Effects of diversity loss on ecosystem function across trophic levels and ecosystems: A test in a detritus‐based tropical food web

Abstract We investigated the effects of biodiversity loss across trophic levels and across ecosystems (terrestrial to aquatic) on ecosystem function, in a detritus‐based tropical food web. Diversities of consumers (stream shredders) and resources (leaf litter) were experimentally manipulated by varying the number of species from 3 to 1, using different species combinations, and the effects on leaf breakdown rates were examined. In single‐species shredder treatments, leaf diversity loss affected breakdown rates, but the effect depended on the identity of the leaves remaining in the system: they increased when the most preferred leaf species remained, but decreased when this species was lost (leaf preferences were the same for all shredders). In multi‐species shredder assemblages, breakdown rates were lower than expected from single‐species treatments, suggesting an important role of interspecific competition. This pattern was also evident when oneleaf species was available but not with higher leaf diversity, suggesting that lowered leaf diversity promotes competitive interactions among shredders. The influence of diversity and identity of species across trophic levels and ecosystems on stream functioning points to complex interactions that may well be reflected in other types of ecosystem.     

How biological diversity influences ecosystem function: a test with a tropical stream detritivore guild

We investigated the relationship between diversity and ecosystem function, which is controversial and has rarely been examined for consumer assemblages, for the process of leaf breakdown by the shredder guild in a tropical stream. We manipulated species richness, evenness and identity of four macroinvertebrate shredder species (three caddisflies and one mayfly) in microcosms and tested their effect on leaf breakdown rates measured as leaf mass loss per capita and per milligram of animal. Species richness, evenness and species identity all affected leaf breakdown rates. Breakdown rates tended to increase with higher richness, but only for the three caddisflies, probably through a release of intraspecific interference, although other mechanisms such as niche complementarity or facilitation cannot be discarded. Leaf breakdown by the caddisflies was reduced in the presence of the mayfly, possibly because of its mode of movement by swimming instead of crawling and its similarity to some predators that are common in leaf litter. Species identity was more important than species richness in determining leaf breakdown rates, indicating that some species within the shredder guild are not redundant, and suggesting important consequences of particular species loss for the functioning of the ecosystem.     

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.     

Leaf litter breakdown, microbial respiration and shredder production in metal-polluted streams

1. If species disproportionately influence ecosystem functioning and also differ in their sensitivities to environmental conditions, the selective removal of species by anthropogenic stressors may lead to strong effects on ecosystem processes. We evaluated whether these circumstances held for several Colorado, U.S.A. streams stressed by Zn. 2. Benthic invertebrates and chemistry were sampled in five second–third order streams for 1 year. Study streams differed in dissolved metal concentrations, but were otherwise similar in chemical and physical characteristics. Secondary production of leaf‐shredding insects was estimated using the increment summation and size‐frequency methods. Leaf litter breakdown rates were estimated by retrieving litter‐bags over a 171 day period. Microbial activity on leaf litter was measured in the laboratory using changes in oxygen concentration over a 48 h incubation period. 3. Dissolved Zn concentrations varied eightfold among two reference and three polluted streams. Total secondary production of shredders was negatively associated with metal contamination. Secondary production in reference streams was dominated by Taenionema pallidum. Results of previous studies and the current investigation demonstrate that this shredder is highly sensitive to metals in Colorado headwater streams. Leaf litter breakdown rates were similar between reference streams and declined significantly in the polluted streams. Microbial respiration at the most contaminated site was significantly lower than at reference sites. 4. Our results supported the hypothesis that some shredder species contribute disproportionately to leaf litter breakdown. Furthermore, the functionally dominant taxon was also the most sensitive to metal contamination. We conclude that leaf litter breakdown in our study streams lacked functional redundancy and was therefore highly sensitive to contaminant‐induced alterations in community structure. We argue for the necessity of simultaneously measuring community structure and ecosystem function in anthropogenically stressed ecosystems.     

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

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Leaf litter breakdown rates in boreal streams: does shredder species richness matter?

1. Leaf litter breakdown rates were assessed in 23 boreal streams of varying size (first–seventh order) in central and northern Sweden. 2. Shredders were most abundant in small streams, while shredder species richness showed a hump-shaped relationship with stream order, with most species in fourth order streams. 3. In a partial least-squares regression analysis, year, water temperature, shredder species richness and shredder abundance were those factors correlating most strongly with leaf breakdown rates. Shredder species richness was more strongly correlated with leaf litter breakdown rates than shredder abundance, and shredder biomass showed no such correlation. 4. These data suggest that shredder species richness is an important variable in terms of leaf litter dynamics in streams.     

Comparison of leaf processing rates under different temperature regimes in three headwater streams

1. We studied the effects of different temperature regimes on leaf litter processing in three forested Appalachian headwater streams of different pH (mean pH = 4.2, 6.5, 7.5).
2. We compared leaf breakdown rates, microbial biomass and macroinvertebrate shredder density and biomass between two 12-week processing periods (October–January and November–February) in each stream. Leaf processing rates were calculated both as k (day^–1) and k _d (degree day^–1).
3. There were no significant differences in processing rates ( k day^–1) between the two study periods for any leaf species in any stream. The average difference in temperature between the two study periods was 175 degree days. Shredder density was significantly higher during the earlier study period on 40% of the sample dates, but shredder biomass was not significantly different between the two study periods. ATP concentration was significantly higher during the early study period for 60% of the sample dates.
4. More significant differences in these variables (shredder density and biomass, ATP concentration) were seen among the three study streams than between the two study periods. This indicates that in this study other factors, particularly stream pH, contributed more to processing rate variation than did differences in thermal regime.     

Drought impact on stream detritivores: experimental effects on leaf litter breakdown and life cycles

Predictions of effects of global climate change include decreased runoff for many parts of the world, which will result in drying of streams. Information of the effects of drought on aquatic ecosystems is limited and little is known of the effects on ecosystem functions. Our main objective was to measure the direct effects of drought on leaf litter breakdown by invertebrate shredders in a controlled laboratory experiment. We hypothesized a decreased breakdown at high drought level. Single-species and multi-species treatments with three shredder species (Asellus aquaticus, Limnephilus bipunctatus, and L. flavicornis) were set up in an experiment with three drought level treatments, control, medium, and high drought (6 cm water level, 1 cm water level, and water level below sediment surface, respectively). Breakdown measured as leaf litter loss was significantly lower in both medium and high drought treatments compared to the control. Previously, decreased breakdown due to drying has been reported, but attributed to low densities of invertebrate shredders. We show that even when shredders are present, drought decreases the breakdown. Drought treatments also induced earlier pupation for the caddisfly L. flavicornis. Shifts in species phenology due to drought, e.g., earlier emergence, may affect species ability to adult survival and reproduction. Shifts in timing of emergence may also affect terrestrial food webs, where emerging aquatic insects may constitute an important food subsidy. Our knowledge of the complex effects of droughts in aquatic systems is limited with an urgent need of extended knowledge of the ecological effects of droughts on freshwater ecosystem functioning.     

Tadpoles enhance microbial activity and leaf decomposition in a neotropical headwater stream

1. Relationships between biodiversity and ecosystem function are of increasing interest, particularly in freshwater ecosystems where species losses are occurring at unprecedented rates. Amphibian declines have been associated with a loss of ecosystem function in neotropical streams, but little is known of the potential roles of stream‐dwelling tadpoles in leaf decomposition. Leaf litter is an important energy source to streams, and the breakdown of this material to fine particulate organic matter (FPOM) is a key ecosystem function. 2. We used mesocosms in a natural stream setting to quantify the effects of grazing tadpoles, shredding macroinvertebrates and a combination of the two on leaf decomposition and associated microbial activity. We measured respiration rates of decomposing leaves, particulate organic matter (POM) and leaf biofilm biomass and C : N : P ratios, and leaf area loss in 4 treatments: Control, tadpole only (TP), tadpole and shredding macroinvertebrates (TP + INV) and shredding macroinvertebrates only (INV). We hypothesised that tadpoles would enhance leaf decomposition by changing nutrient availability and stimulating microbial activity. 3. Respiration rates ranged from 3.1 to 6.0 mg O₂ dry mass^?1 h^?1 and were significantly higher in the TP and TP + INV treatments than in the control. The TP + INV treatment had significantly higher POM in chambers than the control and INV treatments. The TP treatment had significantly lower leaf biofilm biomass than the control and INV treatments. 4. Tadpoles influenced the elemental balance of C and N in POM and leaf biofilm. In contrast to our prediction, molar C : N ratios were higher in the TP + INV treatment than in the control. Mean molar N : P ratios in POM were higher in the TP + INV treatment than in any other treatment. Leaf biofilm followed a similar pattern, but both TP and TP + INV had significantly higher N : P ratios than the control and INV treatments. Leaf area loss was greatest when tadpoles and invertebrates were together (TP + INV = 0.6% leaf area loss per mg organism) than separate (TP = 0.1%, INV = 3%), indicating facilitation. 5. Tadpoles indirectly affected leaf decomposition by influencing microbial communities and macroinvertebrate feeding. As such, ongoing amphibian declines may adversely affect a critical ecosystem function in freshwater habitats.     

Effects of shredding amphipod density on watercress Nasturtium officinale breakdown

Shredding stream invertebrates should have a positive influence on the breakdown rates of leaf litter via direct consumption and particle fragmentation. To determine the effects of shredder density on litter breakdown, breakdown of the emergent stream macrophyte, Nasturtium officinale , was investigated using three litter bag mesh sizes [fine (0.2 mm), medium (1 mm) and coarse (3 mm) mesh] and four stocking densities of the shredder, Gammarus pseudolimnaeus , (0, 4, 8 and 16 per bag). Watercress decayed very rapidly, with breakdown rates ( k values) ranging from 0.075 d^-1 for fine mesh with no shredders to 0.24 d^-1 for coarse mesh. Stocked Gammarus increased breakdown rates significantly in fine mesh bags (p < 0.001), but only marginally in medium mesh bags (p < 0.1). Breakdown rates also increased significantly with mesh size. A regression model showed a significant relation of breakdown rate to Gammarus density and mesh size. These results clearly show that shredders can significantly influence breakdown rates and can account for up to 30% of breakdown, but that mesh size effects such as particle size reduction and loss are also very important.     

Comparison of leaf processing rates under different temperature regimes in three headwater streams

1. We studied the effects of different temperature regimes on leaf litter processing in three forested Appalachian headwater streams of different pH (mean pH = 4.2, 6.5, 7.5).
2. We compared leaf breakdown rates, microbial biomass and macroinvertebrate shredder density and biomass between two 12-week processing periods (October–January and November–February) in each stream. Leaf processing rates were calculated both as k (day^–1) and k _d (degree day^–1).
3. There were no significant differences in processing rates ( k day^–1) between the two study periods for any leaf species in any stream. The average difference in temperature between the two study periods was 175 degree days. Shredder density was significantly higher during the earlier study period on 40% of the sample dates, but shredder biomass was not significantly different between the two study periods. ATP concentration was significantly higher during the early study period for 60% of the sample dates.
4. More significant differences in these variables (shredder density and biomass, ATP concentration) were seen among the three study streams than between the two study periods. This indicates that in this study other factors, particularly stream pH, contributed more to processing rate variation than did differences in thermal regime.     

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 a predatory fish on a tropical detritus-based food web

In contrast to that for grazing systems, relatively little information exists for trophic cascades in detritus-based stream food webs, which are predominant in forested headwater streams. Predator–prey interactions are thought to be weak in these systems, but studies are very scarce, their results are equivocal, and they do not separate the effect of direct consumption from a behavioural response of shredders. We examined the effect of predatory fish on leaf litter breakdown in headwater tropical Australian streams at three levels: (1) the behavioural response of shredder species to predator presence as indicated by chemical cues; (2) the rates of leaf breakdown resulting from shredder activity; and (3) the relationship between shredder species richness and leaf breakdown rates. Our results suggest that predatory fish can have a trait-mediated effect on detritus-based food webs in streams, by reducing consumer activity. We identified reductions in short-term overall activity in response to the presence of predatory fish cues, comparable to those found for grazers. We also observed a visible, albeit statistically non-significant, reduction in consumption rates. Shredder species richness did not affect leaf breakdown rates, and fish presence did not modify this relationship or the differences in breakdown rates among species, suggesting that the overall reduction in leaf breakdown caused by fish presence is due to a reduction in activity in every species. Thus, our laboratory studies have shown that there can be a behavioural basis for trait-mediated trophic cascades linked to fish presence in detrital food webs in streams. However, the strength of fish effects depends on environmental circumstances, and field studies of litter breakdown in streams with and without predatory fish are required if we are to elucidate the ecological significance of our observations.     

Identifying the Relative Importance of Leaf versus Shredder Species Loss on Litter Decomposition in Streams

Understanding how species loss influences ecosystem function is a contemporary issue in ecology. However, most research has focused on species loss at one trophic‐level. We explored the relationship between functional diversity (FD) and species richness separately for trees and aquatic leaf‐shredding detritivores. For trees, we collected information on species‐specific leaf tissue chemistry and species co‐occurrences in the mid‐Atlantic region (USA). For shredders, we used a published trait database with information on communities from 38 streams in the same region. We used a clustering algorithm to estimate FD for each community and for randomly assembled communities. If FD was high, we concluded that species loss was important to change in function; if low, species were functionally redundant and insensitive to species loss. We found tree FD to be significantly different than expected, but shredders exhibited FD levels similar to patterns based on random assembly. Furthermore, there were more leaf species exclusively associated with very high or very low levels of functional diversity compared to shredders. This approach revealed greater implications for leaf than shredder species loss for litter breakdown. (© 2009 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

The Importance of Detritivore Species Diversity for Maintaining Stream Ecosystem Functioning Following the Invasion of a Riparian Plant

The strength of linkages between riparian plants and stream communities can be expected to be influenced by invading plants. While most studies so far have been focussed on the effects of the leaf litter quality of the invader, this study addresses the impact of detritivores on the pool of detritus. In a natural setting, we found that species richness of shredding macroinvertebrates significantly influenced the breakdown rate of an invasive weed species, the Japanese knotweed (Fallopia japonica), which has become a major plant invader along streams and rivers in Europe and North America. Our findings imply that a reduction of the diversity of shredder species, which may be the result of disturbances, could negatively influence stream ecosystems' capacity of processing knotweed leaves. Although the knotweed showed breakdown rates similar to those of common native tree and shrub species, other exotic leaf species might show considerably slower rates and hence have greater consequences for the ecosystems. We have, in this study, indicated a technique by which the effects of other non-indigenous plants on ecosystem functioning might be considered.     

Leaf breakdown in streams differing in catchment land use

1. The impact of changes in land use on stream ecosystem function is poorly understood. We studied leaf breakdown, a fundamental process of stream ecosystems, in streams that represent a range of catchment land use in the Piedmont physiographic province of the south‐eastern United States. 2. We placed bags of chalk maple (Acer barbatum) leaves in similar‐sized streams in 12 catchments of differing dominant land use: four forested, three agricultural, two suburban and three urban catchments. We measured leaf mass, invertebrate abundance and fungal biomass in leaf bags over time. 3. Leaves decayed significantly faster in agricultural (0.0465 day^?1) and urban (0.0474 day^?1) streams than in suburban (0.0173 day^?1) and forested (0.0100 day^?1) streams. Additionally, breakdown rates in the agricultural and urban streams were among the fastest reported for deciduous leaves in any stream. Nutrient concentrations in agricultural streams were significantly higher than in any other land‐use type. Fungal biomass associated with leaves was significantly lower in urban streams; while shredder abundance in leaf bags was significantly higher in forested and agricultural streams than in suburban and urban streams. Storm runoff was significantly higher in urban and suburban catchments that had higher impervious surface cover than forested or agricultural catchments. 4. We propose that processes accelerating leaf breakdown in agricultural and urban streams were not the same: faster breakdown in agricultural streams was due to increased biological activity as a result of nutrient enrichment, whereas faster breakdown in urban streams was a result of physical fragmentation resulting from higher storm runoff.     

The role of timing, duration, and frequency of inundation in controlling leaf litter decomposition in a river-floodplain ecosystem (Tagliamento, northeastern Italy)

Despite growing recognition of the importance of a natural flow regime in river-floodplain systems, researchers struggle to quantify ecosystem responses to altered hydrological regimes. How do frequency, timing, and duration of inundation affect fundamental ecosystem processes such as leaf litter decomposition? Along the semi-natural Tagliamento River corridor, located in northeastern Italy, we employed in situ experiments to separate effects of different inundation components on breakdown rates of black poplar (Populus nigra). We used a litter-bag method with two different mesh sizes to investigate how fungi and macroinvertebrates influence leaf breakdown rates. Ten treatments, each representing a specific combination of duration and frequency of inundation, were deployed in two seasons (summer, winter) to mimic complex inundation patterns. After 30 days of exposure, mean percentage of remaining leaf litter (ash free dry mass) ranged between 51% (permanent wet) and 88% (permanent dry). Leaf breakdown was significantly faster in winter than in summer. Duration of inundation was the main inundation component that controlled leaf breakdown rates. Leaf-shredding macroinvertebrates played only a role in the permanent wet treatment. Fungal parameters explained the faster leaf breakdown in winter. Our study suggests that modifications of the inundation regime will directly modify established decomposition processes. Factors reducing duration of inundation will decelerate leaf breakdown rates, whereas a decrease in flow variation will reduce leaf breakdown heterogeneity.     

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.     

Ecological divergence of <Emphasis Type="Italic">Chaetopteryx rugulosa</Emphasis> species complex (Insecta,Trichoptera) linked to climatic niche diversification

Climate change may affect species diversity and, consequently, ecological processes such as leaf decomposition. We evaluated the effects of increased temperature and carbon dioxide (CO₂) on fungal biomass, leaf breakdown, and on survival and growth of the shredder Phylloicus elektoros. We hypothesized that climatic changes would result in lower survival and growth of shredders and lower leaf consumption by these organisms. On the other hand, we predicted an increase in fungal biomass in response to climatic changes. We conducted an experiment in Manaus, Brazil, using four microcosms that simulate real-time air temperature and CO₂ (control chamber), as well as three other chambers subjected to fixed increases in temperature and CO₂ as compared to the control chamber. The “extreme” condition represented an increase of ~4.5°C in temperature and ~870 ppm in CO₂ in relation to the control chamber. Total and shredder leaf-breakdown rates, fungal biomass, and shredder survival rates were significantly lower in warmer and CO₂ concentrated atmospheres. Shredder growth rate and leaf breakdown by microorganisms were similar among all climatic conditions. With climatic changes, we found an increase in the relative importance of microorganisms on leaf-breakdown rates as compared to shredders. Thus, lower leaf breakdown and a change in the main decomposer due to future climatic conditions may result in major changes in the pathways of organic matter processing and, consequently, in aquatic food webs.     

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.