Shredder Community Research Articles

Functional diversity of shredders, not species richness, drives the decomposition rate of leaf litter in ponds

Leaf litter decomposition is a critical ecosystem-level process in many freshwater habitats. Although ponds are likely to derive a large proportion of their energy from riparian vegetation, allochthonous organic matter decomposition in these water bodies has received little attention. We studied the breakdown rates of black alder (Alnus glutinosa (L.) Gaertn.) litter in ponds and provide the first evidence of the role of the taxonomic and functional diversity of pond-dwelling shredders in this ecosystem process. Despite a strong connection to riparian zones, the litter breakdown rates observed in ponds were generally lower than those reported in headwater streams. It seems that ponds provide less favorable conditions for shredder communities than headwaters. The rate of organic matter decomposition in ponds was significantly positively related to functional diversity, represented by the variability of shredder body size, while shredder species richness did not appear to be a reliable proxy for this ecosystem function. This finding is consistent with theoretical predictions that functional complementarity among species has a systematic effect on ecosystem processes. It also emphasizes that body size is a crucial functional trait mediating the effects of shredder diversity on litter decomposition in ponds.

Chemistry Matters: High Leaf Litter Consumption Does Not Represent a Direct Increase in Shredders' Biomass.

Changes in riparian vegetation can alter the input and quality of leaf litter in aquatic ecosystems, but the effects of these changes on litter fragmentation by invertebrate shredder communities in tropical streams remain poorly studied. The caddisfly genus Phylloicus Müller, 1880 (Trichoptera: Calamoceratidae) is highly abundant in Neotropical streams, representing a great part of shredder biomass, which uses the allochthonous litter as a food resource and for case-building. We investigated leaf consumption by Phylloicus sp. under different leaf conditioning (leached and unleached) and plant species (Eucalyptus grandis, Erythrina falcata, and Inga uruguensis). The effects of leaf conditioning and plant species were measured using microcosm treatments, with one free Phylloicus sp. larva per 2-l microcosm, and a decomposition control to correct for microbial decomposition. Our study suggests that phosphorus and caloric values of leaf litter are more important than leaf hardness and nitrogen in driving leaf consumption by Phylloicus sp. On the one hand, higher consumption was observed in treatment with unleached leaves than in leached leaf treatment due to higher nutrient concentration and caloric values on unleached leaves. On the other hand, Phylloicus sp. larvae preferred leached leaves for case building over unleached leaves, as leached leaves are less prone to the activity of the decomposing community, thus lowering the need for constant case renewal. Finally, high litter consumption is not necessarily converted into biomass by Phylloicus sp. larvae. In this sense, Phylloicus sp. larvae showed selectivity for resources with high caloric content for consumption and low caloric content for case-building.

Leaf breakdown rates as a functional indicator were influenced by an invasive non-native invertebrate in urban ponds

Impacts on biodiversity and the functioning of ecosystems are often overlooked in small waterbodies as they are not regularly monitored as part of the Water Framework Directive (WFD). These small systems are, however, often essential for freshwater biodiversity and are frequently characterized by high beta-diversity. In this study we examined shredder diversity and ecosystem functioning (leaf litter breakdown) as indicators of environmental stress along a gradient of urbanisation. The native isopod Asellus aquaticus and the non-native shrimp Crangonyx pseudogracilis (Amphipoda) were dominant species of the shredder communities in 26 study ponds. Variation in shredder community composition among ponds was explained by pond surface area (P < 0.05) and the coverage of impermeable surface within 500 m (P < 0.05), indicative of urban impact, which was strongly, and negatively correlated with shredder richness. Site mean total leaf breakdown was 0.0020 k d−1 (range 0.0003 – 0.0053), whilst breakdown rates owing to microbial and fungal activity were lower and averaged 0.0010 k d−1 (range 6.72−5 − 0.0036). Both total and microbial activity driven breakdown rates showed relatively weak relationships with the environmental gradient reflecting degrees of urbanization. A model-averaging procedure suggested that total litter breakdown rates could be attributed to activity of shredders with A. aquaticus and C. pseudogracilis contributing most, explaining 65% of the variance (R2 = 0.65, F = 13.7, P < 0.001), which both significantly and similarly predicted total leaf breakdown rates (P < 0.05). Total leaf breakdown rates were significantly correlated with the densities of both C. pseudogracilis (P < 0.001) and A. aquaticus (P < 0.001), however partial correlations indicated that the former showed the strongest association.The process of urbanisation can have significant, and negative impacts upon freshwater ecosystems, including small water bodies such as ponds both regarding biodiversity and ecosystem functioning. The findings of the current study suggest that ecosystem functioning indicators such as leaf litter decomposition should be paired with structural measures. This is because function sustained by tolerant invading species would not otherwise be detected, and their presence is increasingly likely due to climate change and urbanization.

Substrate homogenization affects survival and fitness in the lowland stream caddisflies Micropterna sequax and Potamophylax rotundipennis: a mesocosm experiment

AbstractLoss of substrate heterogeneity or patchiness is common in lowland streams with disturbed hydrological regimes. At the reach scale, peak discharges tend to homogenize the stream bed and decrease the availability of specific microhabitat types. This spatial shift in habitats toward a more homogeneous landscape could have large negative effects on species that perform essential ecosystem processes. We conducted an aquatic mesocosm experiment to test the effect of habitat homogenization on survival and fitness of 2 species of Trichoptera (Micropterna sequax and Potamophylax rotundipennis). We used caddisflies as model organisms because of their abundance in lowland streams, their representativeness of the total shredder community, and their important role in leaf-litter decomposition. We reared larvae in artificial recirculating channels containing leaf and sand patches in 3 spatial configurations, differing in homogeneity of substrates, varying from few large patches to many small patches. We used e...

Shifts in shredder communities and leaf breakdown along a disrupted stream continuum

AbstractStream ecosystem functions are predicted to vary longitudinally along a stream network as a result of changes in physical and biotic processes. Conversion by headwater communities of allochthonous materials, such as leaves, into readily available sources of energy is an essential process for maintenance of downstream communities. We investigated how location along a stream continuum, the presence of a dam, and leaf type affected leaf breakdown. We placed yellow birch (Betula alleghaniensis), sugar maple (Acer saccharum), and white oak (Quercus alba) leaf packs in Sims Creek (North Carolina, USA), at a 1st-order site, a 2nd-order site above a dam, and a 2nd-order site immediately below this dam. Leaf breakdown rates of all species were slower at the above-dam than at the headwater site, and breakdown rates for birch and maple were lower at the below- than the above-dam site. Detritivore composition and abundance differed among sites. Leaf packs had lower detritivore abundance at the below- than the...

Effects of fungicides on decomposer communities and litter decomposition in vineyard streams

Large amounts of fungicides are applied globally and partly enter freshwater ecosystems. A few laboratory studies examined their effects on decomposer communities and the ecosystem process of litter decomposition (LD), whereas the field situation remains largely unknown. We conducted a field study with 17 stream sites in a German vineyard area where fungicides represent the dominant pest control agent. Passive samplers were used to monitor 15 fungicides and 4 insecticides in streams and their toxicity was described using the toxic unit approach, whereas sediment samples were taken to characterise total copper concentrations. Microbial and leaf-shredding invertebrate community composition and related LD rates were assessed at each site. The structure of microbial and shredder communities as well as fungal biomass changed along the fungicide toxicity gradient. The changes in microbial endpoints were associated with a reduction of microbial LD rate of up to 40% in polluted streams. By contrast, neither the invertebrate LD rate nor in-situ measured gammarid feeding rates correlated with fungicide toxicity, but both were negatively associated with sediment copper concentrations. A subsequent laboratory experiment employing field fungicide concentrations suggested that the microbial community changes are causal. Overall, our results suggest that fungicides can affect LD under field conditions.

Leaf-litter breakdown in streams of East Malaysia (Borneo) along an altitudinal gradient: initial nitrogen content of litter limits shredder feeding

AbstractBreakdown of litter in streams is an important terrestrial–aquatic C pathway in forests. Macroinvertebrate detritivore shredders that feed on litter may be more abundant in temperate than tropical streams because the feeding guild is generally restricted to cool waters and because temperate streams may have more high-quality resources. However, rich shredder communities have been found in some high-altitude tropical streams. We investigated litter breakdown and the variation in resource quality of native riparian vegetation in tropical streams at different altitudes (86–3393 m) in eastern Malaysia (Borneo). At each stream, we collected 3 native local leaf species, analyzed their initial resource quality, and incubated them for 56 d in coarse- (shredder and microbial colonization) and fine- (only microbial colonization) mesh bags. We incubated Macaranga tanarius litter in all streams to compare breakdown rates directly. Shredder-mediated breakdown of native litter was detected in only 5 of 24 cases...

Effects of invasive European bird cherry (Prunus padus) on leaf litter processing by aquatic invertebrate shredder communities in urban Alaskan streams

European bird cherry (Prunus padus) (EBC) is an invasive ornamental tree that is spreading rapidly in riparian forests of urban Alaska. To determine how the spread of EBC affects leaf litter processing by aquatic invertebrate shredders, we conducted complementary leaf pack experiments in two streams located in Anchorage, Alaska. The first experiment contrasted invasive EBC with three native tree species—thin-leaf alder (Alnus tenuifolia), paper birch (Betula neoalaskana), and black cottonwood (Populus trichocarpa)—in one reach of Chester Creek; finding that EBC leaf litter broke down significantly faster than birch and cottonwood, but at a similar rate to alder. The second experiment contrasted EBC with alder in four reaches of Campbell and Chester creeks; finding that while EBC leaf litter broke down significantly faster than alder in Chester Creek, EBC broke down at a similar rate to alder in Campbell Creek. Although EBC sometimes supported fewer shredders by both count and mass, shredder communities did not differ significantly between EBC and native plants. Collectively, these data suggest that invasive EBC is not currently exhibiting strong negative impacts on leaf litter processing in these streams, but could if it continues to spread and further displaces native species over time.

The effect of shredder community composition on the production and quality of fine particulate organic matter

AbstractDecomposition of senesced primary production starts processing chains in aquatic systems. Shredding macroinvertebrates convert coarse particulate organic matter (CPOM) to fine particulate organic matter (FPOM) that supports 2 other feeding groups, collecting and filtering macroinvertebrates. This linkage is often invoked by aquatic ecologists, but the effect of detritivore assemblage composition on production of FPOM is relatively understudied. I manipulated detritivore assemblage composition (Limnephilus sp., Caecidotea sp., and Hyalella azteca) in aquatic mesocosms stocked with green speckled alder leaves (Alnus incana rugosa). I measured production rate, size distribution, and stoichiometry of FPOM produced through time. Detritivore species richness had a positive effect on FPOM production resulting from inclusion of the functionally dominant shredder, Limnephilus sp., in mixed-species treatments (e.g., sampling effect). Mixed-species treatments had significantly faster particle production than...

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.

Differential leaf‐litter processing by native (Gammarus pulex) and invasive (Dikerogammarus villosus) freshwater crustaceans under environmental extremes

ABSTRACT There is increasing interest in understanding the implications of biological invasions within the context of ecosystem functioning. Non‐native crustaceans are of particular interest within fresh waters because of their important contributions to leaf‐litter processing. The alien amphipod Dikerogammarus villosus is spreading rapidly through Europe where it has displaced native gammarids including Gammarus pulex. The resultant change in shredder communities has considerable implications for the dynamics of resource availability within invaded systems. The invasive success of D. villosus has been attributed to many characteristics, including its ability to tolerate a wide range of environmental conditions and to exploit a wide feeding niche. These characteristics are thought to give D. villosus an advantage over native species, especially in habitats subject to changing environmental conditions. The leaf (Salix alba) shredding efficiency, defined as the leaf mass consumed over the 4‐day experiment per amphipod‐day, was measured for D. villosus and G. pulex under extreme temperature and conductivity conditions, in single species and mixed species aquaria. At high temperatures (25°C), D. villosus shredded significantly more leaves than size‐matched G. pulex. Conductivity extremes had no species effect; overall leaf‐shredding efficiency was significantly greater at low conductivity (250 μS cm‐1) than high conductivity (1870 μS cm‐1). An inspection of daily leaf disc consumption found that the two species showed significantly different leaf consumption patterns, with D. villosus consuming more leaves earlier in the experiment. These results suggest that D. villosus invasion could lead to ecosystem‐level changes in leaf processing, such as greater leaf processing earlier in autumn and at higher temperatures, which could alter nutrient dynamics and community assemblages within invaded systems. Such effects may not be reflected in biotic indices used to assess ecological status, as these indices do not currently distinguish between native and invasive species in the same family. Copyright © 2013 John Wiley &amp; Sons, Ltd.

Stream ecosystem integrity is impaired by logging and shifting agriculture in a global megadiversity center (Sarawak, Borneo)

In common with most of Borneo, the Bakun region of Sarawak is currently subject to heavy deforestation mainly due to logging and, to a lesser extent, traditional slash-and-burn farming practices. This has the potential to affect stream ecosystems, which are integrators of environmental change in the surrounding terrestrial landscape. This study evaluated the effects of both types of deforestation by using functional and structural indicators (leaf litter decomposition rates and associated detritivores or ‘shredders’, respectively) to compare a fundamental ecosystem process, leaf litter decomposition, within logged, farmed and pristine streams. Slash-and-burn agricultural practices increased the overall rate of decomposition despite a decrease in shredder species richness (but not shredder abundance) due to increased microbial decomposition. In contrast, decomposition by microbes and invertebrates was slowed down in the logged streams, where shredders were less abundant and less species rich. This study suggests that shredder communities are less affected by traditional agricultural farming practices, while modern mechanized deforestation has an adverse effect on both shredder communities and leaf breakdown.

Predicting richness effects on ecosystem function in natural communities: insights from high-elevation streams

. Despite the increased complexity of experimental and theoretical studies on the biodiversity-ecosystem functioning (B-EF) relationship, a major challenge is to demonstrate whether the observed importance of biodiversity in controlled experimental systems also persists in nature. Due to their structural simplicity and their low levels of human impacts, extreme species-poor ecosystems may provide new insights into B-EF relationships in natural systems. We address this issue using shredder invertebrate communities and organic matter decomposition rates in 24 high-altitude (3200-3900 m) Neotropical streams as a study model. We first assessed the effects of stream characteristics and shredder diversity and abundance on organic matter decomposition rates in coarse- and fine-mesh bags. We found the interaction term shredder richness x shredder abundance had the most significant impact on decomposition rates in the field, although water discharge may also play a role locally. We also examined the relative contribution of the three most abundant shredders on decomposition rates by manipulating shredder richness and community composition in a field experiment. Transgressive overyielding was detected among the three shredder species, indicating complementary resource use and/or facilitation. By integrating survey and experimental data in surface response analyses we found that observed B-EF patterns fit those predicted by a linear model that described litter decomposition rates as a function of increasing shredder richness and the relative abundance of the most efficient shredders. Finally, the validity of our approach was tested in a broader context by using two independent but comparable data sets from 49 French and Swedish streams showing more complex shredder community structure. Results revealed that richness and identity effects on decomposition rates were lost with increasing shredder community complexity. Our approach of combining experimental and empirical data with modeling in species-poor ecosystems may serve as an impetus for new B-EF studies. If theory can explain B-EF in low-diversity ecosystems, it may also have credibility in more complex ones.

Invertebrate community structure and ecosystem functioning in European conifer plantation streams

Summary1. Terrestrial leaf‐litter is the dominant energy input to many headwater streams and consequently the nature of the riparian vegetation can have profound effects on in‐stream processes. The impact of conifer plantations on community structure and ecosystem functioning (litter breakdown) was investigated in field experiments in three countries (Britain, Ireland, Poland), each representing a distinct European ecoregion. Twenty‐six streams were used in the trial: half were bordered with broadleaved and the other half with conifer riparian vegetation.2. In a leaf breakdown study using litter bags, two leaf types (oak and alder) were used to assess the impact of resource quality and two mesh sizes (10 and 0.5 mm aperture) were used to gauge the relative importance of invertebrate detritivores and microbial decomposers respectively. Comparisons were made between vegetation types and among regions; pH varied among individual streams but, unlike many previous studies, it was not confounded with vegetation type, enabling us to isolate the effect of vegetation more effectively.3. Overall, riparian vegetation type did not affect breakdown rates but strong regional differences were observed. There was also a significant interaction between these two variables, but this disappeared after fitting pH as a covariable, demonstrating its importance in determining breakdown rates and raising the possibility that in previous studies the impacts of conifer plantations might have been confounded with pH.4. Shredder species composition differed between vegetation types. Small stoneflies were most strongly associated with conifer streams; broadleaved streams generally had a higher proportion of larger taxa, such as limnephilid caddisflies and gammarid shrimps, although the latter were excluded from sites with low pH. However, breakdown rates were maintained irrespective of shredder community composition, suggesting a high degree of functional redundancy in these communities. Similar processing rates were observed between streams with high numbers of nemourids and those with only a few limnephilids or gammarids, suggesting that density compensation among consumers might stabilise process rates.5. Our results suggest that leaf‐litter breakdown can be an effective proxy for assessing stream ecosystem functioning, as rates differed significantly across spatial scales, from between streams to across regions and responded to an environmental gradient (pH). The litter bag technique can also complement traditional assessment methods by providing valuable information on the composition of consumer guilds, thereby providing an important link between structure and function that is needed to help inform management practices.

Grass, mosses, algae, or leaves? Food preference among shredders from open-canopy streams

Shredder feeding is a vital process in making decomposition products available to biota in streams. To investigate which food sources shredders in open-canopy streams exploit, we conducted a feeding preference experiment with the invertebrate detritivores Limnephilus bipunctatus and Nemoura sp., which are commonly found in open-canopy streams on the Swedish island of Oland in the southern Baltic Sea. Leaves of birch, Swedish whitebeam, and shrubby cinquefoil; dead and fresh grass; water moss; and algae were offered to the shredders in multi- and single treatments. We hypothesized that food with high nutritional value would be preferred. Both taxa preferred leaves of shrubby cinquefoil, a bush common in the riparian zone of Oland streams; additionally Nemoura sp. also chose algae. Dead grass, the most abundant food source in the streams during the whole year, was the least consumed food type. The fresh food types had highest nutritional value, measured as carbon to nitrogen content. Therefore, food quality could not alone explain the preference of shrubby cinquefoil. However, among the detritus type offered, shrubby cinquefoil had the highest nutritional value. Shrubby cinquefoil may constitute one important energy source to these open-canopy stream ecosystems and may be essential in maintaining an abundant shredder community in these streams. Thus, the results of this study indicate that detrital resources are indeed important in open-canopy stream systems.

The loss of aquatic and riparian plant communities: Implications for their consumers in a riverine food web.

Human induced alterations to rivers and steams have resulted in significant changes to the structure and diversity of riparian and aquatic plant communities. These changes will impact on the dynamics of riverine carbon cycles and food web structure and function. Here we investigate the principal sources of organic carbon supporting local shredder communities across a gradient in different levels of anthropogenic development along riverine reaches, in South Australia. In forested/wooded reaches with minimum to limited development, semi‐emergent macrophytes were the principal sources of organic carbon supporting the local shredder communities. However, in developed reaches, course particulate organic matter and filamentous algae were the principal food sources. The C:N ratios of the food sources in developed reaches were higher than those of their consumers indicating a stoichiometric mismatch. This imbalanced consumer‐resource nutrient ratio in those developed reaches is likely to impose constraints on the growth and reproduction of their aquatic shredder communities with probable knock‐on effects to higher trophic levels.

Species richness–decomposition relationships depend on species dominance

AbstractHuman disturbances both decrease the number of species in ecosystems and change their relative abundances. Here we present field evidence demonstrating that shifts in species abundances can have effects on ecosystem functioning that are as great as those from shifts in species richness. We investigated spatial and temporal variability of leaf decomposition rates and community metrics of leaf‐eating invertebrates (shredders) in streams. The shredder community composition dramatically influenced the diversity–function relationship; decomposition was much higher for a given species richness at sites with high species dominance than at sites where dominance was low. Decomposition rates also markedly depended on the identity of the dominant species. Further, dominance effects on decomposition varied seasonally and the number of species required for maintaining decomposition increased with increasing evenness. These findings reveal important but less obvious aspects of the biodiversity–ecosystem functioning relationship.

The Relationship Between Reparian Vegetation and the Trophic Structure of Aquatic Macroinvertebrate Communities Within the Greater Yellowstone Area

This study investigated the effects of reduced vegetation cover on stream ecosystems within the Greater Yellowstone Area. The emphasis of the research concerned whether these effects express themselves within the aquatic macroinvertebrate community. Three general vegetation categories, each comprising three to four streams, were used to distinguish gradations of riparian vegetation: dense cover (control), moderate cover, and low cover. Within each site, five aquatic macroinvertebrate samples were collected from riffle areas with a Hess sampler. The organisms were identified to the lowest practicable taxonomic level and assigned to trophic groups on the basis of gut content analysis. Riparian and in-stream variables also were measured. Statistical analyses were performed to elicit significant differences in riparian, stream, and insect community indices among the streams and to relate the structure (i.e., density, diversity, trophic organization) of the insect assemblage to the riparian and stream variables. The analysis of the data yielded no statistically significant differences among the vegetation categories with respect to embeddedness, bank overhang, and several physicochemical indices of water quality. Similarly, macroinvertebrate densities, biomass, and species richness did not differ among the categories. The trophic group analysis, however, revealed that the abundance and biomass indices of the scraper community were significantly greater within the low cover sites than within the other streams. Conversely, the indices of the shredder community were significantly higher within the control streams than within the other sites. It is proposed that a shift from allochthonous to autochthonous food resources, corresponding to the different vegetation gradients, accounted for these observations. Of critical importance is the fact that the nominally shredder genera were not excluded from or even reduced in abundance within the low cover streams, but rather seemed to have adapted to the decline in allochthonous matter by shifting their mode of food acquisition from shredding to gathering. The implications of such trophic generalism for understanding the structure and function of aquatic macroinvertebrate communities are discussed.

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

SUMMARY1. We compare the rates and mechanisms of processing of tussock (Chionochloa spp.) leaf litter in six New Zealand streams draining grassland catchments that contrast in the extent to which they have been developed for pasture.2. Rates of processing, measured as rate of weight loss of leaf packs and rate of leaf softening, were at the slow end of the spectrum for vascular plant processing. Processing was faster at developed sites, mediated mainly through the influence of oxidized nitrogen concentration on microbial activity.3. Few invertebrate shredders colonized leaf packs and it is unlikely that invertebrates had an appreciable effect on leaf processing in our study streams, which do not effectively retain leaf litter. Very small headwater tributaries appear to retain leaf litter and possess a more abundant shredder community.4. Measures of leaf processing in our six streams were significantly correlated with Petersen's (1992) RCE score of stream condition. We discuss the potential for using rate of leaf litter processing as a method of bioassessment.5. Even the most degraded stream in our study is classed as ‘good’ using the RCE inventory system. Human impact in the Taieri River is relatively small compared with the degradation observed in some parts of the world.

Secondary production and trophic relationships in a spring invertebrate community

The invertebrate fauna of a shaded area of the Danish spring Rold Kilde was sampled for a year. There was a large input of allochthonous beech litter, and the water trickled between waterlogged beech leaves. Discharge was constant and temperature amplitudes were small. Primary consumer biomass and production were 3.24 g AFDM m−2 and 8.65 g AFDM m−2 yr−1 and were dominated by shredders (70 and 69%) and gathering collectors (29 and 30%). The seasonal variation in shredder biomass and production was considerably larger within each shredder species than within the total shredder population, implying a relatively uniform processing of energy during the year. The shredder community processed about half of the total leaf input. The primary consumer P : B̄ was as low as 2.67 yr−1 due to many species with long life cycles. Average P : B̄ for eight univoltine species and five semivoltine species was as low as 3.6 and 2.3 yr−1, mainly due to low rates of elimination. Invertebrate biomass and production in low‐order streams are compared, and it is concluded that streams with a shading canopy have a significantly lower mean biomass, production, and P : B̄ ratio than streams of similar order with reduced or no canopy.