Verónica Ferreira

Continental-scale effects of nutrient pollution on stream ecosystem functioning.

Reading the Leaves Excess inputs of nutrients—a type of pollution known as eutrophication—threatens biodiversity and water quality in rivers and streams. Woodward et al. (p. 1438; see the Perspective by Palmer and Febria) studied how one key ecosystem process—leaf-litter decomposition—responds to eutrophication across a large nutrient pollution gradient in 100 European streams. Leaf breakdown was stimulated by low to moderate nutrient concentrations but was inhibited at high rates of nutrient loading. Leaf-litter breakdown rates across 100 European streams offer insights into ecosystem health during eutrophication. Excessive nutrient loading is a major threat to aquatic ecosystems worldwide that leads to profound changes in aquatic biodiversity and biogeochemical processes. Systematic quantitative assessment of functional ecosystem measures for river networks is, however, lacking, especially at continental scales. Here, we narrow this gap by means of a pan-European field experiment on a fundamental ecosystem process—leaf-litter breakdown—in 100 streams across a greater than 1000-fold nutrient gradient. Dramatically slowed breakdown at both extremes of the gradient indicated strong nutrient limitation in unaffected systems, potential for strong stimulation in moderately altered systems, and inhibition in highly polluted streams. This large-scale response pattern emphasizes the need to complement established structural approaches (such as water chemistry, hydrogeomorphology, and biological diversity metrics) with functional measures (such as litter-breakdown rate, whole-system metabolism, and nutrient spiraling) for assessing ecosystem health.

A global experiment suggests climate warming will not accelerate litter decomposition in streams but might reduce carbon sequestration

The decomposition of plant litter is one of the most important ecosystem processes in the biosphere and is particularly sensitive to climate warming. Aquatic ecosystems are well suited to studying warming effects on decomposition because the otherwise confounding influence of moisture is constant. By using a latitudinal temperature gradient in an unprecedented global experiment in streams, we found that climate warming will likely hasten microbial litter decomposition and produce an equivalent decline in detritivore-mediated decomposition rates. As a result, overall decomposition rates should remain unchanged. Nevertheless, the process would be profoundly altered, because the shift in importance from detritivores to microbes in warm climates would likely increase CO(2) production and decrease the generation and sequestration of recalcitrant organic particles. In view of recent estimates showing that inland waters are a significant component of the global carbon cycle, this implies consequences for global biogeochemistry and a possible positive climate feedback.

Global distribution of a key trophic guild contrasts with common latitudinal diversity patterns

Most hypotheses explaining the general gradient of higher diversity toward the equator are implicit or explicit about greater species packing in the tropics. However, global patterns of diversity within guilds, including trophic guilds (i.e., groups of organisms that use similar food resources), are poorly known. We explored global diversity patterns of a key trophic guild in stream ecosystems, the detritivore shredders. This was motivated by the fundamental ecological role of shredders as decomposers of leaf litter and by some records pointing to low shredder diversity and abundance in the tropics, which contrasts with diversity patterns of most major taxa for which broad-scale latitudinal patterns haven been examined. Given this evidence, we hypothesized that shredders are more abundant and diverse in temperate than in tropical streams, and that this pattern is related to the higher temperatures and lower availability of high-quality leaf litter in the tropics. Our comprehensive global survey (129 stream sites from 14 regions on six continents) corroborated the expected latitudinal pattern and showed that shredder distribution (abundance, diversity and assemblage composition) was explained by a combination of factors, including water temperature (some taxa were restricted to cool waters) and biogeography (some taxa were more diverse in particular biogeographic realms). In contrast to our hypothesis, shredder diversity was unrelated to leaf toughness, but it was inversely related to litter diversity. Our findings markedly contrast with global trends of diversity for most taxa, and with the general rule of higher consumer diversity at higher levels of resource diversity. Moreover, they highlight the emerging role of temperature in understanding global patterns of diversity, which is of great relevance in the face of projected global warming.

A meta-analysis of the effects of nutrient enrichment on litter decomposition in streams

The trophic state of many streams is likely to deteriorate in the future due to the continuing increase in human‐induced nutrient availability. Therefore, it is of fundamental importance to understand how nutrient enrichment affects plant litter decomposition, a key ecosystem‐level process in forest streams. Here, we present a meta‐analysis of 99 studies published between 1970 and 2012 that reported the effects of nutrient enrichment on litter decomposition in running waters. When considering the entire database, which consisted of 840 case studies, nutrient enrichment stimulated litter decomposition rate by approximately 50%. The stimulation was higher when the background nutrient concentrations were low and the magnitude of the nutrient enrichment was high, suggesting that oligotrophic streams are most vulnerable to nutrient enrichment. The magnitude of the nutrient‐enrichment effect on litter decomposition was higher in the laboratory than in the field experiments, suggesting that laboratory experiments overestimate the effect and their results should be interpreted with caution. Among field experiments, effects of nutrient enrichment were smaller in the correlative than in the manipulative experiments since in the former the effects of nutrient enrichment on litter decomposition were likely confounded by other environmental factors, e.g. pollutants other than nutrients commonly found in streams impacted by human activity. However, primary studies addressing the effect of multiple stressors on litter decomposition are still few and thus it was not possible to consider the interaction between factors in this review. In field manipulative experiments, the effect of nutrient enrichment on litter decomposition depended on the scale at which the nutrients were added: stream reach > streamside channel > litter bag. This may have resulted from a more uniform and continuous exposure of microbes and detritivores to nutrient enrichment at the stream‐reach scale. By contrast, nutrient enrichment at the litter‐bag scale, often by using diffusing substrates, does not provide uniform controllable nutrient release at either temporal or spatial scales, suggesting that this approach should be abandoned. In field manipulative experiments, the addition of both nitrogen (N) and phosphorus (P) resulted in stronger stimulation of litter decomposition than the addition of N or P alone, suggesting that there might be nutrient co‐limitation of decomposition in streams. The magnitude of the nutrient‐enrichment effect on litter decomposition was higher for wood than for leaves, and for low‐quality than for high‐quality leaves. The effect of nutrient enrichment on litter decomposition may also depend on climate. The tendency for larger effect size in colder regions suggests that patterns of biogeography of invertebrate decomposers may be modulating the effect of nutrient enrichment on litter decomposition. Although studies in temperate environments were overrepresented in our database, our meta‐analysis suggests that the effect of nutrient enrichment might be strongest in cold oligotrophic streams that depend on low‐quality plant litter inputs.

Stream Ecosystem Functioning in an Agricultural Landscape: The Importance of Terrestrial-Aquatic Linkages

The loss of native riparian vegetation and its replacement with non-native species or grazing land for agriculture is a worldwide phenomenon, but one that is prevalent in Europe, reflecting the heavily-modified nature of the continents landscape. The consequences of these riparian alterations for freshwater ecosystems remain largely unknown, largely because bioassessment has traditionally focused on the impacts of organic pollution on community structure. We addressed the need for a broader perspective, which encompasses changes at the catchment scale, by comparing ecosystem processes in woodland reference sites with those with altered riparian zones. We assessed a range of riparian modifications, including clearance for pasture and replacement of woodland with a range of low diversity plantations, in 100 streams to obtain a continental-scale perspective of the major types of alterations across Europe. Subsequently, we focused on pasture streams, as an especially prevalent widespread riparian alteration, by characterising their structural (e.g. invertebrate and fish communities) and functional (e.g. litter decomposition, algal production, herbivory) attributes in a country (Ireland) dominated by this type of landscape modification, via field and laboratory experiments. We found that microbes became increasingly important as agents of decomposition relative to macrofauna (invertebrates) in impacted sites in general and in pasture streams in particular. Resource quality of grass litter (e.g., carbon : nutrient ratios, lignin and cellulose content) was a key driver of decomposition rates in pasture streams. These systems also relied more heavily on autochthonous algal production than was the case in woodland streams, which were more detrital based. These findings suggest that these pasture streams might be fundamentally different from their native, ancestral woodland state, with a shift towards greater reliance on autochthonous-based processes. This could have a destabilizing effect on the dynamics of the food web relative to the slower, detrital-based pathways that dominate in woodland streams.

Future increase in temperature more than decrease in litter quality can affect microbial litter decomposition in streams

The predicted increase in atmospheric CO2 concentration for this century is expected to lead to increases in temperature and changes in litter quality that can affect small woodland streams, where water temperature is usually low and allochthonous organic matter constitutes the basis of the food web. We have assessed the individual and interactive effect of water temperature (5 and 10°C) and alder litter quality produced under ambient CO2 levels (ambient litter) or under CO2 concentrations predicted for 2050 (elevated litter) on litter decomposition and on fungal activity and assemblage structure. Litter decomposition rates and fungal respiration rates were significantly faster at 10 than at 5°C, but they were not affected by litter quality. Litter quality affected mycelial biomass accrual at 5 but not at 10°C, while increases in temperature stimulated biomass accrual on ambient but not on elevated litter. A similar pattern was observed for conidial production. All variables were stimulated on elevated litter at 10°C (future scenario) compared with ambient litter at 5°C (present scenario), but interactions between temperature and litter quality were additive. Temperature was the factor that most strongly affected the structure of aquatic hyphomycete assemblages. Our results indicate that if future increases in atmospheric CO2 lead to only slight modifications in litter quality, the litter decomposition and fungal activities and community structure will be strongly controlled by increased water temperature. This may have serious consequences for aquatic systems as faster litter decomposition may lead to food depletion for higher trophic levels.

Role of physical fragmentation and invertebrate activity in the breakdown rate of leaves

We evaluated the relative importance of current velocity and invertebrate activities in the breakdown rate of alder (Alnus glutinosa (L.) Gaertner) leaves. Decomposition experiments were carried out in artificial channels, where current velo- city and shredder presence were manipulated, and in a 4 th order stream, in both summer and autumn, where litter bags were incubated in several reaches differing in both depth and current velocity. Alder leaves incubated in artificial channels decomposed signifi- cantly faster in the presence of shredders than in their absence (k = 0.0368/d vs. k = 0.0210/d in low current and k = 0.0472/d vs. k = 0.0219/d in high current). However, cur- rent (up to 2.35 m/s) had no significant effect on decomposition rates. In channels with- out invertebrates, no significant differences in k values were found between coarse and fine mesh bags in high (0.20 m/s) and low (0.05 m/s) current. Leaves incubated in the stream during summer, in reaches with current velocity ranging from 0.003 to 1.185 m/s, did not differ in their decomposition rates (k = 0.0489/d to k = 0.0645/d). In autumn, leaves exposed to high current (1.228 m/s) had faster decomposition rate (k = 0.0417/d vs. k = 0.0136/d), which may be related to sediment transport during this time of the year or to the tendency for higher number of shredders in high current-shallow reaches.

Effects of litter diversity on decomposition and biological colonization of submerged litter in temperate and tropical streams

Abstract.  Detrital food webs of woodland streams depend on terrestrial litter input and, thus, are susceptible to changes in riparian cover. We assessed effects of litter species richness and quality on decomposition and associated biological communities in temperate deciduous forest and tropical rainforest streams. Three native litter species were incubated in each stream in all combinations (7 litter treatments, 3 richness levels) in coarse- (invertebrate access) and fine-mesh bags (no invertebrate access) and were sampled 5 times over 74 (temperate stream) or 94 d (tropical stream). Decomposition, and fungal biomass, sporulation, and species richness were measured for each treatment. Alnus glutinosa litter was incubated in both streams to assess effects of environmental and biological differences between streams on litter decomposition. Biological colonization (number of fungal species, fungal biomass) and activity (conidial production) were lower in the tropical than the temperate stream, despite its higher water temperature (24 vs 8°C). Mass loss for individual species reached 95% in the temperate and 60% in the rainforest stream. Decomposition rates in mixtures were unaffected by litter richness but could be predicted from their initial N, phenol, and lignin concentrations (leaf quality). In the temperate stream, Alnus decomposition in coarse-mesh bags was positively related to litter richness, and Alnus stimulated decomposition of mixtures. Microbial O2 consumption, fungal biomass accrual, aquatic hyphomycete sporulation rate and richness, and shredder abundance and richness were insensitive to litter richness. In the temperate stream, presence of tough litter inhibited invertebrate colonization of mixtures, whereas in the tropical stream, presence of soft litter stimulated invertebrate colonization of mixtures. Litter quality (species identity), not richness, was the main controller of decomposition of litter mixtures, and decomposition of litter in mixtures may differ from decomposition of individual species. Thus, disappearance or introduction of key species might affect organic matter processing in streams.

Eucalyptus plantations affect fungal communities associated with leaf-litter decomposition in iberian streams

The replacement of diverse deciduous forests by eucalyptus plantations changes the timing, quality and quantity of litter inputs to streams, which has the potential to affect the activity of decomposers and thus ecosystem functioning. Here, we compared (a) the decomposition rate of alder and oak leaves incubated in deciduous and eucalyptus streams in Spain and Portugal, (b) the activity (fungal biomass and sporulation) and diversity (species richness and Pielous evenness index) of the associated fungal communities and (c) changes in N and P content of leaves. Alder and oak leaves decomposed at similar rates in both stream types and countries, with the exception of oak leaves in the Spanish eucalyptus stream, which decomposed faster than in the corresponding deciduous stream or in the Portuguese eucalyptus stream. This difference was attributed to physical fragmentation due to flooding and not to forest cover. Higher nitrogen and phosphorus content and higher fungal biomass and sporulation were generally found on leaves from eucalyptus rather than from deciduous streams. The higher fungal activity in eucalyptus streams was attributed to higher water temperature and benthic organic matter storage. The Spanish eucalyptus stream had higher species richness of aquatic hyphomycetes than the deciduous one (27 vs. 20) while in Portugal the opposite was true (16 vs. 20). Fungal community evenness was significantly higher on alder leaves in eucalyptus than in deciduous streams. The community structure (MDS analysis) discriminated both stream types in Portugal much better than it did in Spain. At least for Portugal, differences between stream types can be explained by higher litter diversity in deciduous than in eucalyptus streams. In conclusion, stream fungal communities in Portugal were more affected by eucalyptus plantations than in Spain. In both countries, fungal diversity and activity were more affected by eucalyptus plantations than decomposition rates of submerged litter. We suggest therefore that, to mitigate the effect of eucalyptus plantations, deciduous trees could be planted on the river banks or, preferably, riparian strips of native vegetation should be left unmodified.

A predictive model for freshwater bioassessment (Mondego River, Portugal)

We sampled macroinvertebrates at 75 locations in the Mondego river catchment, Central Portugal, and developed a predictive model for water quality assessment of this basin, based on the Reference Condition Approach. Sampling was done from June to September 2001. Fifty-five sites were identified as “Reference sites” and 20 sites were used as “Test sites” to test the model. At each site we also measured 40 habitat variables to characterize water physics and chemistry, habitat type, land use, stream hydrology and geographic location. Macroinvertebrates were generally identified to species or genus level; a total of 207 taxa were found. By Unweighted Pair Group Method with Arithmetic mean (UPGMA) clustering and analysis of species contribution to similarities percentage (SIMPER), two groups of reference sites were established. Using Discriminant Analysis (stepwise forward), four variables correctly predicted 78% of the reference sites to the appropriate group: stream order, pool quality, substrate quality and current velocity. Test sites’ environmental quality was established from their relative distance to reference sites, in MDS ordination space, using a series of bands (BEAST methodology). The model performed well at upstream sites, but at downstream sites it was compromised by the lack of reference sites. As with the English RIVPACS predictive model, the Mondego model should be continually improved with the addition of new reference sites. The adaptation of the Mondego model methodology to the Water Framework Directive is possible and would consist mainly of the integration of the WFD typology and increasing the number of ellipses that define quality bands.