Duration of water flow interruption drives the structure and functional diversity of stream benthic diatoms.

Flow cessation affects river ecosystems submitted to low precipitation and increased water demand, and creates unfavourable conditions to aquatic biological communities. Diatoms are amongst the most sensitive biological groups to hydric stress, making them good indicators of preceding hydrological conditions. We here analyse the response of diatom assemblages to the duration and frequency of non-flow periods in 23 Mediterranean temporary streams. All of them experienced a strong decrease in water flow during summer, leading to a period of flow cessation. In addition, other ten permanent streams (zero dry days during the study period) were included in the study for comparative purposes. Temporary and permanent streams showed similar diatom species richness, evenness, and alpha diversity. However, beta diversity was higher in temporary than permanent streams, regardless rare taxa (<1% occurrence) had similar numbers in the two types of streams. Alpha diversity, richness, and evenness of diatom assemblages in the temporary streams changed with the duration of the non-flow period. Durations of 50-100 days were associated to higher alpha diversity, richness, and evenness, but longer non-flow periods caused their decrease. Diatoms thriving under the most extreme conditions were mostly aerophilic, pioneer, and motile taxa. The proportion of aerophilic diatom taxa increased beyond 100 dry days, particularly in those sites receiving more intense solar radiation. Overall, the taxonomical and functional composition of diatom assemblages mostly responded to the duration of the non-flow period, irrespectively of these being consecutive or not. This study shows that diatom assemblages from temporary streams contain taxa adapted to non-flow conditions, but remain vulnerable to further reductions of water flow associated to climate or global change. 'Unimpacted' watercourses in the Mediterranean region, independently of their flow regime, should be urgently protected and used as taxonomical and functional early indicators of climate change.

Unravelling the effects of multiple stressors on diatom and macroinvertebrate communities in European river basins using structural and functional approaches.

Rivers suffer from more severe decreases in species diversity compared to other aquatic and terrestrial ecosystems due to a variety of pressures related to human activities. Species provide different roles in the functioning of the ecosystem, and their loss may reduce the capacity of the ecosystems to respond to multiple stressors. The effects on diversity will differ based on the type, combination and severity of stressors, as well as on the characteristics of the community composition and tolerance. Multiple trait-based approaches (MTBAs) can help to unravel the effects of multiple stressors on communities, providing a mechanistic interpretation, and, thus, complementing traditional biodiversity assessments using community structure. We studied the relationships between diversity indexes and trait composition of macroinvertebrate and diatom communities, as well as environmental variables that described the hydrological and geomorphological alterations and toxic pollution (pesticides and pharmaceuticals) of three different European river basins: the Adige, the Sava, and the Evrotas. These river basins can be considered representative cases of different situations in European freshwater systems. Hydrological variables were the main drivers determining the community structure and function in the rivers, for both diatoms and macroinvertebrates. For diatom communities, pharmaceutical active compound (PhAC) toxic units were also identified as a very important driver of diversity changes, explaining up to 57% of the variance in taxonomic richness. For macroinvertebrates, river geomorphology was an important driver of structural changes, particularly affecting Plecoptera richness. In addition, PhAC and pesticide toxic units were also identified as stressors for macroinvertebrate communities. MTBA provided a detailed picture of the effects of the stressors on the communities and confirmed the importance of hydrological variables in shaping the functional attributes of the communities.

Effects of olive mill wastewater discharge on benthic biota in Mediterranean streams.

Olive mill wastewaters (OMW) discharging in river ecosystems cause significant adverse effects on their water chemistry and biological communities. We here examined the effects of OMW loads in four streams of a Mediterranean basin characterized by changing flow. The diatom and macroinvertebrate community structures were compared between upstream (control) and downstream (impacted) sites receiving OMW discharge. We also tested if effects occurred at the organism level, i.e. the occurrence of deformities in diatom valves, and the sediment toxicity on the midge Chironomus riparius. We evaluated these effects through a two-year analysis, at various levels of chemical pollution and dilution capacity. The impacted sites had high phenol concentrations and organic carbon loads during and after olive mill (OM) operation, and were characterized by higher abundances of pollution-tolerant diatom and macroinvertebrate taxa. Diatom valve deformities occurred more frequently at the impacted sites. The development of C. riparius was affected by phenolic compounds and organic carbon concentrations in the sediments. The similarity in the diatom and macroinvertebrate assemblages between control and impacted sites decreased at lower flows. Diatoms were more sensitive in detecting deterioration in the biological status of OMW receiving waterways than macroinvertebrates. Our results indicate that the negative effects of OMW extended to the whole benthic community, at both assemblage and organism level.

Multiple stressor effects on biodiversity and ecosystem functioning in a Mediterranean temporary river.

The hydrological and biological complexity of temporary rivers as well as their importance in providing goods and services is increasingly recognized, as much as it is the vulnerability of the biotic communities in view of climate change and increased anthropogenic pressures. However, the effects of flow intermittency (resulting from both seasonal variations and rising hydrological pressure) and pollution on biodiversity and ecosystem functioning have been overlooked in these ecosystems. We explore the way multiple stressors affect biodiversity and ecosystem functioning, as well as the biodiversity-ecosystem functioning (B-EF) relationship in a Mediterranean temporary river. We measured diversity of benthic communities (i.e. diatoms and macroinvertebrates) and related ecosystem processes (i.e. resource use efficiency-RUE and organic matter breakdown-OMB) across a pollution and flow intermittency gradient. Our results showed decreases in macroinvertebrate diversity and the opposite trend in diatom assemblages, whereas ecosystem functioning was negatively affected by both pollution and flow intermittency. The explored B-EF relationships showed contrasting results: RUE decreased with higher diatom diversity, whereas OMB increased with increased macroinvertebrate diversity. The different responses suggest contrasting operating mechanisms, selection effects possibly driving the B-EF relationship in diatoms and complementarity effects driving the B-EF relationship in macroinvertebrates. The understanding of multiple stressor effects on diversity and ecosystem functioning, as well as the B-EF relationship in temporary rivers could provide insights on the risks affecting ecosystem functioning under global change.

Diatom responses to sewage inputs and hydrological alteration in Mediterranean streams.

We analyzed the conjoint effects of sewage inputs and hydrological alteration on the occurrence of teratological forms and on the assemblage composition of stream benthic diatoms. The study was performed in 11 Mediterranean streams which received treated or untreated urban sewage (Impact sites, I), whose composition and morphological anomalies were compared to upstream unaffected (Control, C) sites. The impact sites had high concentrations of ammonium, phosphorus, and pharmaceutical compounds (antibiotics, analgesics, and anti-inflammatories), particularly in those receiving untreated sewage. Impact sites had a higher proportion of teratological forms as well as a prevalence of diatom taxa tolerant to pollution. The differences in the diatom assemblage composition between the paired C and I sites were the largest in the impacted sites that received untreated sewage inputs as well as in the systems with lower dilution capacity. In these sites, the diatom assemblage was composed by a few pollution-tolerant species. Mediterranean river systems facing hydrological stress are highly sensitive to chemical contamination, leading to the homogenization of their diatom assemblages.

Dam regulation and riverine food-web structure in a Mediterranean river.

Flow regimes are a major driver of community composition and structure in riverine ecosystems, and flow regulation by dams often induces artificially-stable flow regimes downstream. This represents a major source of hydrological alteration, particularly in regions where biota is adapted to strong seasonal and interannual flow variability. We hypothesized that dam-induced hydrological stability should increase the availability of autochthonous resources at the base of the food web. This, in turn, should favour herbivorous over detritivorous strategies, increasing the diversity of primary consumers, and the food-web width and length. We tested this hypothesis by studying the longitudinal variation in food-web structure in a highly-seasonal Mediterranean river affected by an irrigation dam. We compared an unregulated reach to several reaches downstream of the dam. Hydrological and sedimentological stability increased downstream of the dam, and altered the type and quantity of available resources downstream, prompting a change from a detritus-based to an algae-based food web. The fraction of links between top and intermediate species also increased, and the food web became longer and wider at the intermediate trophic levels. Food-web structure did not recover 14km downstream of the dam, despite a partial restitution of the flow regime. Our results advance the notion that hydrologic alteration affects riverine food webs via additions/deletions of taxa and variation in the strength and distribution of food-web interactions. Thus, flow regulation by dams may not only impact individual facets of biodiversity, but also food-web level properties across river networks.

Assessing the ecological effects of water stress and pollution in a temporary river - Implications for water management.

Temporary rivers are dynamic and complex ecosystems that are widespread in arid and semi-arid regions, such as the Mediterranean. Biotic communities adapted in their intermittent nature could withstand recurrent drought events. However, anthropogenic disturbances in the form of water stress and chemical pollution challenge biota with unpredictable outcomes, especially in view of climate change. In this study we assess the response of the biotic community of a temporary river to environmental stressors, focusing on water stress and pollution. Towards this aim, several metrics of four biotic groups (diatoms, macrophytes, macroinvertebrates and fish) were applied. All biotic groups responded to a pollution gradient mainly driven by land use, distinct functional groups of all biota responded to water stress (a response related to the rheophilic nature of the species and their resistance to shear stress), while the combined effects of water stress and pollution were apparent in fish. Biotic groups presented a differential temporal response to water stress, where diatom temporal assemblage patterns were explained by water stress variables of short-time response (15days), while the responses of the other biota were associated to longer time periods. There were two time periods of fish response, a short (15days) and a long-time response (60-75days). When considering management decisions, our results indicate that, given the known response of river biota to pollution, biomonitoring of temporary rivers should also involve metrics that can be utilized as early warnings of water stress.

Flood disturbance effects on benthic diatom assemblage structure in a semiarid river network.

Disturbances such as floods and droughts play a central role in determining the structure of riverine benthic biological assemblages. Extreme disturbances from flash floods are often restricted to part of the river network and the magnitude of the flood disturbance may lessen as floods propagate downstream. The present study aimed to characterize the impact of summer monsoonal floods on the resistance and resilience of the benthic diatom assemblage structure in nine river reaches of increasing drainage size within the Gila River in the southwestern United States. Monsoonal floods had a profound effect on the diatom assemblage in the Gila River, but the effects were not related to drainage size except for the response of algal biomass. During monsoons, algal biomass was effectively reduced in smaller and larger systems, but minor changes were observed in medium systems. Resistance and resilience of the diatom assemblage to floods were related to specific species traits, mainly to growth forms. Tightly adhered, adnate and prostrate species (Achnanthidium spp., Cocconeis spp.) exhibited high resistance to repeated scour disturbance. Loosely attached diatoms, such as Nitzschia spp. and Navicula spp., were most susceptible to drift and scour. However, recovery of the diatom assemblage was very quick indicating a high resilience, especially in terms of biomass and diversity. Regional hydroclimatic models predict greater precipitation variability, which will select for diatoms resilient to bed-mobilizing disturbances. The results of this study may help anticipate future benthic diatom assemblage patterns in the southwestern United States resulting from a more variable climate.

Water quality assessment of rivers using diatom metrics across Mediterranean Europe: a methods intercalibration exercise.

The European Water Framework Directive establishes a framework for the protection of water resources. However, common water management tools demand common understanding of assessment methods, so quality goals are equally met. Intercalibration of methods ensures the comparability of biological elements across similar geographical areas. Many aspects can influence the outcome of intercalibration: data sampling, treatment methods, taxonomic reliability of databases, choice of metrics for ecological quality status classification, and criteria for selecting reference sites. This study describes the potentials and constraints of the intercalibration of indices using diatoms for assessment of Mediterranean rivers. Harmonisation of diatom taxonomy and nomenclature was based on a previous ring test which took place at the European level. Four diatom indices (Indice de Polluosensibilité Spécifique-IPS, Indice Biologique Diatomées-IBD 2007, Intercalibration Common Metric Italy-ICMi and Slovenian Ecological Status assessment system) were intercalibrated using data from six European Mediterranean countries (Cyprus, France, Italy, Portugal, Slovenia and Spain). Boundaries between High/Good and Good/Moderate quality classes were harmonised by means of the Intercalibration Common Metric (ICM). Comparability between countries was assured through boundary bias and class agreement. The national boundaries were adjusted when they deviated more than a quarter of a class equivalent (0.25) from the global mean. All national methods correlated well with the ICM, which was sensitive to water quality (negatively correlated to nutrients). Achnanthidium minutissimum sensu lato was the most discriminative species of Good ecological status class. Planothidium frequentissimum, Gomphonema parvulum and Nitzschia palea were the most contributive to Moderate ecological status class. Some taxa were discriminative for both Good and Moderate ecological status classes due to low indication and ecological discriminative power but also due to differences in taxonomy between countries. This intercalibration exercise allowed establishment of common water quality goals across Mediterranean Europe, which is substantiated with the ICM.

Longitudinal development of chlorophyll and phytoplankton assemblages in a regulated large river (the Ebro River).

The distribution of chlorophyll and phytoplankton communities were compared to nutrient concentrations and hydrological parameters along the main stretch of the river Ebro. A progressive increase in planktonic chlorophyll was observed from the upper reaches to the middle section of the river. Chlorophyll reached a maximum (60-80 microg L(-1)) in the meandering section (downstream of the city of Zaragoza), where nutrient inputs (both N and P) and the residence time of the water are very high. In this meandering section phytoplankton assemblages consisted of large centric diatoms and Scenedesmus sp.pl. These longitudinal patterns were interrupted by the presence of three large reservoirs in the lower section of the river. In the section below the reservoirs, the shorter residence water time, the presence of the invasive zebra mussel, and the massive macrophyte development may explain the historical decrease in chlorophyll-a (from 20-45 microg L(-1) in the 1990s to the present 2-5 microg L(-1)). Phytoplankton densities were extremely poor in this section of the river, where large colonial Coelastrum sp.pl. and Pediastrum sp.pl. were the most characteristic taxa.

ALGAL RESPONSE TO NUTRIENT ENRICHMENT IN FORESTED OLIGOTROPHIC STREAM(1).

Nutrient input in streams alters the density and species composition of attached algal communities in open systems. However, in forested streams, the light reaching the streambed (rather than the local nutrient levels) may limit the growth of these communities. A nutrient-enrichment experiment in a forested oligotrophic stream was performed to test the hypothesis that nutrient addition has only minor effects on the community composition of attached algae and cyanobacteria under light limitation. Moderate nutrient addition consisted of increasing basal phosphorus (P) concentrations 3-fold and basal nitrogen (N) concentrations 2-fold. Two upstream control reaches were compared to a downstream reach before and after nutrient addition. Nutrients were added continuously to the downstream reach for 1 year. Algal biofilms growing on ceramic tiles were sampled and identified for more than a year before nutrient addition to 12 months after. Diatoms were the most abundant taxonomic group in the three stream reaches. Nutrient enrichment caused significant variations in the composition of the diatom community. While some taxa showed significant decreases (e.g., Achnanthes minutissima, Gomphonema angustum), increases for other taxa (such as Rhoicosphenia abbreviata and Amphora ovalis) were detected in the enriched reach (for taxonomic authors, see Table 2). Epiphytic and adnate taxa of large size were enhanced, particularly during periods of favorable growth conditions (spring). Nutrients also caused a change in the algal chl a, which increased from 0.5-5.8 to 2.1-10.7 µg chl ·â€Šcm(-2) . Our results indicate that in oligotrophic forested streams, long-term nutrient addition has significant effects on the algal biomass and community composition, which are detectable despite the low light availability caused by the tree canopy. Low light availability moderates but does not detain the long-term tendency toward a nutrient-tolerant community. Furthermore, the effects of nutrient addition on the algal community occur in spite of seasonal variations in light, water flow, and water chemical characteristics, which may confound the observations. [Table: see text].