Summary 1. Mediterranean climate regions are characterised by long summer droughts that usually involve flow intermittency in low‐ to mid‐order streams. Flow intermittency implies flow cessation, drying and subsequent rewetting of the streambed, and affects both autotrophic and heterotrophic processes. The balance between these processes, as well as the balance in the use of carbon (C), nitrogen (N) and phosphorus (P) may change because of the ongoing increase in stream flow intermittency caused by global change in many regions. It is therefore crucial to understand better the consequences of this phenomenon. 2. Our two initial hypotheses were (i) that flow intermittency would impact more on autotrophic than on heterotrophic processes in stream biofilms owing to the higher water dependence of autotrophs, as well as differences in the water storage capacity of the stream biofilm compartments where autotrophic and heterotrophic processes mainly occur (surface cobbles versus hyporheic sediments) and (ii) that the C‐N‐P use by biofilms would change during the dry period (terrestrial phase) owing to the extreme water stress conditions. These hypotheses were tested by analysing the functional response of the main stream biofilms (epilithic, epipsammic and hyporheic) during flow cessation, desiccation and rewetting in a Mediterranean forested stream. The autotrophic response was characterised through changes in the photon yield, whereas the heterotrophic response was characterised by changes in the extracellular enzyme activities. 3. Streambed desiccation had clear effects on the functioning of stream biofilms. Autotrophic biomass decreased by 80% with streambed desiccation, but recovered rapidly after flow resumption. Heterotrophs were more resistant to water stress, especially in the epipsammic and hyporheic biofilms where bacterial cell density decreased only by 20%. 4. Extracellular enzyme activities remained relatively high, and the balance in the C‐N‐P use by biofilms changed during the dry period. The C and P breakdown capacities were maintained during dry conditions, especially in the epipsammic and hyporheic biofilms, but the degradation of N compounds sharply decreased. Elemental molar ratios (C:N and C:P) of the different biofilms also changed with streambed desiccation. C:P ratios increased from 80 to 300, while the C:N ratios increased from 10 to 16. 5. Given the contrasting responses of autotrophic and heterotrophic processes in the different biofilms, our results suggest that the current increase in flow intermittency extent is likely to increase the relative importance of heterotrophic processes in stream ecosystems, as well as the relative contribution of the hyporheic biofilm to C‐N‐P use. Our results further suggest that the longer streams remain dry, the more the biofilm stoichiometry will change.
Temporary streams are a dominant surface water type in the Mediterranean region. As a consequence of their hydrologic regime, these ecosystems contract and fragment as they dry, and expand after rewetting. Global change leads to a rapid increase in the extent of temporary streams, and more and more permanent streams are turning temporary. Consequently, there is an urgent need to better understand the effects of flow intermittency on the biogeochemistry and ecology of stream ecosystems. Our aim was to investigate how stream nutrient availability varied in relation to ecosystem contraction, fragmentation and expansion due to hydrologic drying and rewetting. We quantified the temporal and spatial changes in dissolved nitrogen (N) and phosphorus (P) concentrations along a reach of a temporary Mediterranean forest stream during an entire contraction-fragmentation-expansion hydrologic cycle. We observed marked temporal changes in N and P concentrations, in the proportion of organic and inorganic forms as well as in stoichiometric ratios, reflecting shifts in the relative importance of in-stream nutrient processing and external nutrient sources. In addition, the spatial heterogeneity of N and P concentrations and their ratios increased substantially with ecosystem fragmentation, reflecting the high relevance of in-stream processes when advective transport was lost. Overall, changes were more pronounced for N than for P. This study emphasizes the significance of flow intermittency in regulating stream nutrient availability and its implications for temporary stream management. Moreover, our results point to potential biogeochemical responses of these ecosystems in more temperate regions under future water scarcity scenarios.
Temporary streams are characterized by the alternation of dry and wet hydrological phases, creating both a harsh environment for the biota as well as a high diversity of opportunities for adaptation. These systems are mainly microbial-based during several of these hydrological phases, and those growing on all solid substrata (biofilms) accordingly change their physical structure and community composition. Biofilms experience large decreases in cell densities and biomass, both of bacteria and algae, during dryness. Algal and bacterial communities show remarkable decreases in their diversity, at least locally (at the habitat scale). Biofilms also respond with significant physiological plasticity to each of the hydrological changes. The decreasing humidity of the substrata through the drying process, and the changing quantity and quality of organic matter and nutrients available in the stream during that process, causes unequal responses on the biofilm bacteria and algae. Biofilm algae are affected faster than bacteria by the hydric stress, and as a result the ecosystem respiration resists longer than gross primary production to the increasing duration of flow intermittency. This response implies enhancing ecosystem heterotrophy, a pattern that can be exacerbated in temporary streams suffering of longer dry periods under global change.Keywords: biofilm, dry-rewetting cycle, temporary, intermittency, bacteria, algae, Mediterranean THE RELEVANCE OF TEMPORARY STREAMS IN THE WORLDFlow intermittency is part of the natural hydrology for streams and rivers, especially in arid and semi-arid landscapes. Temporary streams, or those that cease to flow at some points in space and time along their course, are a large fraction of many river networks, basically in tributaries of small order, but also in segments in the middle and lower parts of river networks. The number of temporary streams has been probably underestimated , as it appears from the application of novel on-site sensors, advances in remote sensing, and new modeling approaches. Flow intermittency has been estimated to account for 69% of first-order streams below 60 • , and up to 34% of larger order rivers (Raymond et al., 2013). Flow intermittency produces periodic or eventrelated loss of hydrologic connectivity between stream compartments, with consequences for all the processes ongoing in a waterway. Thus, flow intermittency triggers a chain of cascading effects eventually affecting water chemistry and biogeochemistry, as well as biodiversity, and ultimately community structure and ecosystem functioning.The direct implications of climate change and anomalous climatic events on flow regime are behind of the increasing temporality of many streams and rivers. Complex planetary scale processes, as well as local processes, may justify these situations. Anomalies such as the El Niño
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