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.     

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.     

Leaf diversity influences in-stream litter decomposition through effects on shredders

1. The functioning of many aquatic ecosystems is controlled by surrounding terrestrial ecosystems. In a view of growing interest in linking biodiversity to ecosystem‐level processes, we examined whether and how leaf diversity influences litter decomposition and consumers in streams. 2. We tested experimentally the hypothesis that the effects of leaf diversity on decomposition are determined by the responses of leaf consumers to resource–habitat heterogeneity. Leaves from three common riparian trees, beech (Fagus sylvatica), hazel (Corylus avellana) and ash (Fraxinus excelsior), were exposed alone and in all possible mixtures of two and three species in a stream. We analysed individual leaf species for decomposition rate, microbial respiration and mycelial biomass, and we determined the species composition, abundance and biomass of shredders in leaf bags. 3. We found that the decomposition of the fastest decomposing leaves (hazel and ash) was substantially stimulated (up to twofold higher than single species leaf packs) in mixtures containing beech leaves, which are refractory. In contrast, the decomposition of beech leaves was not affected by leaf mixing. Such species‐specific behaviour of leaves in species mixtures has been overlooked in previous studies that examined the overall decomposition of litter mixtures. 4. The effects of leaf diversity on decomposition varied with the abundance and biomass of shredders but not with microbial parameters. Beech leaves alone were less attractive to shredders than leaf packs made of hazel, ash or any mixture of species. Moreover, the presence of beech leaves in mixtures led to higher shredder abundance and biomass than we had expected from data from single species exposed alone. Lastly, we found that early instars of the caddisfly Potamophylax (the dominant shredder in terms of biomass) almost exclusively used the toughest material (i.e. beech leaves) to construct their cases. 5. Leaf pack heterogeneity may have altered shredder‐mediated decomposition. Shredders colonising diverse leaf packs benefited from the stable substratum provided by beech leaves, whereas ash and hazel leaves were primarily used as food. Thus, our findings provide strong evidence for an intimate linkage between the diversity of riparian vegetation and aquatic communities.     

Effects of crayfish on leaf processing and invertebrate colonisation of leaves in a headwater stream: decoupling of a trophic cascade

I performed a field experiment to test the hypotheses that omnivorous crayfish both promote breakdown of leaves (basal resources) and decouple any potential trophic cascade by simultaneously affecting intermediate consumers as well as their basal resource. Leaf packs were placed inside in situ artificial channels, which excluded or allowed access to crayfish. During a 4-week period, crayfish greatly promoted leaf processing, with decomposition rates among the fastest ever recorded from temperate streams. Crayfish also affected invertebrate abundance in the leaf packs. As a result of resource consumption, predation and bioturbation, crayfish treatments contained significantly lower densities of invertebrates. In contrast, exclusion of crayfish did not promote leaf decay via increased colonisation by detritivores, primarily because of the conspicuous lack of shredder insects in New Zealand streams. The results support the hypothesis that omnivorous top consumers decouple cascading chains through simultaneous direct and indirect effects on intermediate consumers and basal resources. Decapod consumers, which have been largely ignored in leaf decomposition studies, can be key leaf processors in temperate streams where shredder insects are poorly represented. Received: 8 February 2000 / Accepted: 14 April 2000     

Do macroconsumers affect insect responses to a natural stream phosphorus gradient?

We assessed the role of macroconsumers (e.g. fishes and shrimps) in affecting the response of insect assemblages to a natural phosphorus gradient formed by six streams with different phosphorus levels (range 12–350 μg/l). We hypothesized that insect responses to the phosphorus gradient would be strongest in the absence of macroconsumers. Within each stream, macroconsumers were allowed access to, or were excluded from, leaf packs using electric `fences'. Macroconsumers did not have significant effects on insects, but there was a significant phosphorus effect. Insect assemblages in high-phosphorus streams had 3–12-fold greater biomass and 3–11-fold greater abundance than assemblages in low-phosphorus streams. We also found that insect responses to phosphorus were more variable when assessed on the natural benthic substrate (e.g. mixed detritus) than in standardized leaf packs. In both substrates, the relationship between insects and phosphorus was not linear: abundance and biomass increased with phosphorus concentration to an asymptote. This suggests that insects were responding to a food resource gradient (e.g. fungi and bacteria). The Michaelis–Menten model provided a good fit for the relationship between insects and phosphorus concentrations, with half-saturation constants ranging from 12 to 60 μg SRP/l. The asymptotic relationship observed between phosphorus and insects suggests that phosphorus saturation occurred above a threshold of ca. 100 μg SRP/l. Our results provide support for the hypothesis that detritus-based food webs are mainly controlled by bottom-up forces.     

Patterns of macroinvertebrate colonization on fresh and senescent alder leaves in two Michigan streams

SUMMARY.

• 1 Communities of invertebrates colonizing senescent autumn and fresh summer alder leaves (Alnus rugosa) were compared. Leaf packs for each treatment were placed in two hardwater streams in the Upper Peninsula of Michigan in late summer and early autumn. One stream has a cobble-bottom and the other a sand-bottom and both receive fresh leaf inputs by beaver fellings.
• 2 Fresh leaf packs remained intact after 26 days immersion, but thereafter were processed faster than were the autumn leaf packs in both streams.
• 3 In the cobble-bottom stream taxon richness (S), numbers of individuals and biomass were higher on fresh than on autumn leaves.
• 4 Fresh leaves in the sand-bottom stream supported a more diverse (H'), richer (S) and more equitably distributed (J') insect fauna than did the autumn leaves.
• 5 We discuss the simultaneous lack of fresh leaf loss and the presence of more complex insect communities on those leaves during the first 26 days of the study. Invertebrates in both mid-latitude heterotrophic streams and in tropical lowland wet forest streams may rely on fresh leaf inputs, which have received little attention.

     

Macroconsumers are more important than specialist macroinvertebrate shredders in leaf processing in urban forest streams of Rio de Janeiro,Brazil

Coarse particulate organic matter is often broken down by specialist shredder invertebrates in temperate streams. In some tropical streams, larger, non-specialist, omnivorous fauna, (macroconsumers), particularly decapod shrimps and crabs, have been found to process coarse particulate matter. Larger shrimps and fish may also prey on or inhibit smaller invertebrates. Depending on the relative importance of larger and smaller fauna in leaf processing and in predatory interactions, we could expect that exclusion of larger fauna could either result in a decrease in leaf processing (if they were important in shredding or bioturbation) or increase in leaf processing if they negatively affected smaller shredders. We tested this by excluding fauna of different sizes from leaf peaks using bags with different sizes of mesh –0.2 mm (exclusion of most fauna), 2 mm (exclusion of larger fauna), and 10 mm (access to most fauna). Bag effect on leaf processing was minimized by constructing the bags of the same, fine, material, and sewing a relatively small window of the required mesh size. The experiment was conducted on two occasions in three streams of the urban forest of Parque Estadual da Pedra Branca, city of Rio de Janeiro. The three streams differed in larger fauna of shrimps (Macrobrachium potiuna), crabs, tadpoles, and fish. Packs were incubated for six time intervals and the rate of leaf processing calculated as the exponential rate of loss of leaf material. Rate of leaf processing was faster in bags with the largest mesh size; the rates in the other two mesh sizes were not statistically different. Rates varied between experiments and among streams. We could not attribute the faster leaf processing to any particular component of the larger fauna; the patterns of differences among streams and between experiments were not associated with particular taxa. There was no general trend of fewer smaller fauna in the presence of macroconsumers; the few smaller taxa that were different between mesh sizes were variously less and more abundant in the 10-mm mesh bags compared to the 2-mm. Known shredders were rare in the smaller fauna; the mining chironomid Stenochironomus was common, but was apparently not affected by larger fauna and apparently did not increase leaf processing. We conclude that macroconsumers and not smaller fauna had a positive effect on leaf processing, and this confirms a pattern observed in some other coastal Neotropical 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.     

Temporal niches of shredders in lake littorals with possible implications on ecosystem functioning

Temporal growth separation of shredders is known in streams but has not been reported from lakes. In the present study, temporal niche/trait differentiations among shredders in lakes were investigated. We sampled quantitatively three lakes in SE Sweden over a period of 18 months. Dry weight and number of individuals of the collected shredders were measured monthly. Standing stock of detritus types was also monitored. The same 10 species of lentic shredders were found in each lake, one isopod and larvae of nine trichopterans. Functionally, the shredders could be categorized into two main groups; winter and summer growing species. However, also within these groups, temporal differences in growth pattern existed. The main input of detritus occurred during the autumnal leaf fall and a majority of winter shredders had the start of their lives tied to this period. A succession in loss of detritus types was evident with easily degraded matter disappearing first followed by more resistant matter. Shredder species richness, shredder biomass per m² and the ratio coarse/fine detritus all reached its annual low in late summer. We propose a temporal link between the shredder groups and the organic matter subject to decomposition; the successive palatability of coarse detritus is likely to make a temporally separated community of shredders efficient in terms of decomposition. We believe that a temporal differentiation per se is sufficient to conclude that different impact on ecosystem function exists among shredders. Additionally we discuss impacts of differences in abundance and shredding capacity among the species.     

Nutrient addition accelerates leaf breakdown in an alpine springbrook

This study assessed the effect of nutrient enrichment on organic matter breakdown in an alpine springbrook, using alder leaf packs to which phosphorus and nitrogen were added in the form of slow-release fertilizer briquettes. The breakdown of leaf packs with nutrients added (k=0.0284 day^–1) was significantly faster than that of unfertilized packs (k=0.0137 day^–1), resulting in a 30% higher mass loss after 42 days. Unfertilized leaves enclosed in fine-mesh bags broke down at an even slower rate (k=0.0062 day^–1). Phosphorus and nitrogen concentrations were initially higher in leaf packs with nutrients added, but this difference disappeared within 3 weeks. Fungal biomass developing in decomposing leaves was substantial (c. 55 mg dry mass per 1 g leaf dry mass) although similar between fertilized and unfertilized packs, as was the sporulation activity of aquatic hyphomycetes. There was a significantly greater number and higher biomass of macroinvertebrates (shredding nemourid stoneflies in particular) on the fertilized packs, suggesting that the increased leaf mass loss was brought about by shredder feeding. Received: 11 March 1999 / Accepted: 6 September 1999     

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

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Is invertebrate shredding critical for collector invertebrates? A test of the shredder–collector facilitation hypothesis

A laboratory experiment was conducted to test the hypothesis that leaf processing by Japanese crayfish (Cambaroides japonicus de Haan) enhances the growth and reduces the mortality of collector amphipods (Sternomoera yezoensis Ueno) through fine particulate organic matter (FPOM) production. Two treatments (crayfish shredding, no shredding control) were established using flow-through artificial streams and permitting differential FPOM production rates. An amphipod enclosure, associated with juvenile amphipods and preconditioned leaf packs, was placed at the downstream end of each channel. During a 4-week period, shredding by crayfish greatly increased the amount of drifting FPOM compared to that in the control channels, in which only shredder-independent FPOM occurred. Despite the substantial contribution of crayfish to FPOM production, amphipods in both treatments grew during the 4-week period and neither amphipod survival nor growth differed between the two treatments. Furthermore, the residual leaf mass in the amphipod enclosures did not differ between the two treatments. However, residual leaf mass in amphipod enclosures was about 20% less than that expected from the processing rates of the leaf packs in the absence of invertebrate shredding. It is likely that amphipods were directly utilizing leaf packs as their food source regardless of FPOM abundance. These results indicate that juvenile amphipods, despite their classification as collectors, were plastic in their foraging habits, and that the FPOM produced by crayfish was not critical for amphipod growth or survivorship.     

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.     

Functional redundancy of stream macroconsumers despite differences in catchment land use

1. The conversion of forested landscapes to agriculture and, increasingly, to suburban and urban development significantly affects the structure and function of both terrestrial and aquatic ecosystems. While a growing body of research is examining how biotic communities change in response to human alteration of landscapes, less is known about how these changes in community structure affect biotic interactions. 2. The objective of this study was to examine top‐down control by macroconsumers (fish and crayfish) across a human‐impacted landscape. We predicted that changes in stream macroconsumers and physicochemical characteristics associated with increased catchment development (e.g. decreased abundance of fish that are obligate benthic invertivores, increased sedimentation) would diminish top‐down control of benthic insects. We expected that effects on algal assemblages would be more variable, with increased top‐down control at sites dominated by algivorous fish and diminished control elsewhere. To test these predictions, we experimentally excluded fish and crayfish from areas of the bed of five streams whose catchments ranged from 100% to <50% forested, and examined the effects of exclusion on benthic insects and algae. 3. Despite cross‐site differences in physical, chemical and biological characteristics, the outcome of our experiments was consistent across five sites representing a range of catchment development. Across all sites, macroconsumers reduced total insect biomass, largely due to decreases in Chironomidae and Hydropsychidae larvae. Macroconsumers also affected algal assemblages, reducing chlorophyll‐a and the proportion of upright and filamentous diatoms (e.g. Melosira, Cymbella) but increasing the proportion of adnate diatoms (e.g. Achnanthes) across all sites. 4. We expected that differences in factors such as macroconsumer assemblage composition, nutrient and light availability and sedimentation would result in variable responses to macroconsumer exclusion in the five streams. Contrary to these expectations, only one response variable (ash‐free dry mass) showed a statistically significant interaction (i.e. site × exclusion) effect. Most responses to exclusion were relatively consistent, suggesting functional redundancy in assemblages of macroconsumers among the sites despite differences in catchment land use.     

Positive effect of shredders on microbial biomass and decomposition in stream microcosms

1. Animals play a major role in nutrient cycling via excretory processes. Although the positive indirect effects of grazers on periphytic algae are well understood, little is known about top‐down effects on decomposers of shredders living on leaf litter. 2. Nutrient cycling by shredders in oligotrophic forest streams may be important for the microbial‐detritus compartment at very small spatial scales (i.e. within the leaf packs in which shredders feed). We hypothesised that insect excretion may cause local nutrient enrichment, so that microorganism growth on leaves is stimulated. 3. We first tested the effect of increasing concentration of ammonium (+10, +20 and +40 μg NH₄⁺ L^?1) on fungal and bacterial biomass on leaf litter in a laboratory experiment. Then we performed two experiments to test the effect of the presence and feeding activity of shredder larvae. We used two species belonging to the trichopteran family Sericostomatidae: the Palaearctic Sericostoma vittatum and the Neotropical Myothrichia murina, to test the effect of these shredders on fungal and bacterial biomass and decomposition on leaves of Quercus robur and Nothofagus pumilio, respectively. All experiments were run in water with low ammonium concentrations (2.4 ± 0.34 to 14.47 ± 0.95 μg NH₄⁺ L^?1). 4. After 5 days of incubation, NH₄ concentrations were reduced to near‐ambient streamwater concentrations in all treatments with leaves. Fungal biomass was positively affected by increased ammonium concentration. On the other hand, bacteria abundance was similar in all treatments, both in terms of abundance (bacteria cells mg^?1 leaf DW) and biomass. However, there was a tendency towards larger mean cell size in treatments with 20 μg NH₄ L^?1. 5. In the experiment with S. vittatum, fungal biomass in the treatment with insects was more than twice that in the control after 15 days. Bacteria were not detected in treatments with insects, where hyphae were abundant, but they were abundant in treatments without larvae. In the decomposition experiment run with M. murina, leaf‐mass loss was significantly higher in treatments with larvae than in controls. 6. Our hypothesis of a positive effect of shredders on fungal biomass and decomposition was demonstrated. Insect excretion caused ammonium concentration to increase in the microcosms, contributing to microbial N uptake in leaf substrata, which resulted in structural and functional changes in community attributes. The positive effect of detritivores on microbes has been mostly neglected in stream nutrient‐cycling models; our findings suggest that this phenomenon may be of greater importance than expected in stream nutrient budgets.     

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.     

Relative influence of shredders and fungi on leaf litter decomposition along a river altitudinal gradient

We compared autumn decomposition rates of European alder leaves at four sites along the Lasset–Hers River system, southern France, to test whether changes in litter decomposition rates from upstream (1,300 m elevation) to downstream (690 m) could be attributed to temperature-driven differences in microbial growth, shredder activity, or composition of the shredder community. Alder leaves lost 75–87% of original mass in 57 days, of which 46–67% could be attributed to microbial metabolism and 8–29% to shredder activity, with no trend along the river. Mass loss rates in both fine-mesh (excluding shredders) and coarse-mesh (including shredders) bags were faster at warm, downstream sites (mean daily temperature 7–8°C) than upstream (mean 1–2°C), but the difference disappeared when rates were expressed in heat units to remove the temperature effect. Mycelial biomass did not correlate with mass loss rates. Faster mass loss rates upstream, after temperature correction, evidently arise from more efficient shredding by Nemourid stoneflies than by the Leuctra-dominated assemblage downstream. The influence of water temperature on decomposition rate is therefore expressed both directly, through microbial metabolism, and indirectly, through the structure of shredder communities. These influences are evident even in cold water where temperature variation is small.     

Eucalypt leaf decomposition in an intermittent stream in south-eastern Australia

Eucalypt leaf packs were placed at two sites in an intermittent stream during summer to examine the hypothesis that terrestrially-exposed leaf litter accumulates a richer microbial flora than submerged leaves — a phenomenon observed in Canadian temporary vernal pools. This did not occur; during the experiment, microbial biomass (as ATP) rose steadily on submerged leaves but remained low on terrestrially-exposed leaves. Densities of most functional feeding groups on the submerged leaves increased with time. Scrapers appeared to be more important than shredders in eucalypt leaf breakdown at both sites.     

Detrital processing in streams exposed to acidic precipitation in the Central Appalachian Mountains

Continuing high rates of acidic deposition in the eastern United States may lead to long-term effects on stream communities, because sensitive catchments are continuing to lose anions and cations. We conducted a two-year study of the effects of pH and associated water chemistry variables on detrital processing in three streams with different bedrock geology in the Monongahela National Forest, West Virginia. We compared leaf pack processing rates and macroinvertebrate colonization and microbial biomass (ATP concentration) on the packs in the three streams. Breakdown rates of red maple and white oak leaf packs were significantly lower in the most acidic stream. The acidic stream also had significantly lower microbial and shredder biomass than two more circumneutral streams. Shredder composition differed among streams; large-particle detritivores dominated the shredder assemblages of the two circumneutral streams, and smaller shredders dominated in the acidic stream. Within streams, processing rates for three leaf species were not significantly different between the two years of the study even though invertebrate and microbial communities were different in the two years. Thus, macroinvertebrate and microbial communities differed both among streams that differed in their capacity to buffer the effects of acidic precipitation and among years in the same stream; these differences in biotic communities were not large enough to affect rates of leaf processing between the two years of the study, but they did significantly affect processing rates between acidic and circumneutral streams.The Unit is jointly sponsored by the National Biological Service, the West Virginian Division of Natural Resources, West Virginia University, and the Wildlife Management Institute.The Unit is jointly sponsored by the National Biological Service, the West Virginian Division of Natural Resources, West Virginia University, and the Wildlife Management Institute.     

Production and litter processing by crayfish in an Appalachian mountain stream

SUMMARY 1. Mean annual density and biomass () of Cambarus bartonii in an Appalachian mountain stream (U.S.A.) was 12 individuals m⁻² and 1669 mg (ash-free dry weight) m⁻².
2. Annual production ( P ) of C bartonii was 961 mg AFDW m⁻². Despite high biomass, low growth rates resulted in low production and a low P/ ratio of 0.58.
3. While C bartonii constituted 61% of the total macroinvertebrate biomass, it contributed only 13% of annual community secondary production.
4. Litter processing was positively related to temperature and crayfish size. Cambariis bartotnii was estimated to comminute 36 g m⁻² y⁻¹ of leaf litter (>1 mm²) to 24 g m−^>2 y⁻¹ fine particulate material (<1 mm²). The annual pattern of litter comminution by crayfish was regulated by temperature. As a result, >5()% of shredding activity by crayfish occurred from June to September which was also the period of lowest litter standing crops and activity of other shredding macroinvertebrates.
5. We speculate that during summer crayfish play an important role in temperate woodland streams by converting slowly processed leaf litter species (e.g. Rhododendron ) to fine particles which are then available to collector-gatherers (e.g. Chironomidae, Oligochaeta).