Nonadditive effects of leaf litter species diversity on breakdown dynamics in a detritus-based stream

Since species loss is predicted to be nonrandom, it is important to understand the manner in which those species that we anticipate losing interact with other species to affect ecosystem function. We tested whether litter species diversity, measured as richness and composition, affects breakdown dynamics in a detritus-based stream. Using full-factorial analyses of single- and mixed-species leaf packs (15 possible combinations of four dominant litter species; red maple [Acer rubrum], tulip poplar [Liriodendron tulipifera], chestnut oak [Quercus prinus], and rhododendron [Rhododendron maximum]), we tested for single-species presence/absence (additive) or species interaction (nonadditive) effects on leaf pack breakdown rates, changes in litter chemistry, and microbial and macroinvertebrate biomass. Overall, we found significant nonadditive effects of litter species diversity on leaf pack breakdown rates, which were explained both by richness and composition. Leaf packs containing higher litter species richness had faster breakdown rates, and antagonistic effects of litter species composition were observed when any two or three of the four litter species were mixed. Less-consistent results were obtained with respect to changes in litter chemistry and microbial and macroinvertebrate biomass. Our results suggest that loss of litter species diversity will decrease species interactions involved in regulating ecosystem function. To that end, loss of species such as eastern hemlock (Tsuga canadensis) accompanied by predicted changes in riparian tree species composition in the southeastern United States could have nonadditive effects on litter breakdown at the landscape scale.     

Mitigating the impact of oil‐palm monoculture on freshwater fishes in Southeast Asia

Anthropogenic land‐cover change is driving biodiversity loss worldwide. At the epicenter of this crisis lies Southeast Asia, where biodiversity‐rich forests are being converted to oil‐palm monocultures. As demand for palm oil increases, there is an urgent need to find strategies that maintain biodiversity in plantations. Previous studies found that retaining forest patches within plantations benefited some terrestrial taxa but not others. However, no study has focused on aquatic taxa such as fishes, despite their importance to human well‐being. We assessed the efficacy of forested riparian reserves in conserving freshwater fish biodiversity in oil‐palm monoculture by sampling stream fish communities in an oil‐palm plantation in Central Kalimantan, Indonesia. Forested riparian reserves maintained preconversion local fish species richness and functional diversity. In contrast, local and total species richness, biomass, and functional diversity declined markedly in streams without riparian reserves. Mechanistically, riparian reserves appeared to increase local species richness by increasing leaf litter cover and maintaining coarse substrate. The loss of fishes specializing in leaf litter and coarse substrate decreased functional diversity and altered community composition in oil‐palm plantation streams that lacked riparian reserves. Thus, a land‐sharing strategy that incorporates the retention of forested riparian reserves may maintain the ecological integrity of fish communities in oil‐palm plantations. We urge policy makers and growers to make retention of riparian reserves in oil‐palm plantations standard practice, and we encourage palm‐oil purchasers to source only palm oil from plantations that employ this practice.     

Headwater riparian forest-floor invertebrate communities associated with alternative forest management practices.

Headwater streams and their riparian zones are a common, yet poorly understood, component of Pacific Northwest, USA, landscapes. We describe the ecological importance of headwater stream riparian zones as habitat for forest-floor invertebrate communities and assess how alternative management strategies for riparian zones may impact these communities. We compared community composition of forest-floor invertebrates at increasing distances along trans-riparian (stream edge to upslope) transects in mature forests, clearcuts, and riparian buffers of approximately 30-m width with upslope clearcuts. Invertebrates were collected using pitfall traps in five replicate blocks of three treatments each in the Willamette National Forest, Oregon, USA. We measured microclimate and microhabitat variables at pitfall locations. Despite strong elevation and block effects on community composition, community analyses revealed a distinct "riparian" invertebrate community within 1 m of the stream edge in mature forest treatments, which was strongly related to cool, humid microclimate conditions. Invertebrate community composition in buffer treatments was far more similar to that of mature forests than to clearcuts; a pattern mirrored by microclimate. These results suggest that, within our study sites, forest-floor invertebrate distributions are strongly associated with microclimate and that riparian buffers of approximately 30-m width do provide habitat for many riparian and forest species. Riparian reserves may serve as effective forest refugia and/or dispersal corridors for invertebrates and other taxa, and their incorporation into watershed management plans likely will contribute to meeting persistence and connectivity objectives.     

Urbanization affects stream ecosystem function by altering hydrology, chemistry, and biotic richness.

Catchment urbanization can alter physical, chemical, and biological attributes of stream ecosystems. In particular, changes in land use may affect the dynamics of organic matter decomposition, a measure of ecosystem function. We examined leaf-litter decomposition in 18 tributaries of the St. Johns River, Florida, USA. Land use in all 18 catchments ranged from 0% to 93% urban which translated to 0% to 66% total impervious area (TIA). Using a litter-bag technique, we measured mass loss, fungal biomass, and macroinvertebrate biomass for two leaf species (red maple [Acer rubrum] and sweetgum [Liquidambar styraciflua]). Rates of litter mass loss, which ranged from 0.01 to 0.05 per day for red maple and 0.006 to 0.018 per day for sweetgum, increased with impervious catchment area to levels of approximately 30-40% TIA and then decreased as impervious catchment area exceeded 40% TIA. Fungal biomass was also highest in streams draining catchments with intermediate levels of TIA. Macroinvertebrate biomass ranged from 17 to 354 mg/bag for red maple and from 15 to 399 mg/bag for sweetgum. Snail biomass and snail and total invertebrate richness were strongly related to breakdown rates among streams regardless of leaf species. Land-use and physical, chemical, and biological variables were highly intercorrelated. Principal-components analysis was therefore used to reduce the variables into several orthogonal axes. Using stepwise regression, we found that flow regime, snail biomass, snail and total invertebrate richness, and metal and nutrient content (which varied in a nonlinear manner with impervious surface area) were likely factors affecting litter breakdown rates in these streams.     

Decomposition of diverse litter mixtures in streams

In view of growing interest in understanding how biodiversity affects ecosystem functioning, we investigated effects of riparian plant diversity on litter decomposition in forest streams. Leaf litter from 10 deciduous tree species was collected during natural leaf fall at two locations (Massif Central in France and Carpathians in Romania) and exposed in the field in litter bags. There were 35 species combinations, with species richness ranging 1-10. Nonadditive effects on the decomposition of mixed-species litter were minor, although a small synergistic effect was observed in the Massif Central stream where observed litter mass remaining was significantly lower overall than expected from data on single-species litter. In addition, variability in litter mass remaining decreased with litter diversity at both locations. Mean nitrogen concentration of single- and mixed-species litters (0.68-4.47% of litter ash-free dry mass) accounted for a large part of the variation in litter mass loss across species combinations. For a given species or mixture, litter mass loss was also consistently faster in the Massif Central than in the Carpathians, and the similarity in general stream characteristics, other than temperature, suggests that this effect was largely due to differences in thermal regimes. These results support the notion that decomposition of litter mixtures is primarily driven by litter quality and environmental factors, rather than by species richness per se. However, the observed consistent decrease in variability of decomposition rate with increasing plant species richness indicates that conservation of riparian tree diversity is important even when decomposition rates are not greatly influenced by litter mixing.     

Effects of riparian land use on water quality and fish communities in the headwater stream of the Taizi River in China

Riparian land use remains one of the most significant impacts on stream ecosystems. This study focuses on the relationship between stream ecosystems and riparian land use in headwater regions. Four riparian land types including forest, grassland, farmland, and residential land were examined to reveal the correlation between stream water and fish communities in headwater streams of the Taizi River in north-eastern China. Four land types along riparian of 3 km in length were evaluated at 25, 50, 100, 200 and 500 m widths, respectively. Generally, the results found a significant relationship between riparian land uses and stream water quality. Grassland was positively correlated with water quality parameters (conductivity and total dissolved solids) at scales from 100 to 500 m riparian width. Farmland and residential land was negatively correlated with water quality parameters at scales from 25 to 500 m and from 50 to 200 m riparian widths, respectively. Although the riparian forest is important for maintaining habitat diversity and fish communities, the results found that only fish communities were significantly correlated with the proportion of riparian farmland. Farmland had a positive correlation with individual fish abundance within a riparian corridor of 25 to 50 m, but a negative correlation with fish diversity metrics from 25 to 100 m. This study indicates that effective riparian management can improve water quality and fish communities in headwater streams.     

Plant genes link forests and streams

Although it is understood that the composition of riparian trees can affect stream function through leaf litter fall, the potential effects of genetic variation within species are less understood. Using a naturally hybridizing cottonwood system, we examined the hypothesis that genetic differences among two parental species (Populus fremontii and P. angustifolia) and two groups of their hybrids (F1 and backcrosses to P. angustifolia) would affect litter decomposition rates and the composition of the aquatic invertebrate community that colonizes leaves. Three major findings emerged: (1) parental and hybrid types differ in litter quality, (2) decomposition differs between two groups, a fast group (P. fremontii and F1 hybrid), and a slow group (P. angustifolia and backcross hybrids), and (3) aquatic invertebrate communities colonizing P. fremontii litter differed significantly in composition from all other cross types, even though P. fremontii and the F1 hybrid decomposed at similar rates. These findings are in agreement with terrestrial arthropod studies in the same cottonwood system. However, the effects are less pronounced aquatically than those observed in the adjacent terrestrial community, which supports a genetic diffusion hypothesis. Importantly, these findings argue that genetic interactions link terrestrial and aquatic communities and may have significant evolutionary and conservation implications.     

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.     

Effects of Drainage‐Basin Geomorphology on Insectivorous Bird Abundance in Temperate Forests

Abstract: Interfaces between terrestrial and stream ecosystems often enhance species diversity and population abundance of ecological communities beyond levels that would be expected separately from both the ecosystems. Nevertheless, no study has examined how stream configuration within a watershed influences the population of terrestrial predators at the drainage‐basin scale. We examined the habitat and abundance relationships of forest insectivorous birds in eight drainage basins in a cool temperate forest of Japan during spring and summer. Each basin has different drainage‐basin geomorphology, such as the density and frequency of stream channels. In spring, when terrestrial arthropod prey biomass is limited, insectivorous birds aggregated in habitats closer to streams, where emerging aquatic prey was abundant. Nevertheless, birds ceased to aggregate around streams in summer because terrestrial prey became plentiful. Watershed‐scale analyses showed that drainage basins with longer stream channels per unit area sustained higher densities of insectivorous birds. Moreover, such effects of streams on birds continued from spring through summer, even though birds dispersed out of riparian areas in the summer. Although our data are from only a single year, our findings imply that physical modifications of stream channels may reduce populations of forest birds; thus, they emphasize the importance of landscape‐based management approaches that consider both stream and forest ecosystems for watershed biodiversity conservation.     

Macroinvertebrate Colonization of Woody Debris in Canadian Shield Lakes following Riparian Clearcutting

Deployment of litterfall traps revealed that clearcut logging of boreal riparian forests in northwestern Ontario, Canada resulted in a dramatic shift from once dominant conifers to regrowth composed largely of deciduous trees and reduced the allochthonous inputs of small woody debris to lake littoral zones by over 90%. Due to the rarity of macrophytes in these oligotrophic lakes, littoral macroinvertebrates were found to actively colonize woody debris placed within mesh litter bags. The recalcitrant nature of small woody debris in these lakes (average median persistence time of about 5 years estimated from mass loss data) indicates, however, that this important habitat resource will probably never completely disappear in relation to its projected rate of resupply during post-disturbance forest regeneration. Colonization rates of twigs and bark contained within the litter bags were not found to differ between coniferous and deciduous species. This indicates that macroinvertebrates in these boreal lakes are merely opportunistic colonizers of woody debris, probably for its use as either a biofilm substrate or a predation refuge. As a result, shifts in tree species composition following riparian clearcutting should not detrimentally affect the taxa richness or organism abundance of aquatic macroinvertebrates in these lakes.     

Twenty years of stream restoration in Finland: little response by benthic macroinvertebrate communities

The primary focus of many in-stream restoration projects is to enhance habitat diversity for salmonid fishes, yet the lack of properly designed monitoring studies, particularly ones with pre-restoration data, limits any attempts to assess whether restoration has succeeded in improving salmonid habitat. Even less is known about the impacts of fisheries-related restoration on other, non-target biota. We examined how restoration aiming at the enhancement of juvenile brown trout (Salmo trutta L.) affects benthic macroinvertebrates, using two separate data sets: (1) a before-after-control-impact (BACI) design with three years before and three after restoration in differently restored and control reaches of six streams; and (2) a space-time substitution design including channelized, restored, and near-natural streams with an almost 20-year perspective on the recovery of invertebrate communities. In the BACI design, total macroinvertebrate density differed significantly from before to after restoration. Following restoration, densities decreased in all treatments, but less so in the controls than in restored sections. Taxonomic richness also decreased from before to after restoration, but this happened similarly in all treatments. In the long-term comparative study, macroinvertebrate species richness showed no difference between the channel types. Community composition differed significantly between the restored and natural streams, but not between restored and channelized streams. Overall, the in-stream restoration measures used increased stream habitat diversity but did not enhance benthic biodiversity. While many macroinvertebrates may be dispersal limited, our study sites should not have been too distant to reach within almost two decades. A key explanation for the weak responses by macroinvertebrate communities may have been historical. When Fennoscandian streams were channelized for log floating, the loss of habitat heterogeneity was only partial. Therefore, habitat may not have been limiting the macroinvertebrate communities to begin with. Stream restoration to support trout fisheries has strong public acceptance in Finland and will likely continue to increase in the near future. Therefore, more effort should be placed on assessing restoration success from a biodiversity perspective using multiple organism groups in both stream and riparian ecosystems.     

Geology-dependent impacts of forest conversion on stream fish diversity

Forest conversion is one of the greatest global threats to biodiversity, and land-use change and subsequent biodiversity declines sometimes occur over a variety of underlying geologies. However, how forest conversion and underlying geology interact to alter biodiversity is underappreciated, although spatial variability in geology is considered an integral part of sustaining ecosystems. We aimed to examine the effects of forest conversion to farmland, the underlying geology, and their interaction on the stream fishes’ diversity, evenness, and abundance in northeastern Japan. We disentangled complex pathways between abiotic and biotic factors with structural equation modeling. Species diversity of stream fishes was indirectly shaped by the interaction of land use and underlying geology. Diversity declined due to nutrient enrichment associated with farmlands, which was mainly the result of changes in evenness rather than by changes in species richness. This impact was strongest in streams with volcanic geology with coarse substrates probably because of the differential responses of abundant stream fishes to nutrient enrichment (i.e., dominance) and the high dependency of these fishes on large streambed materials during their life cycles. Our findings suggest that remediation of deforested or degraded forest landscapes would be more efficient if the interaction between land use and underlying geology was considered. For example, the negative impacts of farmland on evenness were larger in streams with volcanic geology than in other stream types, suggesting that riparian forest restoration along such streams would efficiently provide restoration benefits to stream fishes. Our results also suggest that land clearing around such streams should be avoided to conserve species evenness of stream fishes.     

Effects of Oil‐Palm Plantations on Diversity of Tropical Anurans

Agriculturally altered vegetation, especially oil‐palm plantations, is rapidly increasing in Southeast Asia. Low species diversity is associated with this commodity, but data on anuran diversity in oil‐palm plantations are lacking. We investigated how anuran biological diversity differs between forest and oil‐palm plantation, and whether observed differences in biological diversity of these areas is linked to specific environmental factors. We hypothesized that biological diversity is lower in plantations and that plantations support a larger proportion of disturbance‐tolerant species than forest. We compared species richness, abundance, and community composition between plantation and forest areas and between site types within plantation and forest (forest stream vs. plantation stream, forest riparian vs. plantation riparian, forest terrestrial vs. plantation terrestrial). Not all measures of biological diversity differed between oil‐palm plantations and secondary forest sites. Anuran community composition, however, differed greatly between forest and plantation, and communities of anurans in plantations contained species that prosper in disturbed areas. Although plantations supported large numbers of breeding anurans, we concluded the community consisted of common species that were of little conservation concern (commonly found species include Fejervarya limnocharis, Microhyla heymonsi, and Hylarana erythrea). We believe that with a number of management interventions, oil‐palm plantations can provide habitat for species that dwell in secondary forests. Efectos de las Plantaciones de Palma de Aceite sobre la Diversidad de Anuros Tropicales Faruk et al.     

An invasive riparian tree reduces stream ecosystem efficiency via a recalcitrant organic matter subsidy

A disturbance, such as species invasion, can alter the exchange of materials and organisms between ecosystems, with potential consequences for the function of both ecosystems. Russian olive (Elaeagnus angustifolia) is an exotic tree invading riparian corridors in the western United States, and may alter stream organic matter budgets by increasing allochthonous litter and by reducing light via shading, in turn decreasing in-stream primary production. We used a before-after invasion comparison spanning 35 years to show that Russian olive invasion increased allochthonous litter nearly 25-fold to an invaded vs. a control reach of a stream, and we found that this litter decayed more slowly than native willow. Despite a mean 50% increase in canopy cover by Russian olive and associated shading, there were no significant changes in gross primary production. Benthic organic matter storage increased fourfold after Russian olive invasion compared to pre-invasion conditions, but there were no associated changes in stream ecosystem respiration or organic matter export. Thus, estimated stream ecosystem efficiency (ratio of ecosystem respiration to organic matter input) decreased 14%. These findings show that invasions of nonnative plant species in terrestrial habitats can alter resource fluxes to streams with consequences for whole-ecosystem functions.     

The width of riparian habitats for understory birds in an Amazonian forest

Riparian habitats are important for the maintenance of regional biodiversity. Many studies have compared bird distributions between riparian and non-riparian habitats but have not established how wide riparian habitats used by birds are, as measured by distance from the nearest stream. We investigated the distribution of understory birds along gradients of distance from streams, soil clay content, and slope in a central Amazonian forest, by mist-netting birds three times in 45 plots. We used nonmetric multidimensional scaling (NMDS) to reduce the dimensionality of species quantitative (abundance) and qualitative (presence-absence) composition to one multivariate axis. Estimates of the width of riparian habitats as indicated by understory birds depended on the community attribute considered, measuring 90 m for species quantitative composition and 140 m for species qualitative composition. Species distributions were correlated with clay content but were independent of slope, while distance from streams was positively correlated with clay content but independent of slope. Clay content affects plant species composition, which in turn, may influence bird species composition. However, distribution patterns of birds in relation to distance from streams are consistent among studies carried out in many different temperate and tropical regions, indicating an effect of distance from streams itself. Protection of riparian habitats is one of the most widely used conservation strategies, and Brazilian environmental legislation mandates the protection of a 30 m wide strip of riparian vegetation on either side of small streams. We show that the protected strip should be much wider and recommend strategies to place other forms of land protection contiguous with riparian areas so that Brazilian environmental legislation better fulfills its role of protecting biodiversity associated with riparian habitats.     

Restoration of rivers used for timber floating: effects on riparian plant diversity.

Fluvial processes such as flooding and sediment deposition play a crucial role in structuring riparian plant communities. In rivers throughout the world, these processes have been altered by channelization and other anthropogenic stresses. Yet despite increasing awareness of the need to restore natural flow regimes for the preservation of riparian biodiversity, few studies have examined the effects of river restoration on riparian ecosystems. In this study, we examined the effects of restoration in the Ume River system, northern Sweden, where tributaries were channelized to facilitate timber floating in the 19th and early 20th centuries. Restoration at these sites involved the use of heavy machinery to replace instream boulders and remove floatway structures that had previously lined stream banks and cut off secondary channels. We compared riparian plant communities along channelized stream reaches with those along reaches that had been restored 3-10 years prior to observation. Species richness and evenness were significantly increased at restored sites, as were floodplain inundation frequencies. These findings demonstrate how river restoration and associated changes in fluvial disturbance regimes can enhance riparian biodiversity. Given that riparian ecosystems tend to support a disproportionate share of regional species pools, these findings have potentially broad implications for biodiversity conservation at regional or landscape scales.     

Increasing litter species richness reduces variability in a terrestrial decomposer system

Debate on the relationship between diversity and stability has been driven by the recognition that species loss may influence ecosystem properties and processes. We conducted a litterbag experiment in the Scottish Highlands, United Kingdom, to examine the effects of altering plant litter diversity on decomposition, microbial biomass, and microfaunal abundance. The design of treatments was fully factorial and included five species from an upland plant community (silver birch, Betula pendula; Scots' pine, Pinus sylvestris; heather, Calluna vulgaris; bilberry, Vaccinium myrtillus; wavy-hair grass, Deschampsia flexuosa); species richness ranged from one to five species. We tested the effects of litter species richness and composition on variable means, whether increasing litter species richness reduced variability in the decomposer system, and whether any richness-variability relationships were maintained over time (196 vs. 564 days). While litter species composition effects controlled variable means, we revealed reductions in variability with increasing litter species richness, even after accounting for differences between litter types. These findings suggest that higher plant species richness per se may result in more stable ecosystem processes (e.g., decomposition) and decomposer communities. Negative richness-variation relationships generally relaxed over time, presumably because properties of litter mixtures became more homogeneous. However, given that plant litter inputs continue to enter the belowground system over time, we conclude that variation in ecosystem properties may be buffered by greater litter species richness.     

Microbial community composition and function across an arctic tundra landscape

Arctic landscapes are characterized by a diversity of ecosystems, which differ in plant species composition, litter biochemistry, and biogeochemical cycling rates. Tundra ecosystems differing in plant composition should contain compositionally and functionally distinct microbial communities that differentially transform dissolved organic matter as it moves downslope from dry, upland to wet, lowland tundra. To test this idea, we studied soil microbial communities in upland tussock, stream-side birch-willow, and lakeside wet sedge tundra in arctic Alaska, USA. These are a series of ecosystems that differ in topographic position, plant composition, and soil drainage. Phospholipid fatty acid (PLFA) analyses, coupled with compound-specific 13C isotope tracing, were used to quantify microbial community composition and function; we also assayed the activity of extracellular enzymes involved in cellulose, chitin, and lignin degradation. Surface soil from each tundra ecosystem was labeled with 13C-cellobiose,13C-N-acetylglucosamine, or 13C-vanillin. After a five-day incubation, we followed the movement of 13C into bacterial and fungal PLFAs, microbial respiration, dissolved organic carbon, and soil organic matter. Microbial community composition and function were distinct among tundra ecosystems, with tussock tundra containing a significantly greater abundance and activity of soil fungi. Although the majority of 13C-labeled substrates rapidly moved into soil organic matter in all tundra soils (i.e., 50-90% of applied 13C), microbial respiration of labeled substrates in wet sedge tundra soil was lower than in tussock and birch-willow tundra; approximately 8% of 13C-cellobiose and approximately 5% of 13C-vanillin was respired in wet sedge soil vs. 26-38% of 13C-cellobiose and 18-21% of 13C-vanillin in the other tundra ecosystems. Despite these differences, wet sedge tundra exhibited the greatest extracellular enzyme activity. Topographic variation in plant litter biochemistry and soil drainage shape the metabolic capability of soil microbial communities, which, in turn, influence the chemical composition of dissolved organic matter across the arctic tundra landscape.     

Incubation time, functional litter diversity, and habitat characteristics predict litter-mixing effects on decomposition

Plant diversity influences many fundamental ecosystem functions, including carbon and nutrient dynamics, during litter breakdown. Mixing different litter species causes litter mixtures to lose mass at different rates than expected from component species incubated in isolation. Such nonadditive litter-mixing effects on breakdown processes often occur idiosyncratically because their direction and magnitude change with incubation time, litter species composition, and ecosystem characteristics. Taking advantage of results from 18 litter mixture experiments in streams, we examined whether the direction and magnitude of nonadditive mixing effects are randomly determined. Across 171 tested litter mixtures and 510 incubation time-by-mixture combinations, nonadditive effects on breakdown were common and on average resulted in slightly faster decomposition than expected. In addition, we found that the magnitude of nonadditive effects and the relative balance of positive and negative responses in mixtures change predictably over time, and both were related to an index of functional litter diversity and selected environmental characteristics. Based on these, it should be expected that nonadditive effects are stronger for litter mixtures made of functionally dissimilar species especially in smaller streams. Our findings demonstrate that effects of litter diversity on plant mixture breakdown are more predictable than generally thought. We further argue that the consequences of current worldwide homogenization in the composition of plant traits on carbon and nutrient dynamics could be better inferred from long-duration experiments that manipulate both functional litter diversity and ecosystem characteristics in "hotspots of biodiversity effects," such as small streams.     

Whole-system nutrient enrichment increases secondary production in a detritus-based ecosystem

Although the effects of nutrient enrichment on consumer-resource dynamics are relatively well studied in ecosystems based on living plants, little is known about the manner in which enrichment influences the dynamics and productivity of consumers and resources in detritus-based ecosystems. Because nutrients can stimulate loss of carbon at the base of detrital food webs, effects on higher consumers may be fundamentally different than what is expected for living-plant-based food webs in which nutrients typically increase basal carbon. We experimentally enriched a detritus-based headwater stream for two years to examine the effects of nutrient-induced changes at the base of the food web on higher metazoan (predominantly invertebrate) consumers. Our paired-catchment design was aimed at quantifying organic matter and invertebrate dynamics in the enriched stream and an adjacent reference stream for two years prior to enrichment and two years during enrichment. Enrichment had a strong negative effect on standing crop of leaf litter, but no apparent effect on that of fine benthic organic matter. Despite large nutrient-induced reductions in the quantity of leaf litter, invertebrate secondary production during the enrichment was the highest ever reported for headwater streams at this Long Term Ecological Research site and was 1.2-3.3 times higher than predicted based on 15 years of data from these streams. Abundance, biomass, and secondary production of invertebrate consumers increased significantly in response to enrichment, and the response was greater among taxa with larval life spans < or = 1 yr than among those with larval life spans >1 yr. Production of invertebrate predators closely tracked the increased production of their prey. The response of invertebrates was largely habitat-specific with little effect of enrichment on food webs inhabiting bedrock outcrops. Our results demonstrate that positive nutrient-induced changes to food quality likely override negative changes to food quantity for consumers during the initial years of enrichment of detritus-based stream ecosystems. Longer-term enrichment may impact consumers through eventual reductions in the quantity of detritus.