Effects of consumers and nitrogen availability on heterotrophic microbial activity during leaf decomposition in headwater streams

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Resource–consumer diversity: testing the effects of leaf litter species diversity on stream macroinvertebrate communities

1. Understanding relationships between resource and consumer diversity is essential to predicting how changes in resource diversity might affect several trophic levels and overall ecosystem functioning. 2. We tested for the effects of leaf litter species diversity (i.e. litter mixing) on litter mass remaining and macroinvertebrate communities (taxon diversity, abundance and biomass) during breakdown in a detritus‐based headwater stream (North Carolina, U.S.A.). We used full‐factorial analyses of single‐ and mixed‐species litter from dominant riparian tree species with distinct leaf chemistries [red maple (Acer rubrum), tulip poplar (Liriodendron tulipifera), chestnut oak (Quercus prinus) and rhododendron (Rhododendron maximum)] to test for additivity (single‐species litter presence/absence effects) and non‐additivity (emergent effects of litter species interactions). 3. Significant non‐additive effects of litter mixing on litter mass remaining were explained by species composition, but not richness, and litter‐mixing effects were variable throughout breakdown. Specifically, small differences in observed versus expected litter mass remaining were measured on day 14; whereas observed litter mass remaining in mixed‐species leaf packs was significantly higher on day 70 and lower on day 118 than expected from data for single‐species leaf packs. 4. Litter mixing had non‐additive effects on macroinvertebrate community structure. The number of species in litter mixtures (two to four), but not litter species composition, was a significant predictor of the dominance of particular macroinvertebrates (i.e. indicator taxa) within mixed‐species packs. 5. In addition, the presence/absence of high‐ (L. tulipifera) and low‐quality (R. maximum) litter had additive effects on macroinvertebrate taxon richness, abundance and biomass. The presence of L. tulipifera litter had both positive (synergistic) and negative (antagonistic) effects on invertebrate taxon richness, that varied during breakdown but were not related to litter chemistry. In contrast, the presence/absence of L. tulipifera had a negative relationship with total macroinvertebrate biomass (due to low leaf mass remaining when L. tulipifera was present and higher condensed and hydrolysable tannins associated with leaf packs lacking L. tulipifera). Macroinvertebrate abundance was consistently lower when R. maximum was present, which was partially explained by litter chemistry [e.g., high concentrations of lignin, condensed tannins, hydrolysable tannins and total phenolics and high carbon to nutrient (N and P) ratios]. 6. The bottom‐up effects of litter species diversity on stream macroinvertebrates and litter breakdown are different, which suggests that structural attributes of macroinvertebrate communities may only partially explain the effects of litter‐mixing on organic matter processing in streams. In addition, stream macroinvertebrates colonising decomposing litter are influenced by resource diversity as well as resource availability. Broad‐scale shifts in riparian tree species composition will alter litter inputs to streams, and our results suggest that changes in the diversity and availability of terrestrial litter may alter stream food webs and organic matter processing.     

Nutrients and temperature additively increase stream microbial respiration

Rising temperatures and nutrient enrichment are co‐occurring global‐change drivers that stimulate microbial respiration of detrital carbon, but nutrient effects on the temperature dependence of respiration in aquatic ecosystems remain uncertain. We measured respiration rates associated with leaf litter, wood, and fine benthic organic matter (FBOM) across seasonal temperature gradients before (PRE) and after (ENR1, ENR2) experimental nutrient (nitrogen [N] and phosphorus [P]) additions to five forest streams. Nitrogen and phosphorus were added at different N:P ratios using increasing concentrations of N (~80–650 μg/L) and corresponding decreasing concentrations of P (~90–11 μg/L). We assessed the temperature dependence, and microbial (i.e., fungal) drivers of detrital mass‐specific respiration rates using the metabolic theory of ecology, before vs. after nutrient enrichment, and across N and P concentrations. Detrital mass‐specific respiration rates increased with temperature, exhibiting comparable activation energies (E, electronvolts [eV]) for all substrates (FBOM E = 0.43 [95% CI = 0.18–0.69] eV, leaf litter E = 0.30 [95% CI = 0.072–0.54] eV, wood E = 0.41 [95% CI = 0.18–0.64] eV) close to predicted MTE values. There was evidence that temperature‐driven increased respiration occurred via increased fungal biomass (wood) or increased fungal biomass‐specific respiration (leaf litter). Respiration rates increased under nutrient‐enriched conditions on leaves (1.32×) and wood (1.38×), but not FBOM. Respiration rates responded weakly to gradients in N or P concentrations, except for positive effects of P on wood respiration. The temperature dependence of respiration was comparable among years and across N or P concentration for all substrates. Responses of leaf litter and wood respiration to temperature and the combined effects of N and P were similar in magnitude. Our data suggest that the temperature dependence of stream microbial respiration is unchanged by nutrient enrichment, and that increased temperature and N + P availability have additive and comparable effects on microbial respiration rates.     

Effects of Nutrient Enrichment on Decomposition and Fungal Colonization of Sweet Chestnut Leaves in an Iberian Stream (Central Portugal)

This study assessed the effect of nutrient enrichment on rates of decomposition, ergosterol concentrations (as a measure of fungal biomass), and rates of fungal sporulation of sweet chestnut (Castanea sativa Miller) leaves in a 3rd order stream (Central Portugal), with medium to high background values of nutrients. Coarse and fine mesh leaf bags were attached to nutrient diffusing substrata containing NaNO₃, KH₂PO₄, both nutrients, or no additions. Leaf breakdown rates were similar in the four treatments and in the two mesh sizes (k=−0.0155 to −0.0219 day⁻¹). Phosphorus content of P or N + P enriched leaves was higher than in the other treatments after 28 days, but there were no differences in N concentrations. Ergosterol concentrations associated with decomposing leaves were similar among treatments. The peak sporulation rates of aquatic hyphomycetes were stimulated by the addition of N + P and N but not by P alone. Results from the experiment provide evidence that leaf breakdown in the study stream, as a model for streams with naturally medium to high level of nutrients, was not nutrient-limited, and that fungal reproductive activity was limited by dissolved N but not by dissolved P in stream water.     

Effects of nutrient enrichment and leaf quality on the breakdown of leaves in a hardwater stream

1. The breakdown of leaf litter in streams is influenced strongly by leaf quality and the concentration of dissolved nutrients, primarily inorganic nitrogen (N) and phosphorus (P) in the water. We examined the effect of nutrient enrichment on the breakdown of three species of leaves in a hardwater, nutrient‐rich stream. The rate of microbial respiration was also measured on the decomposing leaves. 2. The breakdown rates of dogwood (Cornus stolonifera), aspen (Populus tremuloides) and birch (Betula occidentalis), k‐values of 0.0461, 0.0307 and 0.0186 day^–1, respectively, were unaffected by nutrient enrichment and generally faster than reported previously. Microbial respiration on the leaves was greater than reported previously for leaves of congeneric species. It appears that leaf breakdown in the study stream was not nutrient limited. 3. Nitrogen‐based measures of leaf quality, such as percentage N and carbon (C)/nitrogen ratio, did not correspond to measured breakdown rates among the three leaf types. The best predictors of relative breakdown rates were percentage lignin and the percentage of the total carbon that occurred as lignin. We suggest that, when leaf breakdown is not nutrient limited, measures of carbon quality (i.e. lignin‐based measures) are a better assessment of overall leaf quality than are N‐based measures. 4. Previous studies have indicated that the enzymes produced by aquatic hyphomycetes (microfungi) operate most efficiently at a basic pH and in the presence of calcium ions. The hardwater conditions (pH=8.6, total hardness > 300 mg CaCO₃ L^–1) and abundance of dissolved NO₃ and soluble reactive phosphorous (SRP) (approximately 50 μg L^–1, each) in the study stream appear to have provided conditions that resulted in a high respiration rate and rapid breakdown of leaf litter.     

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.     

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.     

Shredder abundance and leaf breakdown in an Appalachian Mountain stream

SUMMARY.

• 1 Breakdown rates of dogwood (Cornus florida L.), red maple (Acer rubrum L.) and white oak (Quercus alba L.) leaves were investigated at two first-order and two second-order sites in an Appalachian Mountain stream.
• 2 Leaves exposed in mesh bags were sampled on eight occasions over a 207 day period and breakdown rates were compared using an exponential decay model.
• 3 There was a consistent ranking in leaf breakdown rate within each site, i.e. dogwood > red maple > white oak, and all species broke down faster at second-than at first-order sites.
• 4 Our data suggest that differences in species-specific leaf breakdown rates were largely a function of shredder abundance on the leaves.

     

Leaf litter decomposition and microbial activity in nutrient-enriched and unaltered reaches of a headwater stream

SUMMARY 1. Decomposition of red maple ( Acer rubrum ) and rhododendron ( Rhododendron maximum ) leaves and activity of associated microorganisms were compared in two reaches of a headwater stream in Coweeta Hydrologic Laboratory, NC, U.S.A. The downstream reach was enriched with ammonium, nitrate, and phosphate whereas the upstream reach was not altered.
2. Decomposition rate, microbial respiration, fungal and bacterial biomass, and the sporulation rate of aquatic hyphomycetes associated with decomposing leaf material were significantly higher for both leaf types in the nutrient-enriched reach. Species richness and community structure of aquatic hyphomycetes also exhibited considerable changes with an increase in the number of fungal codominants in the nutrient-enriched reach.
3. Fungal biomass was one to two orders of magnitude greater than bacterial biomass in both reaches. Changes in microbial respiration rate corresponded to those in fungal biomass and sporulation, suggesting a primary role of fungi in leaf decomposition.
4. Nutrient enrichment increased microbial activity, the proportion of leaf carbon channelled through the microbial compartment and the decomposition rate of leaf litter.     

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

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

     

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.     

Effects of organic matter and season on leaf litter colonisation and breakdown in cave streams

1. Low organic matter availability is thought to be a primary factor influencing evolutionary and ecological processes in cave ecosystems. We examined links among organic matter abundance, macroinvertebrate community structure and breakdown rates of red maple (Acer rubrum) and corn litter (Zea mays) in coarse‐ (10 × 8 mm) and fine‐mesh (500‐μm) litter bags over two seasonal periods in four cave streams in the south‐eastern U.S.A. 2. Organic matter abundance differed among cave streams, averaging from near zero to 850 g ash‐free dry mass m^?2. Each cave system harboured a different macroinvertebrate community. However, trophic structure was similar among caves, with low shredder biomass (2–17% of total biomass). 3. Corn litter breakdown rates (mean k = 0.005 day^?1) were faster than red maple (mean k = 0.003 day^?1). Breakdown rates in coarse‐mesh bags (k = 0.001–0.012 day^?1) were up to three times faster than in fine‐mesh bags (k = 0.001–0.004 day^?1). Neither invertebrate biomass in litter bags nor breakdown rates were correlated with the ambient abundance of organic matter. Litter breakdown rates showed no significant temporal variation. Epigean (surface‐adapted) invertebrates dominated biomass in litter bags, suggesting that their effects on cave ecosystem processes may be greater than hypogean (cave‐adapted) taxa, the traditional focus of cave studies. 4. The functional diversity of our cave communities and litter breakdown rates are comparable to those found in previous litter breakdown studies in cave streams, suggesting that the factors that control organic matter processing (e.g. trophic structure of communities) may be broadly similar across geographically diverse areas.     

Resource quality and stoichiometric constraints on stream ecosystem functioning

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

Decomposition of Fire Exposed Eucalyptus Leaves in a Portuguese Lowland Stream

We compared fire exposed with normal abscised eucalyptus leaves incubated in a stream running through eucalyptus plantations in central Portugal, in terms of breakdown rates, microbial activity, diversity and macroinvertebrate abundance. Although leaves exposed to fire had lower nutritional value, mass loss was similar for both leaf types (k = 0.0089–0.0095 d^–1 for fire and k = 0.0084–0.00103 d^–1 for normal leaves). Fungal biomass was similar among treatments, whereas sporulation and microbial respiration were lower in fire exposed leaves. Both leaf types had similar aquatic hyphomycetes communities. Physical fragmentation was important in fire exposed leaves breakdown. Invertebrates colonized leaves in low numbers in both treatments. Alteration of leaf litter quality determined by fires in streams does not seem to determine changes in ecosystem functioning in a short term. (© 2007 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

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.     

Responses of a beaded Arctic stream to short-term N and P fertilisation

1. Oligotrophic Arctic streams are likely to be sensitive to changes in hydrology and nutrient inputs predicted to occur as a consequence of future climate and land use change. To investigate the potential consequences of nutrient enrichment for low‐order Arctic streams, we added ammonium‐N and phosphorous to a second‐order beaded, tundra stream on Alaska's north slope. We measured responses in nutrient chemistry, chlorophyll a standing crop, and in the breakdown and macroinvertebrate colonisation of leaf litter over a 38‐day summer period. 2. During the addition, nutrient concentrations immediately downstream of the dripper averaged 6.4 μm ammonium‐N and 0.45 μm soluble reactive P. Concentrations upstream of the dripper averaged 0.54 μm ammonium‐N and 0.03 μm soluble reactive P. Uptake of both nutrients was rapid. Concentrations were reduced on average to 28% (ammonium‐N) and 15% (inorganic P) of maximum values within 1500 m. Standing crops of chlorophyll a on standardised samplers were significantly higher by the end of the experiment. Breakdown rates of senescent willow (Salix sp.) and sedge (Carex sp.) litter and associated fungal biomass were also significantly increased by nutrient addition. 3. Fertilisation resulted in four‐ to sevenfold higher macroinvertebrate abundance and two‐ to fourfold higher macroinvertebrate biomass in litter bags, as well as an increase in late‐summer body mass of larval Nemoura stoneflies. 4. Our results are consistent with those of similar studies of larger streams in the high‐Arctic region. Based on our short‐term experiment, increased inputs of nutrients into these ecosystems, whether caused by climate change or more local disturbance, are likely to have profound ecological consequences. Longer‐term effects of enrichment, and their interaction with other components of future change in climate or land use, are more difficult to assess.     

Leaf breakdown, detrital resources, and food webs in streams affected by mine drainage

Breakdown of leaf litter is essential for providing detrital resources for food webs but can be impaired by anthropogenic activities, which may disrupt energy flow to consumers. We investigated the relationship between leaf breakdown and food web structure in 12 streams with or without mining impacts on South Island, New Zealand. Six streams received inputs of acid mine drainage (pH 2.5–4.9), three were naturally acidic (pH ~5.0), and three were circumneutral (pH ~6.8). Streams affected by mining either had highly acidic water (pH <3) or iron precipitates present on substrata. Breakdown rates of leaves were significantly lower in mining-affected streams than circumneutral (by almost 50%) but not naturally acidic streams and were driven primarily by microbial activity, as shredding invertebrates were often absent. Mining-affected stream webs were simplified structures with fewer species and links than those in other streams. With few species to process leaf litter and transfer detrital resources, inputs of AMD disrupted both the mechanisms responsible for breakdown and links for energy flow. While faster breakdown rates were associated with larger food webs, limited function maintained in mining-affected streams was sufficient to support primary consumers and small food webs.     

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.     

Detrital nutrient content and leaf species differentially affect growth and nutritional regulation of detritivores

Resource nutrient content and identity are common bottom–up controls on organismal growth and nutritional regulation. One framework to study these factors, ecological stoichiometry theory, predicts that elevated resource nitrogen (N) and phosphorus (P) contents enhance organism growth by alleviating constraints on N and P acquisition. However, the regulatory mechanisms underlying this response – including whether responses depend on resource identity – remain poorly understood. In this study, we tested roles of detrital N and P contents and identity (leaf species) in constraining growth of aquatic invertebrate detritivores. We synthesized results from seven detritivore species fed wide nutrient gradients of oak and maple detritus in the laboratory. Across detritivore taxa, we used a meta‐analytic approach quantifying effects of detrital leaf species and N and P contents on growth, consumption, and N‐ and P‐specific assimilation and growth efficiencies. Detritivore growth rates increased on higher‐N and P detritus and on oak compared to maple detritus. Notably, the mechanisms of improved growth differed between the responses to detrital nutrients versus leaf species, with the former driven by greater consumption rates despite lower assimilation efficiencies on higher‐nutrient detritus, and the latter driven by improved N and P assimilation and N growth efficiencies on oak detritus. These findings suggest animal nutrient acquisition changes flexibly in response to resource changes, altering the fate of detrital N and P throughout regulation. We affirm resource identity and nutrients as important bottom–up controls, but suggest these factors act through separate pathways to affect organism growth and thereby change detrital ecosystems under anthropogenic forest compositional change and nutrient enrichment.     

Effects of nutrient enrichment on boreal streams: invertebrates, fungi and leaf-litter breakdown

1. The effect of nutrient enrichment on structural (invertebrate indices) and functional (leaf‐litter breakdown rates) characteristics of stream integrity was studied in nine boreal streams. 2. The results showed predicted changes in biotic indices and leaf‐litter breakdown along a complex (principal component) nutrient gradient. Biotic indices were better correlated with nutrient effects than leaf‐litter breakdown. 3. Fungal biomass and invertebrate densities in the litter bags were positively correlated with leaf‐litter breakdown, and both were also positively related to the nutrient gradient. 4. Invertebrate community composition influenced breakdown rate. High breakdown rates at one site were associated with the high abundance of the detritivore Asellus aquaticus. 5. This study lends support to the importance of invertebrate and fungi as mediators of leaf‐litter decomposition. However, our study also shows that study design (length of incubation) can confound the interpretation of nutrient‐induced effects on decomposition.