The influence of Mediterranean riparian forests on stream nitrogen dynamics: a review from a catchment perspective

The influence ofMediterranean riparian forests on stream nitrogen dynamics: a review from a catchment perspective Riparian zones are considered natural filters of nitrogen (N) within catchments because they can substantially diminish the exports of N from terrestrial to aquatic ecosystems. However, understanding the influence of riparian zones on regulating N exports at the catchment scale still remains a big challenge in ecology, mainly because upscaling plot scale results is difficult, as it is disentangling the effects of riparian, upland, and in-stream processes on stream water chemistry. In this review, we summarize previous studies examining key hydrological and biogeochemical processes by which Mediterranean riparian zones regulate catchment water and N exports.We focus onMediterranean regions because they experience a marked climatic seasonality that facilitates disentangling the close link between climate, riparian hydrology, and stream N exports. We show that Mediterranean riparian soils can be hot spots of N mineralization and nitrification within catchments given their relatively moist conditions and large stocks of N-rich leaf litter. Extremely large nitrification rates can occur during short-time periods (i.e. hot moments) and lead to increases in stream N loads, suggesting that riparian soils can be a potential source of N to adjacent aquatic systems. Moreover, riparian trees can contribute to decrease riparian groundwater level during the vegetative period, and promote reverse fluxes from the stream to the riparian zone. During periods of high hydrological retention, stream water exports to downstream ecosystem decrease, while stream water chemistry is mostly determined by in-stream processes. Riparian tree phenology can also affect catchment N exports by shaping the temporal pattern of both light and litter inputs into the stream. In spring, light enhances in-stream photoautotrophic N uptake before riparian leaf-out, while riparian leaf litter inputs promote in-stream N mineralization in summer and fall. Finally, we illustrate that the impact of Mediterranean riparian zones on stream hydrology and biogeochemistry increases along the stream continuum, and can ultimately influence catchment N exports to downstream ecosystems. Overall, findings gathered in this review question the well-established idea that riparian zones are efficient N buffers, at least for Mediterranean regions, and stress that an integrated view of upland, riparian, and stream ecosystems is essential for advancing our understanding of catchment hydrology and biogeochemistry

Effects of riparian vegetation removal on nutrient retention in a Mediterranean stream

We examined the effects of riparian vegetation removal on algal dynamics and stream nutrient retention efficiency by comparing NH4-N and PO4-P uptake lengths from a logged and an unlogged reach in Riera Major, a forested Mediterranean stream in northeastern Spain. From June to September 1995, we executed 6 short-term additions of N (as NH4Cl) and P (as Na2HPO4) in a 200-m section to measure nutrient uptake lengths. The study site included 2 clearly differentiated reaches in terms of canopy cover by riparian trees: the first 100 m were completely logged (i.e., the logged reach) and the remaining 100 m were left intact (i.e., the shaded reach). Trees were removed from the banks of the logged reach in the winter previous to our sampling. In the shaded reach, riparian vegetation was dominated by alders (Alnus glutinosa). The study was conducted during summer and fall months when differences in light availability between the 2 reaches were greatest because of forest canopy conditions. Algal biomass and % of stream surface covered by algae were higher in the logged than in the shaded reach, indicating that logging had a stimulatory effect on algae in the stream. Overall, nutrient retention efficiency was higher (i.e., shorter uptake lengths) in the logged than in the shaded reach, especially for PO4-P. Despite a greater increase in PO4-P retention efficiency relative to that of NH4-N following logging, retention efficiency for NH4-N was higher than for PO4-P in both study reaches. The PO4-P mass-transfer coefficient was correlated with primary production in both study reaches, indicating that algal activity plays an important role in controlling PO4-P dynamics in this stream. In contrast, the NH4-N mass-transfer coefficient showed a positive relation-ship only with % of algal coverage in the logged reach, and was not correlated with any algal-related parameter in the shaded reach. The lack of correlation with algal production suggests that mechanisms other than algal activity (i.e., microbial heterotrophic processes or abiotic mechanisms) may also influence NH4-N retention in this stream. Overall, this study shows that logging disturbances in small shaded streams may alter in-stream ecological features that lead to changes in stream nutrient retention efficiency. Moreover, it emphasizes that alteration of the tight linkage between the stream channel and the adjacent riparian zone may directly and indirectly impact biogeochemical processes with implications for stream ecosystem functioning

Biofilm growth and nitrogen uptake responses to increases in nitrate and ammonium availability

Nitrate (NO3 −) and ammonium (NH4 +) are the two major dissolved inorganic nitrogen (DIN) species available in streams. Human activities increase stream DIN concentrations and modify the NO3 −:NH4 + ratio. However, few studies have examined biofilm responses to enrichment of both DIN species. We examined biofilm responses to variation in ambient concentrations and enrichments in either NO3 − or NH4 +. We incubated nutrient diffusing substrata (NDS) bioassays with three treatments (DIN-free, +NO3 − and +NH4 +) in five streams. Biomass-specific uptake rates (U spec ) of NO3 − and NH4 + were then measured using in situ additions of 15N-labeled NO3 − and NH4 +. Biomass (estimated from changes in carbon content) and algal accrual rates, as well as U spec -NO3 − of biofilms in DIN-free treatments varied among the streams in which the NDS had been incubated. Higher ambient DIN concentrations were only correlated with enhanced biofilm growth rates. U spec -NO3 − was one order of magnitude greater and more variable than U spec -NH4 +, however similar relative preference index (RPI) suggested that biofilms did not show a clear preference for either DIN species. Biofilm growth and DIN uptake in DIN-amended NDS (i.e., +NO3 − and +NH4 +) were consistently lower than in DIN-free NDS (i.e., control). Lower values in controls with respect to amended NDS were consistently more pronounced for algal accrual rates and U spec -NO3 − and for the +NH4 + than for the +NO3 − treatments. In particular, enrichment with NH4 + reduced biofilm U spec -NO3 − uptake, which has important implications for N cycling in high NH4 + streams

A round-trip ticket: the importance of release processes for in-stream nutrient spiraling

Most nutrient-spiraling studies have focused on estimates of gross uptake (Ugross), which show that streams take up dissolved inorganic nutrients very efficiently. However, studies based on estimates of net uptake (Unet) emphasize that streams tend to be at biogeochemical steady state (i.e., Unet ≈ 0), at least on a time scale of hours. These findings suggest that streams can be highly reactive ecosystems but remain at short-term biogeochemical steady state if Ugross is counterbalanced by release (R), a process that remains widely unexplored. Here, we propose a novel approach to infer R by comparing Unet and Ugross estimated from ambient and plateau concentrations obtained from standard short-term nutrient additions along a reach. We used this approach to examine the temporal variation of R and its balance with Ugross in 2 streams with contrasting hydrological regime (i.e., perennial vs intermittent) during 2 years. We focused on the spiraling metrics of NH4+ and soluble reactive P (SRP), essential sources of N and P in stream ecosystems. R differed substantially between the 2 streams. The perennial stream had a higher proportion of dates with R > 0 and a 2× higher mean R than the intermittent stream for both nutrients. Despite these differences, the magnitude of R and Ugross tended to be similar for both nutrients within each stream, which lead to Unet ≈ 0 in most cases. A notable exception occurred for SRP in the intermittent stream, where R tended to be higher than Ugross during most of the winter period, probably because of desorption of P from stream sediments. Together, our findings shed light on the contribution of release processes to the dynamics of nutrient spiraling and support the idea that streams can be active ecosystems with high spiraling fluxes while simultaneously approaching short-term biogeochemical steady-state

In-stream net uptake regulates inorganic nitrogen export from catchments under base flow conditions

We aimed to investigate the temporal variation of in‐stream net dissolved inorganic nitrogen (DIN) areal uptake rates (UDIN, in μg N m−2 min−1) and its implications on regulating catchment N export, under base flow conditions. To do so, we estimated UDINfrom longitudinal profiles of ambient DIN concentration (nitrate + ammonium) in two streams on a monthly basis during two hydrological years (n = 45). We found that in‐stream DIN uptake and release did not offset each other (UDIN ≠ 0) in half of the dates, and that UDIN> 0 occurred mostly in autumn. Based on these reach‐scale uptake rates, we performed empirical calculations and model simulations to assess the potential of stream network DIN retention to regulate DIN export from catchments on an annual scale. The empirical approach consisted in up‐scalingUDIN by means of a dynamic stream network analysis that considered temporal and spatial variation of UDIN. The modeling approach consisted in applying different scenarios with the INCA model based on the natural range of empirical UDIN values. Our results showed that the contribution of stream network DIN retention to catchment DIN export increased when calculations accounted for the temporal variation of UDIN. Both approaches suggested that stream network DIN retention can significantly reduce DIN export from headwater catchments under base flow conditions (from 4% to 38%)

Does soil moisture overrule temperature dependence of soil respiration in Mediterranean riparian forests?

Soil respiration (SR) is a major component of ecosystems' carbon cycles and represents the second largest CO2 flux in the terrestrial biosphere. Soil temperature is considered to be the primary abiotic control on SR, whereas soil moisture is the secondary control factor. However, soil moisture can become the dominant control on SR in very wet or dry conditions. Determining the trigger that makes soil moisture as the primary control factor of SR will provide a deeper understanding on how SR changes under the projected future increase in droughts. Specific objectives of this study were (1) to investigate the seasonal variations and the relationship between SR and both soil temperature and moisture in a Mediterranean riparian forest along a groundwater level gradient; (2) to determine soil moisture thresholds at which SR is controlled by soil moisture rather than by temperature; (3) to compare SR responses under different tree species present in a Mediterranean riparian forest (Alnus glutinosa, Populus nigra and Fraxinus excelsior). Results showed that the heterotrophic soil respiration rate, groundwater level and 30 cm integral soil moisture (SM30) decreased significantly from the riverside moving uphill and showed a pronounced seasonality. SR rates showed significant differences between tree species, with higher SR for P. nigra and lower SR for A. glutinosa. The lower threshold of soil moisture was 20 and 17% for heterotrophic and total SR, respectively. Daily mean SR rate was positively correlated with soil temperature when soil moisture exceeded the threshold, with Q10 values ranging from 1.19 to 2.14; nevertheless, SR became decoupled from soil temperature when soil moisture dropped below these thresholds

Experimental evidence reveals impact of drought periods on dissolved organic matter quality and ecosystem metabolism in subalpine streams

Subalpine streams are predicted to experience lower summer discharge following climate change and water extractions. In this study, we aimed to understand how drought periods impact dissolved organic matter (DOM) processing and ecosystem metabolism of subalpine streams. We mimicked a gradient of drought conditions in stream‐side flumes and evaluated implications of drought on DOM composition, gross primary production, and ecosystem respiration. Our experiment demonstrated a production and release of DOM from biofilms and leaf litter decomposition at low discharges, increasing dissolved organic carbon concentrations in stream water by up to 50%. Absorbance and fluorescence properties suggested that the released DOM was labile for microbial degradation. Dissolved organic carbon mass balances revealed a high contribution of internal processes to the carbon budget during low flow conditions. The flumes with low discharge were transient sinks of atmospheric CO2 during the first 2 weeks of drought. After this autotrophic phase, the metabolic balance of these flumes turned heterotrophic, suggesting a nutrient limitation for primary production, while respiration remained high. Overall our experimental findings suggest that droughts in subalpine streams will enhance internal carbon cycling by transiently increasing primary production and more permanently respiration as the drought persists. We propose that the duration of a drought period combined with inorganic nutrient availability are key variables that determine if more carbon is respired in situ or exported downstream

Soil water content drives spatiotemporal patterns of CO2 and N2O emissions from a Mediterranean riparian forest soil

Riparian zones play a fundamental role in regulating the amount of carbon (C) and nitrogen (N) that is exported from catchments. However, C and N removal via soil gaseous pathways can influence local budgets of greenhouse gas (GHG) emissions and contribute to climate change. Over a year, we quantified soil effluxes of carbon dioxide (CO2) and nitrous oxide (N2O) from a Mediterranean riparian forest in order to understand the role of these ecosystems on catchment GHG emissions. In addition, we evaluated the main soil microbial processes that produce GHG (mineralization, nitrification, and denitrification) and how changes in soil properties can modify the GHG production over time and space. Riparian soils emitted larger amounts of CO2 (1.2-10 g C m−2 d−1) than N2O (0.001-0.2 mg N m−2 d−1) to the atmosphere attributed to high respiration and low denitrification rates. Both CO2 and N2O emissions showed a marked (but antagonistic) spatial gradient as a result of variations in soil water content across the riparian zone. Deep groundwater tables fueled large soil CO2 effluxes near the hillslope, while N2O emissions were higher in the wet zones adjacent to the stream channel. However, both CO2 and N2O emissions peaked after spring rewetting events, when optimal conditions of soil water content, temperature, and N availability favor microbial respiration, nitrification, and denitrification. Overall, our results highlight the role of water availability on riparian soil biogeochemistry and GHG emissions and suggest that climate change alterations in hydrologic regimes can affect the microbial processes that produce GHG as well as the contribution of these systems to regional and global biogeochemical cycles

Riparian corridors: A new conceptual framework for assessingt nitrogen buffering across biomes

Anthropogenic activities have more than doubled the amount of reactive nitrogen circulating on Earth, creating excess nutrients across the terrestrial-aquatic gradient. These excess nutrients have caused worldwide eutrophication, fundamentally altering the functioning of freshwater and marine ecosystems. Riparian zones have been recognized to buffer diffuse nitrate pollution, reducing delivery to aquatic ecosystems, but nutrient removal is not their only function in river systems. In this paper, we propose a new conceptual framework to test the capacity of riparian corridors to retain, remove, and transfer nitrogen along the continuum from land to sea under different climatic conditions. Because longitudinal, lateral, and vertical connectivity in riparian corridors influences their functional role in landscapes, we highlight differences in these parameters across biomes. More specifically, we explore how the structure of riparian corridors shapes stream morphology (the river's spine), their multiple functions at the interface between the stream and its catchment (the skin), and their biogeochemical capacity to retain and remove nitrogen (the kidneys). We use the nitrogen cycle as an example because nitrogen pollution is one of the most pressing global environmental issues, influencing directly and indirectly virtually all ecosystems on Earth. As an initial test of the applicability of our interbiome approach, we present synthesis results of gross ammonification and net nitrification from diverse ecosystems

Nitrogen processing and the role of epilithic biofilms downstream of a wastewater treatment plant

We investigated how dissolved inorganic N (DIN) inputs from a wastewater treatment plant (WWTP) effluent are processed biogeochemically by the receiving stream. We examined longitudinal patterns of NH4+ and NO3− concentrations and their 15N signatures along a stream reach downstream of a WWTP. We compared the δ15N signatures of epilithic biofilms with those of DIN to assess the role of stream biofilms in N processing. We analyzed the δ15N signatures of biofilms coating light- and dark-side surfaces of cobbles separately to test whether light constrains functioning of biofilm communities. We sampled during 2 contrasting periods of the year (winter and summer) to explore whether changes in environmental conditions affected N biogeochemical processes. The study reach had a remarkable capacity for transformation and removal of DIN, but the magnitude and relevance of different biogeochemical pathways of N processing differed between seasons. In winter, assimilation and nitrification influenced downstream N fluxes. These processes were spatially segregated at the microhabitat scale, as indicated by a significant difference in the δ15N signature of light- and dark-side biofilms, a result suggesting that nitrification was mostly associated with dark-side biofilms. In summer, N processing was intensified, and denitrification became an important N removal pathway. The δ15N signatures of the light- and dark-side biofilms were similar, a result suggesting less spatial segregation of N cycling processes at this microhabitat scale. Collectively, our results highlight the capacity of WWTP-influenced streams to transform and remove WWTP-derived N inputs and indicate the active role of biofilms in these in-stream processes