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Forest clearance is a pervasive disturbance worldwide, but many of its impacts are regarded as transient, diminishing in intensity as forest recovers. However, forests can take decades to centuries to recover after severe disturbances, and temporal lags in recovery of ecosystem properties for different forest habitats are mostly unknown. This includes forest streams, where most studies of the impacts of forest clearance are restricted to the first years of recovery, typically finding that temporary increases in light and nutrient run-off diminish as forest recovers. Implications of longer term changes remain little investigated.

In a space-for-time substitution experiment, we assessed changes in organic matter processing and in the functional and taxonomic composition of litter-consuming detritivores along a riparian forest age gradient ranging from 1 to 120 years since last timber harvesting.

Variation in organic matter processing and detritivore functional diversity along the forest succession gradient were both expressed as second-order polynomial relationships (peaking at ~50 years along the forest age gradient). Decomposition rates were lowest in both the more recently clear-cut and older riparian forest streams.

Variation of litter decomposition rates among litter bags within streams, measured by the coefficient of variation, was lowest in recent clear-cuts and increased linearly along the succession gradient. This result indicates higher within-stream heterogeneity in decomposition rates in older forest streams.

Synthesis and applications: We found that the decomposition of leaf litter, a component of carbon cycling in forests, was higher in streams flowing through intermediately aged forest, and that several key attributes of the organisms regulating litter decomposition also varied systematically with forest age. These findings highlight the longer term consequences of forest succession following forest clear-cutting for stream habitats. Our findings further illustrate complications arising from the use of forested sites as references for newly cleared sites without properly accounting for forest age, given conclusions regarding biotic responses will depend on the age of the reference forests. Finally, our results emphasise the potential of intensive forest management centred on vast, one-time clear-cutting events to drive long-term homogenisation not only in forest age structure but also in the functioning of associated forest stream habitats. ​.

1. Habitat restoration is increasingly undertaken in degraded streams and rivers to help improve biodiversity and ecosystem functioning. Follow-up assessments focused on outcomes for biodiversity have often found scant evidence for recovery, raising concerns about the efficacy of habitat restoration for improving ecological integrity. However, responses of other ecological variables, such as ecosystem process rates and the functional trait composition of biological assemblages, have been little evaluated.

2. We assessed how the restoration of habitat heterogeneity affected multiple functional parameters in 20 boreal stream reaches encompassing both more and less extensively restored sites, as well as channelised and natural reference sites. We further assessed relationships between our functional parameters and a fluvial geomorphic measure of habitat heterogeneity.

3. Leaf decomposition was positively related to habitat heterogeneity. This was associated with shifts in the functional composition of detritivore assemblages, with the most obligate litter consumers more prominent in reaches showing higher habitat heterogeneity. The deposition of fine particulate organic matter was consistently higher in restored than channelised sites, and was positively related to the heterogeneity gradient. Algal biomass accrual per unit area did not vary either with restoration or the heterogeneity gradient.

4. Synthesis and applications. Our findings demonstrate that restoration of river habitat heterogeneity can enhance retention and decomposition of organic matter, key ecosystem properties underpinning ecosystem functioning and service delivery. Significantly, enhanced litter decomposition was linked with a change in the functional composition rather than diversity of detritivore assemblages. Future evaluation of the success of habitat restorations should incorporate quantification of ecosystem processes and the functional traits of biota, in addition to measures of fluvial geomorphology and more traditional biotic metrics, to facilitate a more comprehensive and mechanistic assessment of ecological responses.

Plant litter represents a major basal resource in streams, where its decomposition is partly regulated by litter traits. Litter-trait variation may determine the latitudinal gradient in decomposition in streams, which is mainly microbial in the tropics and detritivore-mediated at high latitudes. However, this hypothesis remains untested, as we lack information on large-scale trait variation for riparian litter. Variation cannot easily be inferred from existing leaf-trait databases, since nutrient resorption can cause traits of litter and green leaves to diverge. Here we present the first global-scale assessment of riparian litter quality by determining latitudinal variation (spanning 107 degrees) in litter traits (nutrient concentrations; physical and chemical defences) of 151 species from 24 regions and their relationships with environmental factors and phylogeny. We hypothesized that litter quality would increase with latitude (despite variation within regions) and traits would be correlated to produce 'syndromes' resulting from phylogeny and environmental variation. We found lower litter quality and higher nitrogen: phosphorus ratios in the tropics. Traits were linked but showed no phylogenetic signal, suggesting that syndromes were environmentally determined. Poorer litter quality and greater phosphorus limitation towards the equator may restrict detritivore-mediated decomposition, contributing to the predominance of microbial decomposers in tropical streams.

How are resource consumption and growth rates of litter-consuming detritivores affected by imbalances between consumer and litter C:N:P ratios? To address this question, we offered leaf litter as food to three aquatic detritivore species, which represent a gradient of increasing body N: P ratios: a crustacean, a caddisfly and a stonefly. The detritivores were placed in microcosms and submerged in a natural stream. Four contrasting leaf species were offered, both singly and in two-species mixtures, to obtain different levels of stoichiometric imbalance between the resources and their consumers. The results suggest that detritivore growth was constrained by N rather than C or P, even though 1) the N: P ratios of the consumers' body tissue was relatively low and 2) microbial leaf conditioning during the experiment reduced the N:P imbalance between detritivores and leaf litter. This surprisingly consistent N limitation may be a consequence of cumulative N-demand arising from the production of N-rich chitin in the exoskeletons of all three consumer species, which is lost during regular moults, in addition to N-demand for silk production by the caddisfly. These N requirements are not commonly quantified in stoichiometric analyses of arthropod consumers. There was no evidence for compensatory feeding, but when offered mixed-species litter varying in C:N:P ratios, detritivores consumed more of the litter species showing the highest N:P and lowest C:N ratio, accelerating the mass loss of the preferred leaf species in the litter mixture. These results show that imbalances in consumer-resource stoichiometry can have contrasting effects on coupled processes, highlighting a challenge in developing a mechanistic understanding of the role of stoichiometry in regulating ecosystem processes such as leaf litter decomposition.

Biodiversity and ecosystem-functioning theory suggest that litter mixtures composed of dissimilar leaf species can enhance decomposition due to species trait complementarity. Here we created a continuous gradient of litter chemistry trait variability within species mixtures to assess effects of litter dissimilarity on three related processes in a natural stream: litter decomposition, fungal biomass accrual in the litter, and nitrogen and phosphorus immobilization. Litter from a pool of eight leaf species was analyzed for chemistry traits affecting decomposition (lignin, nitrogen, and phosphorus) and assembled in all of the 28 possible two-species combinations. Litter dissimilarity was characterized in terms of a range of trait-diversity measures, using Euclidean and Gower distances and dendrogram-based indices. We found large differences in decomposition rates among leaf species, but no significant relationships between decomposition rate of individual leaf species and litter trait dissimilarity, irrespective of whether decomposition was mediated by microbes alone or by both microbes and litter-consuming invertebrates. Likewise, no effects of trait dissimilarity emerged on either fungal biomass accrual or changes during decomposition of nitrogen or phosphorus concentrations in individual leaf species. In line with recent meta-analyses, these results provide support for the contention that litter diversity effects on decomposition, at least in streams, are less pronounced than effects on terrestrial primary productivity.

Species functional traits provide an important conceptual link between the effects of disturbances on community composition and diversity, and their ultimate outcomes for ecosystem functioning. Across 10 boreal streams covering a gradient of increasing intensity of land-use management, from forested to agricultural sites, we analysed relationships between leaf decomposition, the feeding traits of detritivores and measures of anthropogenic disturbances in two seasons. The direct effect of increasing land-use intensity on decomposition was positive and was associated with increases in nutrient concentrations and current velocities. However, this relationship was countered by negative effects associated with a loss of detritivore functional diversity along the gradient during autumn and shifts in species trait composition during spring, limiting the net change in functioning associated with increasing land-use management overall. Our results highlight the key roles that trait identity and diversity can play in mediating the effects of human disturbance on ecosystem functioning.

Despite ample experimental evidence indicating that biodiversity might be an important driver of ecosystem processes, its role in the functioning of real ecosystems remains unclear. In particular, the understanding of which aspects of biodiversity are most important for ecosystem functioning, their importance relative to other biotic and abiotic drivers, and the circumstances under which biodiversity is most likely to influence functioning in nature, is limited. We conducted a field study that focussed on a guild of insect detritivores in streams, in which we quantified variation in the process of leaf decomposition across two habitats (riffles and pools) and two seasons (autumn and spring). The study was conducted in six streams, and the same locations were sampled in the two seasons. With the aid of structural equations modelling, we assessed spatiotemporal variation in the roles of three key biotic drivers in this process: functional diversity, quantified based on a spe- cies trait matrix, consumer density and biomass. Our models also accounted for variability related to different litter resources, and other sources of biotic and abiotic variability among streams. All three of our focal biotic drivers influenced leaf decomposition, but none was important in all habitats and seasons. Functional diversity had contrasting effects on decomposition between habitats and seasons. A positive relationship was observed in pool habitats in spring, associated with high trait dispersion, whereas a negative relationship was observed in riffle habitats during autumn. Our results demonstrate that functional biodiversity can be as significant for functioning in natural ecosystems as other important biotic drivers. In particular, variation in the role of functional diversity between seasons highlights the importance of fluctuations in the relative abundances of traits for ecosystem process rates in real ecosystems.

What will happen to ecosystems if species continue to go extinct at the high rates seen today? Although ecosystems are often threatened by a myriad of physical or chemical stressors, recent evidence has suggested that the loss of species may have impacts on the functions and services of ecosystems that equal or exceed other major environmental disturbances. The underlying causes that link species diversity to ecosystem functioning include species niche complementarity, facilitative interactions, or selection effects, which cause process rates to be enhanced in more diverse communities. Interference competition, antagonistic interactions, or negative selection effects may otherwise reduce the efficiency or resource processing in diverse communities. While several of these mechanisms have been investigated in controlled experiments, there is an urgent need to understand how species diversity affects ecosystem functioning in nature, where variability of both biotic and abiotic factors is usually high. Species functional traits provide an important conceptual link between the effects of disturbances on community composition and diversity, and their ultimate outcomes for ecosystem functioning. Within this framework, I investigated relationships between the decomposition of leaf litter, a fundamental ecosystem process in stream ecosystems, and the composition and diversity of functional traits within the detritivore feeding guild. These include traits related to species habitat and resource preferences, phenology, and size. I focused on disentangling the biotic and abiotic drivers, including functional diversity, regulating ecosystem functioning in streams in a series of field experiments that captured real-world environmental gradients. Leaf decomposition rates were assessed using litter-bags of 0.5 and 10 mm opening size which allow the quantification of microbial and invertebrate + microbial contributions, respectively, to litter decomposition. I also used PVC chambers where leaf litter and a fixed number of invertebrate detritivores were enclosed in the field for a set time-period. The chemical characterisation of stream detritivores and leaf litter, by means of their nitrogen, phosphorus, and carbon concentration, was used to investigate how stoichiometric imbalance between detritivores and leaf litter may affect consumer growth and resource consumption. I found that the diversity and composition of functional traits within the stream detritivore feeding guild sometimes had effects on ecosystem functioning as strong as those of other major biotic factors (e.g. detritivore density and biomass), and abiotic factors (e.g. habitat complexity and agricultural stressors). However, the occurrence of diversity-functioning relationships was patchy in space and time, highlighting ongoing challenges in predicting the role of diversity a priori. The stoichiometric imbalance between consumers and resource was also identified as an important driver of functioning, affecting consumer growth rates, but not leaf decomposition rates. Overall, these results shed light on the understanding of species functional diversity effect on ecosystems, and indicate that the shifts in the functional diversity and composition of consumer guilds can have important outcomes for the functioning of stream ecosystems.

Stream restoration is a multi-million dollar business that aims at rehabilitating systems impacted by hydrogeomorphological modifications, such as channelization, and ameliorating physical or ecological degradation caused by catchment-scale impacts, such as agriculture or urbanization. Despite extensive programs aimed at restoring habitat complexity in channelized streams, there is little evidence for a recovery of biological diversity, and functional responses have been little assessed. Notably large-scale habitat restorations have recently been undertaken in a river catchment in northern Sweden, including rehabilitation of large habitat structures (massive boulders, large woody debris) originally removed to facilitate timber floating. Based on a hydrogeomorphological measure of habitat complexity, we characterised variability in habitat complexity across 20 stream reaches in the catchment, including reference, channelised and restored sites. We assessed whether increased habitat complexity following restoration affected retention of organic matter (FPOM), the functional diversity and organisation of the detritivore feeding guild, and two ecosystem processes: algal productivity and litter decomposition. Deposition of FPOM increased along the complexity gradient, as did leaf litter decomposition mediated by invertebrates. The increase in invertebrate-mediated decomposition was associated with shifts in the functional composition of detritivore assemblages, with feeding traits associated with more efficient decomposition more prominent in the restored reaches. There was no change in algal productivity at local scales, but increases in shallow, well- lit habitats favourable for algal growth indicate a possible increase in algal productivity at the stream reach scale. Increases in habitat complexity enhanced functioning within the detritital foodweb at local scales, without any changes in the biodiversity of detritivores. Our findings indicate that aspects of functional diversity and ecosystem functioning may be better than measures of community structure for assessing stream restoration projects.

Biodiversity and ecosystem functioning theory suggests that litter mixtures composed of dissimilar leaf species can enhance decomposition due to species trait complementarity. Here we create a continuous gradient of litter chemistry trait variability within species mixtures to assess effects of litter dissimilarity on three related processes in a natural stream: litter decomposition, fungal biomass accrual in the litter, and nitrogen and phosphorus immobilization. Litter from a pool of eight leaf species was analyzed for chemistry traits affecting decomposition (lignin, nitrogen, and phosphorus) and assembled in all of the 28 possible two-species combinations. Litter dissimilarity was characterized in terms of a range of functional diversity measures, using Euclidean and Gower distances and dendrogram-based indices. Despite favorable experimental conditions for litter dissimilarity effects to arise, we found no significant relationships between decomposition rate of individual leaf species and litter functional dissimilarity, irrespective of whether decomposition was mediated by microbes alone or by both microbes and litter-consuming invertebrates. Likewise, no effects of functional dissimilarity emerged on either fungal biomass accrual or changes during decomposition of nitrogen or phosphorus concentrations in individual leaf species. These results provide support for the contention that litter diversity effects on decomposition in streams are less pronounced than in forest environments, and in particular are less common than effects on terrestrial primary productivity. Key to understanding these discrepancies may be the constant supply of nutrients provided by the flowing water in streams, exacerbated by very large-scale cultural nutrient enrichment of surface waters in many parts of the world.