Aingeru Martínez

Stream regulation by small dams affects benthic macroinvertebrate communities: from structural changes to functional implications

We studied benthic macroinvertebrate communities upstream and downstream of five small reservoirs (surface release in autumn–winters) (north Spain) to assess the effect of flow regulation on structural and functional characteristics of stream ecosystems. We based our approach on the use of structural metrics (density, biomass, richness and diversity) in combination with two functional diversity indices based on biological and ecological traits: FDPG index, related to species richness, and FDQ, which incorporates evenness across taxa. Although water physicochemical parameters were unaffected by the reservoirs during the study period (autumn–winter), macroinvertebrate metrics were lower below the dams, with detritivores (shredders and collector-gatherers) being the most affected. The alder leaf breakdown rate estimated by the litter-bag technique was related to the density, biomass, richness, diversity and FDPG index of shredders, compromising the ecosystem functioning. The most plausible origin for the observed differences in macroinvertebrate metrics between upstream and downstream reaches was the change of the flow regime caused by the impoundments at downstream sites, leading to droughts in summer in those naturally permanently flowing streams. The observed functional diversity loss might reduce the chances of the community to override natural or man-induced fluctuations in their environment with possible repercussions on important ecosystem functions and services.

Temperature affects leaf litter decomposition in low-order forest streams: field and microcosm approaches.

Despite predicted global warming, the temperature effects on headwater stream functioning are poorly understood. We studied these effects on microbial-mediated leaf decomposition and the performance of associated aquatic hyphomycete assemblages. Alder leaves were incubated in three streams differing in winter water temperature. Simultaneously, in laboratory, leaf discs conditioned in these streams were incubated at 5, 10 and 15 °C. We determined mass loss, leaf N and sporulation rate and diversity of aquatic hyphomycete communities. In the field, decomposition rate correlated positively with temperature. Decomposition rate and leaf N presented a positive trend with dissolved nutrients, suggesting that temperature was not the only factor determining the process velocity. Under controlled conditions, it was confirmed that decomposition rate and leaf N were positively correlated with temperature, leaves from the coldest stream responding most clearly. Sporulation rate correlated positively with temperature after 9 days of incubation, but negatively after 18 and 27 days. Temperature rise affected negatively the sporulating fungi richness and diversity only in the material from the coldest stream. Our results suggest that temperature is an important factor determining leaf processing and aquatic hyphomycete assemblages and that composition and activity of fungal communities adapted to cold environments could be more affected by temperature rises. Highlight: Leaf decomposition rate and associated fungal communities respond to temperature shifts in headwater streams.

Effects of flow scarcity on leaf-litter processing under oceanic climate conditions in calcareous streams.

Although temporary streams represent a high proportion of the total number and length of running waters, historically the study of intermittent streams has received less attention than that of perennial ones. The goal of the present study was to assess the effects of flow cessation on litter decomposition in calcareous streams under oceanic climate conditions. For this, leaf litter of alder was incubated in four streams (S1, S2, S3 and S4) with different flow regimes (S3 and S4 with zero-flow periods) from northern Spain. To distinguish the relative importance and contribution of decomposers and detritivores, fine- and coarse-mesh litter bags were used. We determined processing rates, leaf-C, -N and -P concentrations, invertebrate colonization in coarse bags and benthic invertebrates. Decomposition rates in fine bags were similar among streams. In coarse bags, only one of the intermittent streams, S4, showed a lower rate than that in the other ones as a consequence of lower invertebrate colonization. The material incubated in fine bags presented higher leaf-N and -P concentrations than those in the coarse ones, except in S4, pointing out that the decomposition in this stream was driven mainly by microorganisms. Benthic macroinvertebrate and shredder density and biomass were lower in intermittent streams than those in perennial ones. However, the bags in S3 presented a greater amount of total macroinvertebrates and shredders comparing with the benthos. The most suitable explanation is that the fauna find a food substrate in bags less affected by calcite precipitation, which is common in the streambed at this site. Decomposition rate in coarse bags was positively related to associated shredder biomass. Thus, droughts in streams under oceanic climate conditions affect mainly the macroinvertebrate detritivore activity, although macroinvertebrates may show distinct behavior imposed by the physicochemical properties of water, mainly travertine precipitation, which can override the flow intermittence effects.

Leaf litter decomposition of native and introduced tree species of contrasting quality in headwater streams: how does the regional setting matter?

Terrestrial plant litter is important in sustaining stream food webs in forested headwaters. Leaf litter quality often decreases when native species are replaced by introduced species, and a lower quality of leaf litter inputs may alter litter decomposition at sites afforested with non-native species. However, since detritivore composition and resource use plasticity may depend on the prevalent litter inputs, the extent of the alteration in decomposition can vary between streams. We tested 2 hypotheses using 2 native and 3 introduced species of tree differing in quality in 4 Iberian regions with contrasting vegetational traits: 1) decomposition rates of all plant species would be higher in regions where streams normally receive litter inputs of lower rather than higher quality; 2) a higher resource-use plasticity of detritivores in regions vegetated with plants of lower litter quality will cause a greater evenness in decomposition rates among plant species compared to regions where streams normally receive higher-quality plant litter inputs. Results showed a highly consistent interspecific ranking of decomposition rates across regions driven by litter quality, and a significant regional effect. Hypothesis 1 was supported: decomposition rates of the five litter types were generally higher in streams from regions vegetated with species producing leaf litter of low quality, possibly due to the profusion of caddisfly shredders in their communities. Hypothesis 2 was not supported: the relative differences in decomposition rates among leaf litter species remained essentially unaltered across regions. Our results suggest that, even in regions where detritivores can be comparatively efficient using resources of low quality, caution is needed particularly when afforestation programs introduce plant species of lower litter quality than the native species.

River ecosystem processes: a synthesis of approaches, criteria of use and sensitivity to environmental stressors

River ecosystems are subject to multiple stressors that affect their structure and functioning. Ecosystem structure refers to characteristics such as channel form, water quality or the composition of biological communities, whereas ecosystem functioning refers to processes such as metabolism, organic matter decomposition or secondary production. Structure and functioning respond in contrasting and complementary ways to environmental stressors. Moreover, assessing the response of ecosystem functioning to stressors is critical to understand the effects on the ecosystem services that produce direct benefits to humans. Yet, there is more information on structural than on functional parameters, and despite the many approaches available to measure river ecosystem processes, structural approaches are more widely used, especially in management. One reason for this discrepancy is the lack of synthetic studies analyzing river ecosystem functioning in a way that is useful for both scientists and managers. Here, we present a synthesis of key river ecosystem processes, which provides a description of the main characteristics of each process, including criteria guiding their measurement as well as their respective sensitivity to stressors. We also discuss the current limitations, potential improvements and future steps that the use of functional measures in rivers needs to face.

Leaf-litter quality effects on stream ecosystem functioning: a comparison among fi ve species

The replacement of native vegetation by exotic species has the potential to impact detritus based stream ecosystems. To test the possible consequences of vegetation alteration on stream ecosystem functioning we com- pared the processing rate and invertebrate colonisation of leaves from fi ve different tree species, two natives (Alnus glutinosa and Quercus pyrenaica) and three exotics (Populus × canadensis, Platanus hispanica and Pinus radiata) in three unaltered forested streams in the Atlantic region of northern Spain. Breakdown rates, nitrogen concentra- tion and C:N ratio of incubated leaves, and invertebrate communities in the benthos and bags were analysed. Break- down rates differed among leaf species and were positively related to initial nitrogen concentration and negatively to the C:N ratio. After the incubation period (6 -7 weeks), the differences in nutritional quality among leaf species and the relationship between breakdown rates and litter quality persisted. The density and biomass of macroinverte- brates and shredders associated with litter bags correlated with leaf-nitrogen concentration and explained the differ- ences in breakdown rates. The results indicate that the replacement of native deciduous forest by plant commercial species with leaf-litter of low quality such as plane and pine can affect in-stream organic matter processing and the subsequent transference of energy throughout the food web.

In-stream litter decomposition along an altitudinal gradient: does substrate quality matter?

In temperate streams, water temperature and organic matter inputs from surrounding forest vary along the altitude. We tested if the different features of streams of similar size determined by an altitudinal gradient might differentially affect the processing rate of different quality leaves (alder, oak and beech). To distinguish the relative contribution of microbial decomposition from overall decomposition, fine- and coarse-mesh bags were used. We determined decomposition rates, leaf-N and -P concentration, microbial respiration (fine bags), invertebrate colonisation (coarse bags) and density and identity of benthic invertebrates in three second-order streams. Alder decomposed faster than the other species in all three streams and regardless of mesh size due to its lower values of C:N, C:P and N:P. Unexpectedly, microbial decomposition rate did not vary among streams for any of the leaf species. The total decomposition rate of alder and oak showed a negative trend along the altitudinal gradient, the magnitude of the change in decomposition rates being similar for both species. The density and structure of the invertebrate community differed along the altitudinal gradient, related to temperature and surrounding vegetation, determining the decomposition rate. Unexpectedly, sensitivity of decomposition rate of different quality leaves to temperature does not differ along the gradient.

Climate modulates the magnitude of the effects of flow regulation on leaf-litter decomposition

The need of water for human use has led the impact on running waters of flow regulation to be of a global-scale. Although the effects of this impact have been widely investigated, efforts have been focused on large dams, so information about small reservoirs and their effects on ecosystem functioning is lacking. A recent collaborative project (IMPARIOS) addressed the effects of flow regulation by small impoundments on leaf-litter decomposition, a key function in low order streams which contributes greatly to the global carbon cycle. Flow regulation was found to affect ecosystem functioning reducing decomposition rate by altering shredders, but the magnitude of change varied among the different sub-climatic regions. The current project examined whether climatic variables modulate the effect of flow regulation on decomposition. For this, 19 bioclimatic variables were studied in relation to the leaf-litter decomposition rate and associated variables (sporulation rate and richness of aquatic hyphomycetes, and richness, density and biomass of total macroinvertebrates and shredders) in 17 streams impacted by regulation structures distributed in four sub-climatic regions within Spain. Overall, decomposition was slower below structures and climate influenced the magnitude of reduction. Effect sizes were negatively related to the seasonal changes in temperature and precipitation and to the general water deficit of the locations. In the future, the forecasted increase of seasonality in precipitation and temperature and the expected increase of number of dams to meet the needs of growing population may exacerbate the effects of flow regulation, altering nutrient recycling and the carbon cycle globally.

Leaf Traits Drive Plant Diversity Effects On Litter Decomposition And FPOM Production In Streams

Biodiversity loss in riparian forests has the potential to alter rates of leaf litter decomposition in stream ecosystems. However, studies have reported the full range of positive, negative and no effects of plant diversity loss on decomposition, and there is currently no explanation for such inconsistent results. Furthermore, it is uncertain whether plant diversity loss affects other ecological processes related to decomposition, such as fine particulate organic matter production or detritivore growth, which precludes a thorough understanding of how detrital stream food webs are impacted by plant diversity loss. We used a microcosm experiment to examine the effects of plant diversity loss on litter decomposition, fine particulate organic matter production, and growth of a dominant leaf-shredding detritivore, using litter mixtures varying in species composition. We hypothesized that plant diversity loss would decrease the rates of all studied processes, but such effects would depend on the leaf traits present in litter mixtures (both their average values and their variability). Our findings partly supported our hypotheses, showing that plant diversity loss had a consistently negative effect on litter decomposition and fine particulate organic matter production (but not on detritivore growth) across litter mixtures, which was mediated by detritivores. Importantly, the magnitude of the diversity effect and the relative importance of different mechanisms underlying this effect (i.e., complementarity vs. selection) varied depending on the species composition of litter mixtures, mainly because of differences in litter nutritional quality and trait variability. Complementarity was prevalent but varied in size, with positive selection effects also occurring in some mixtures. Our results support the notion that loss of riparian plant species is detrimental to key stream ecosystem processes that drive detrital food webs, but that the magnitude of such effects largely depends on the the order of species loss.

Responses of Aquatic Hyphomycetes to Temperature and Nutrient Availability: a Cross-transplantation Experiment

Aquatic hyphomycetes represent a large component of the microbial assemblage that decomposes submerged leaf-litter in fluvial ecosystems. The structure and activity of these fungal decomposers depend on environmental factors. Fungal communities may adapt to local habitat conditions; however, little is known about how fungal communities respond to abrupt changes in factors such as nutrient availability and temperature. To respond to this question, we carried out a cross-transplantation experiment, which assessed the decomposer activity and structure of this microbial community on decaying leaves transplanted from a cold and oligotrophic stream (S1) to a warmer and nitrogen-richer one (S2) and vice versa. Results were compared to those from untransplanted leaves decomposing either at S1 or at S2. In terms of days, untransplanted leaves were decomposed at a similar rate in both streams; the change to warmer and nitrogen-richer waters (S1 ➔ S2) significantly enhanced the decomposition process while the reciprocal transplantation (S2 ➔ S1) did not alter decomposition rate. However, when standardizing the temperature effects by using degree-days, microbial decomposers under colder conditions were more efficient in terms of accumulated heat, independent of the initial or final incubation site. Regarding community structure, taxa richness and diversity of aquatic hyphomycetes appear to be favoured under warmer and richer conditions, increasing after transplantation to S2 but with little effect on the predominant taxa. However, the reciprocal transplantation (S2 ➔ S1) yielded a clear decline of the dominant taxa at S2 (Lunulospora curvula) in favour of the local dominant ones. Thus, effects of environmental changes on activity and community structure can be highly variable and not always clearly linked or reciprocal. Therefore, results from simplified experimental designs (e.g. artificial assemblages under laboratory conditions) must be taken with caution. Additional field studies and manipulative experimentation dealing with natural communities are required when trying to extend individual results to complex scenarios such as those projected by global change.