with the leaf bags. We also identified the macroinvertebrate community and examined differences in functional feeding guilds and taxa under ambient- and shaded-light treatments and low- and high-nutrient concentrations in relation to measured leaf characteristics.
3. Algal biomass on leaves was greatest in high-nutrient streams and ambient-light treatments in both seasons. Fungal biomass on leaves was greatest in high-nutrient streams and showed a moderate marginally significant positive correlation with algae during the winter. Leaf C:N was negatively correlated to algae in winter and fungi in both seasons, while leaf N:P and C:P were negatively correlated to fungi in winter and algae in spring. Interactions between fungi and algae on leaves and the nutritional importance of each for macroinvertebrates likely change across seasons, potentially impacting macroinvertebrate community composition.
4. Macroinvertebrate diversity did not differ, but biomass was significantly greater in shaded-light treatments during spring. Abundance was highest in the high-nutrient ambient-light conditions in both seasons, corresponding to greatest algal biomass. Functional feeding guild biomass and abundance were related to different leaf characteristics by season and guild. Higher algal biomass was an important factor for colonization of certain macroinvertebrates (e.g., Ephemerella (Ephemeroptera: Ephemerellidae) and Stenonema (Ephemeroptera: Heptageniidae)), while others were more abundant under shaded treatments with lower algal biomass (e.g., Tipula (Diptera: Tipulidae)), indicating taxa-specific responses.
5. Leaf-associated algae may be an important factor mediating macroinvertebrate communities associated with leaves in temperate headwater streams. Our results demonstrate that green and brown food webs intersect within leaf packs, and they cannot be easily disentangled. We therefore should consider both autochthonous and allochthonous resources within headwater streams when examining their communities or developing water management strategies.

     

Leaf litter processing and exoenzyme production on leaves in streams of different pH

We examined microbial colonization, exoenzyme activity, and processing of leaves of yellow poplar (Liriodendron tulipifera), red maple (Acer rubrum), and white oak (Quercus alba) in three streams on the Allegheny Plateau of West Virginia, United States. Leaf packs were placed in streams that varied in their underlying bedrock geology, and therefore in their sensitivity to the high level of acidic precipitation that occurs in this region. The mean pH of the streams was 4.3 in the South Fork of Red Run (SFR), 6.2 in Wilson Hollow Run (WHR), and 7.7 in the North Fork of Hickman Slide Run (HSR). Through time, the patterns of microbial biomass and exoenzyme activity were generally similar among leaf species, but the magnitude of microbial biomass and exoenzyme activity differed among leaf species. Pectinase activity was greatest in HSR, the most alkaline stream, whereas the activity of exocellulase and xylanase was greatest in WHR and SFR, the intermediate and acidic streams. This variation in the activity of different exoenzymes was consistent with published pH optima for these exoenzymes. Variation in processing rates, both among leaf species and among streams, seems to be related to the level of microbial exoenzyme activity on the leaf detritus.     

Macroconsumer effects on insect detritivores and detritus processing in a tropical stream

1. Few studies have assessed the effects of macroconsumers, such as fishes and shrimps, on detritus and detritivores.
2. We used an underwater electric field to prevent macroconsumers from feeding in and on leaf packs in a lowland stream in Costa Rica and thus to determine their effects on the density of insect detritivores and decay rates of leaves.
3. Exclusion of macroconsumers resulted in significantly higher densities of small invertebrates inhabiting leaf packs. Most of these were collector–gatherers, none were shredders.
4. Despite the increase in invertebrate density, decay rates of leaves were not statistically different. These findings contrast with results from temperate streams showing that increases in the density of invertebrates in leaf packs typically result in an increased rate of decay.
5. Leaf decay rates and invertebrate densities were also compared between leaf packs placed in electric exclusion treatments and those placed in coarse (2 cm) plastic net bags (as used in many previous studies). Our results suggest that using such netting in tropical streams may deter macroconsumers, which can affect insect density and, potentially, decay rates of organic matter.     

The influence of elevated nitrate concentration on rate of leaf decomposition in a stream

SUMMARY.

• 1 Leaf decomposition was compared in two streams at the Coweeta Hydrologic Laboratory, North Carolina. U.S.A. One stream drains an undisturbed hardwood watershed, while the other drains a successional watershed subject to an insect outbreak. The successional watershed has elevated nitrate concentrations in the streamwater.
• 2 Both black locust (Robinia pseudo-acacia) and sweet birch (Betula lenta) leaf litter decomposed 2.8 times more rapidly in the stream with high nitrate concentrations.
• 3 The more rapid decay rates appeared to be partly due to accelerated microbial processing in response to nitrate enrichment, because microbial biomass (as ATP) was higher in the nitrate-enriched stream.
• 4 At each point in time, nitrogen and phosphorus content of the litter was lower in the high nitrate stream; however, there was no significant difference in nitrogen or phosphorus content at the same state of leaf decay in the two streams.

     

Breakdown and detritivore colonisation of leaves in three New Zealand streams

Breakdown of leaves from three native riparian tree species, and their colonisation by shredding and collecting insect larvae, were investigated in three streams on Banks Peninsula, New Zealand. Leaves were introduced in baskets at the time of leaf fall. Breakdown rates of leaves were faster than previously recorded in New Zealand streams and were comparable to those of many northern hemisphere deciduous species. Shredder and total detritivore densities and biomass in leaf baskets were also greater than previously found in New Zealand streams. Peaks of shredder biomass on red beech and mahoe leaves were found when only about 20% of leaf biomass remained. No shredder peak was recorded on fuchsia leaves, and no collector peaks occurred in any of the streams. Relative shredder and collector biomass (per g DW leaf) in leaf baskets did not exceed or was smaller than in leaf litter accumulations of mixed origin and conditioning throughout the streams during leaf breakdown although absolute shredder and collector biomass (per m² stream bottom) was occasionally larger in baskets than in the rest of the stream. These findings support contentions that spatial and temporal relationships between detrital inputs and detritivore biomass and life histories are weak in New Zealand streams.     

Future increase in temperature may stimulate litter decomposition in temperate mountain streams: evidence from a stream manipulation experiment

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Bioturbation by a dominant detritivore in a headwater stream: litter excavation and effects on community structure

Bioturbation can affect community structure by influencing resource distribution and habitat heterogeneity. Bioturbation by detritivores in small headwater streams could affect community structure by reintroducing buried detrital resources into the food web and could also affect the distribution of various taxa on detritus. We evaluated the ability of the caddisfly Pycnopsyche gentilis to uncover experimentally buried leaves in a headwater stream. Packs of leaves were placed in enclosures and covered with a known volume of sediment. We added 0, 3 or 6 large Pycnopsyche to the enclosures which were permeable to most other invertebrate taxa. Leaf packs were sampled after 23 days and leaf pack mass, the amount of sediment covering the leaf packs, and macro‐ and microinvertebrate densities on leaf packs were quantified. There was a significant negative relationship between Pycnopsyche density and leaf pack mass. Pycnopsyche also reduced the volume of sediment covering leaf packs. Pycnopsyche had complex effects on the abundance of invertebrate taxa associated with the leaves. Some taxa exhibited their highest abundance in the 3 Pycnopsyche treatment while others exhibited non‐significant increases as Pycnopsyche density increased. These results suggest that the beneficial effects of Pycnopsyche (e.g. uncovering leaves which increases the availability of habitat and food) outweigh any negative effects (e.g. disturbance, encounter competition) of the caddisfly when it is present at lower densities. However, the negative impacts of Pycnopsyche appear to outweigh the positive effects via sediment removal at higher caddisfly densities for some taxa. Our results suggest that bioturbating organisms in streams have the potential to reintroduce organic matter to detrital food webs and affect the distribution and abundance of benthic taxa associated with organic matter.