Leaf‐litter colonisation and breakdown in relation to stream typology: insights from Mediterranean low‐order streams

1. Scant information is available on leaf breakdown in streams of arid and semiarid regions, including the Mediterranean, where environmental heterogeneity can be high and the relationship between stream characteristics and leaf breakdown is poorly known. We tested the hypotheses that differences in leaf breakdown metrics would be substantially higher between mountain and lowland Mediterranean streams than among streams within each subregion and that variability among streams would be substantially higher in the lowlands, because permanent reaches in the semiarid lowland streams are rare and isolated. 2. We compared leaf breakdown and associated dynamics of nutrients, fungi and invertebrates in low‐order Mediterranean streams draining sub‐humid forests in the Sierra Nevada Mountains and nearby semiarid lowlands of south‐eastern Spain. Streams differed between the two subregions mainly in water ion content, temperature and riparian tree cover. We detected higher environmental heterogeneity among streams within the lowlands compared to the Sierra Nevada mountain range. In the lowlands, breakdown coefficients (k) of alder leaves spanned almost the entire range reported for this species from temperate streams, overlapping with less variable breakdown coefficients in the Sierra Nevada. 3. The high variability of k values among the lowland sites appeared to be caused primarily by variability in the composition and abundance of a few leaf‐consuming invertebrate taxa, particularly the snail Melanopsis praemorsa. Fungal and nutrient dynamics were less variable among sites within each subregion. 4. These results indicate that the critical condition for stream functional assessment of well‐constrained breakdown rates, or related metrics, could be met at reference sites within homogenous bio‐geo‐climatic regions such as the Sierra Nevada. By contrast, in heterogeneous areas such as the semiarid lowland streams, natural variability of breakdown rates can greatly exceed the magnitude of effects expected in response to anthropogenic disturbances.     

Patterns of hydrologic control over stream water total nitrogen to total phosphorus ratios

Many ecologists and biogeochemists explore the interaction of the nitrogen (N) and phosphorus (P) cycles by addressing N:P ratios. While N:P ratios are recognized as broadly important to the composition and functioning of lotic ecosystems, the fundamental controls on stream water N:P ratio variation remains poorly understood. Low N:P ratio (less than 16) streams appear more likely in arid climates than in mesic climates, suggesting possible hydrologic or landscape controls. We explored the importance of watershed hydrology to the variation of total N to total P (TN:TP) ratios in stream water, and whether such variation is characteristically different across watershed classes based on mean annual precipitation and median observed TN:TP ratio. Nonparametric scatter plot analysis was applied to normalized TN:TP ratios and associated discharge (Q) measurements from 57 minimally-impacted watersheds from the contiguous United States. At the seasonal scale, TN:TP ratios showed a negative relationship with Q in semiarid climates and a positive relationship with Q in humid climates. Over storm event scales, TN:TP ratios decline with increasing Q across all watershed classes. The results broadly indicate hydrology is an important driver of TN:TP ratio variation over multiple time scales. We hypothesize that the broad differences across watershed classes are driven by variation in the nature of connectivity (frequency and magnitude of connections) of the landscape to streams. A strong physical control of N:P ratios in stream water is in stark contrast to the biological control of N:P ratios in the oceans, suggesting that application of stoichiometric theory—developed using marine systems—to lotic systems requires a broader consideration of controlling factors.     

Analysis of nitrogen saturation potential in Rocky Mountain tundra and forest: implications for aquatic systems

We employed grass and forest versions of the CENTURY model under a range of N deposition values (0.02–1.60 g N m^–2 y^–1) to explore the possibility that high observed lake and stream N was due to terrestrial N saturation of alpine tundra and subalpine forest in Loch Vale Watershed, Rocky Mountain National Park, Colorado. Model results suggest that N is limiting to subalpine forest productivity, but that excess leachate from alpine tundra is sufficient to account for the current observed stream N. Tundra leachate, combined with N leached from exposed rock surfaces, produce high N loads in aquatic ecosystems above treeline in the Colorado Front Range. A combination of terrestrial leaching, large N inputs from snowmelt, high watershed gradients, rapid hydrologic flushing and lake turnover times, and possibly other nutrient limitations of aquatic organisms constrain high elevation lakes and streams from assimilating even small increases in atmospheric N. CENTURY model simulations further suggest that, while increased N deposition will worsen the situation, nitrogen saturation is an ongoing phenomenon.     

An unexpected source of invertebrate community recovery in intermittent streams from a humid continental climate

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Smog nitrogen and the rapid acidification of forest soil, San Bernardino Mountains, southern California

We report the rapid acidification of forest soils in the San Bernardino Mountains of southern California. After 30 years, soil to a depth of 25 cm has decreased from a pH (measured in 0.01 M CaCl2) of 4.8 to 3.1. At the 50-cm depth, it has changed from a pH of 4.8 to 4.2. We attribute this rapid change in soil reactivity to very high rates of anthropogenic atmospheric nitrogen (N) added to the soil surface (72 kg ha(-1) year(-1)) from wet, dry, and fog deposition under a Mediterranean climate. Our research suggests that a soil textural discontinuity, related to a buried ancient landsurface, contributes to this rapid acidification by controlling the spatial and temporal movement of precipitation into the landsurface. As a result, the depth to which dissolved anthropogenic N as nitrate (NO3) is leached early in the winter wet season is limited to within the top approximately 130 cm of soil where it accumulates and increases soil acidity.     

Factors affecting denitrification in agricultural headwater streams in Northeast Ohio,USA

As a result of increased anthropogenic nitrogen (N) loading in surface waters of agricultural watersheds, there is enhanced interest to understand and quantify N removal mechanisms. Denitrification, an important N removal mechanism in aquatic systems, may contribute to reducing N pollution in agricultural headwater streams. However, the key factors controlling this process in lotic systems remain unclear. The objective of our study was to examine the factors regulating rates of denitrification in the sediments of agricultural headwater streams in the mid-western USA. Denitrification rates were variable among streams and treatments (<0.1–28.0 μg N g AFDM⁻¹ h⁻¹) and on average, were higher than those reported for similar headwater streams. Carbon quantity and quality, and pH had no effect on denitrification, while temperature and nitrate ( ) concentrations had a positive effect on rates of denitrification. Specifically, controlled denitrification following Michaelis-Menten kinetics. We calculated a value of k_m (1.0 mg -N L^-1) that was comparable to other studies in aquatic sediments but was well below the median in-stream concentrations (5.2–17.4 mg -N L⁻¹) observed at the study sites. Despite high rates of denitrification, this removal mechanism is most likely saturated in the agricultural headwater streams we examined, suggesting that these systems are not effective at removing in-stream N. Handling editor: D. Ryder     

Deposition of atmospheric nitrogen compounds in humid tropical Cuba

Acid deposition, a direct effect of gaseous air pollutants, is causing widespread damage to terrestrial and aquatic ecosystems. Further, these pollutants are responsible for the corrosion of building materials and cultural objects, as well as having an impact on human health. In Cuba, main atmospheric deposition of nitrogen compounds varies from approximately 12.0 to 65.0 kg N ha(-1) year(-1) in rural areas. Ammonia and ammonium are the most important elements in Cuba's tropical conditions.     

Inorganic and organic losses of nitrogen from upland regions of Britain: concentrations and fluxes

The nitrogen (N) composition of streams draining eight upland regions of Britain was compared using monthly samples collected between April 1997 and April 1998. Stream samples were analysed for total N (TN), particulate N (PN), nitrate (NO3), ammonium (NH4), and dissolved organic nitrogen (DON). Concentrations of TN were small, generally less than 1.5 mg N l(-1), were dominated by dissolved forms of N, and varied significantly between regions. NO3 accounted for the majority of variability. Concentrations of DON also varied between regions but to a smaller extent than those of NO3. There were considerable variations in TN fluxes between upland regions, which ranged between 3.8 and 16.1 kg N ha(-1) year(-1). The majority of the variation was due to NO3 fluxes, which were largest in regions receiving largest inputs of atmospheric N deposition and ranged between 1.4 and 13.5 kg N ha(-1) year(-1). Fluxes of DON ranged between 1 and 3.5 kg N ha(-1) year( -1), while fluxes of PN were generally less than 0.5 kg N ha(-1) year(-1) , and NH4 fluxes ranged between 0.1 and 0.4 kg N ha(-1) year(-1). NO3 was the dominant fraction (47-84%) of N exported from all upland regions except the Highlands, where DON accounted for 52% of the TN flux. This study has shown that the DON fraction is an important component of the total N transported by upland streams in Britain.     

Ecosystem linkages revealed by experimental lake-derived isotope signal in heathland food webs

Cross-ecosystem movement of nutrients and biomass can have important effects on recipient systems. Emerging aquatic insects are subsidies to terrestrial ecosystems and can influence foodweb interactions in riparian systems. In a 2-year field experiment, we simulated aquatic insect deposition by adding adult midge carcasses (150?g dry mass m^?2 year^?1) to 1-m² heathland plots at a site with low natural midge deposition. We established four levels of midge-addition treatments and measured stable isotopes (??¹³C and ??¹⁵N) in plants and arthropods within each treatment. We used a multiple-source isotope Bayesian mixing model to estimate the terrestrial versus aquatic contribution to the diets of arthropods. Aquatic resources were incorporated into plant, detritivore, and predator biomass. Detritivorous Collembola showed the greatest difference in isotope values (+3??? ??¹⁵N and +4??? ??¹³C) between midge-addition and reference treatments. Isotope values of small spiders followed the same trend of enrichment as Collembola while other arthropods (mites and large spiders) were only enriched after 2?years of midge addition. Although predator diets did not change, they became isotopically enriched via their likely prey (Collembola). Plants also had elevated ??¹⁵N (+1???) in midge-addition treatments. The time required and amount of midge-derived C and N detected varied and depended on trophic position. Midge-derived nutrients were no longer present in arthropod biomass in the year following midge addition. Aquatic insect carcasses can be rapidly incorporated into terrestrial food webs in nearshore habitats, and repeated inputs can be detected at multiple trophic levels, thus highlighting the importance of the detrital pathway for aquatic to terrestrial cross-ecosystem subsidies.     

Cloudwater deposition as a source of fixed nitrogen in a Hawaiian montane forest

Precipitation, dry deposition, and cloud water deposition of fixed nitrogen (nitrate and ammonium ions) were measured on the Island of Hawaii. The first two were small N inputs, averaging 0.6±0.2 and 0.15±0.08 kg N ha-1 yr- 1 respectively. We estimate cloud water deposition to be 8–22 kg inorganic N ha-1 yr-1. If an estimate of organic N is included, it may be as high as 50 kg N ha-1 yr-1. Additionally, cloud water deposition is of hydrological significance as it comprised 37% of the total water input. Cloud water interception was certainly lower in the past, as the developing ecosystem would have had less collection surface area.Cloud water samples that were associated with volcanic haze events were very concentrated in NO3- and constituted 60% of the annual cloud water N deposition. This NO3- probably originated from atmospheric N fixed thermally at the hot lava surface or from an interaction between lava and the ocean. If volcanically influenced samples are excluded from the analysis, the resulting cloud water deposition would have been 4–9 kg N ha-1 yr-1. The Pacific Ocean and Kilauea volcano may have a profound impact on the development of this terrestrial ecosystem, as sources of fixed nitrogen.     

Exchange of gaseous nitrogen compounds between agricultural systems and the atmosphere

Crop and livestock agricultural production systems are important contributors to local, regional and global budgets of NH₃, NO_x (NO + NO₂) and N₂O. Emissions of NH₃ and NO_x (which are biologically and chemically active) into the atmosphere serve to redistribute fixed N to local and regional aquatic and terrestrial ecosystems that may otherwise be disconnected from the sources of the N gases. The emissions of NO_x also contribute to local elevated ozone concentrations while N₂O emissions contribute to global greenhouse gas accumulation and to stratospheric ozone depletion.Ammonia is the major gaseous base in the atmosphere and serves to neutralize about 30% of the hydrogen ions in the atmosphere. Fifty to 75% of the 55 Tg N yr^–1 NH₃ from terrestrial systems is emitted from animal and crop-based agriculture from animal excreta and synthetic fertilizer application. About half of the 50 Tg N yr^–1 of NO_x emitted from the earth surface annually arises from fossil fuel combustion and the remainder from biomass burning and emissions from soil. The NO_x emitted, principally as nitric oxide (NO), reacts rapidly in the atmosphere and in a complex cycle with light, ozone and hydrocarbons, and produces nitric acid and particulate nitrate. These materials can interact with plants and the soil locally or be transported form the site and interact with atmospheric particulate to form aerosols. These salts and aerosols return to fertilize terrestrial and aquatic systems in wet and dry deposition. A small fraction of this N may be biologically converted to N₂O. About 5% of the total atmospheric greenhouse effect is attributed to N₂O from which 70% of the annual global anthropogenic emissions come from animal and crop production.The coupling of increased population with a move of a large sector of the world population to diets that require more energy and N input, will lead to continued increases in anthropogenic input into the global N cycle. This scenario suggests that emissions of NH₃, NO_x and N₂O from agricultural systems will continue to increase and impact global terrestrial and aquatic systems, even those far removed from agricultural production, to an ever growing extent, unless N resources are used more efficiently or food consumption trends change.     

Growth, nitrogen uptake, and metabolism in two semiarid shrubs grown at ambient and elevated atmospheric CO2 concentrations: effects of nitrogen supply and source

The effect of differences in nitrogen (N) availability and source on growth and nitrogen metabolism at different atmospheric CO(2) concentrations in Prosopis glandulosa and Prosopis flexuosa (native to semiarid regions of North and South America, respectively) was examined. Total biomass, allocation, N uptake, and metabolites (e.g., free NO(3)(-), soluble proteins, organic acids) were measured in seedlings grown in controlled environment chambers for 48 d at ambient (350 ppm) and elevated (650 ppm) CO(2) and fertilized with high (8.0 mmol/L) or low (0.8 mmol/L) N (N(level)), supplied at either 1 : 1 or 3 : 1 NO(3)(-) : NH(4)(+) ratios (N(source)). Responses to elevated CO(2) depended on both N(level) and N(source), with the largest effects evident at high N(level). A high NO(3)(-) : NH(4)(+) ratio stimulated growth responses to elevated CO(2) in both species when N was limiting and increased the responses of P. flexuosa at high N(level). Significant differences in N uptake and metabolites were found between species. Seedlings of both species are highly responsive to N availability and will benefit from increases in CO(2), provided that a high proportion of NO(3)- to NH(4)-N is present in the soil solution. This enhancement, in combination with responses that increase N acquisition and increases in water use efficiency typically found at elevated CO(2), may indicate that these semiarid species will be better able to cope with both nutrient and water deficits as CO(2) levels rise.     

森林和沼泽对溪流水化学特征的影响

以小兴安岭北部公别拉河上游为研究区,于2004年7～9月对森林溪流和沼泽溪流水样进行水化学特征对比分析.结果表明,森林和沼泽溪流水化学类型均为重碳酸盐类钙组Ⅰ型水(C^Ca_Ⅰ).森林溪流水的pH、矿化度、总硬度、HCO₃^-、SO₄^2-、Ca²⁺、Mg²⁺、Fe均低于沼泽溪流,而总氮、总磷、Cl^-、K⁺、Na⁺则高于沼泽溪流.森林溪流和沼泽溪流中重金属元素Fe、Mn、Cu、Zn、Cd、Hg和Pb含量较低,均未超过我国Ⅰ类地表水环境质量标准.森林溪流中总氮含量为（0.27±0.04） mg·L^-1、总磷含量为（0.040±0.005） mg·L^-1,明显高于沼泽溪流中总氮含量(（0.21±0.02） mg·L^-1)和总磷含量(（0.025±0.004） mg·L^-1),沼泽湿地对N、P有较强的储存和吸附能力,且对NH₄⁺-N的吸附作用远大于对NO₃^--N的吸附.沼泽溪流中Fe含量为（0.26±0.05） mg·L^-1,显著高于森林溪流Fe含量,沼泽湿地对Fe起到还原释放作用.     

The effects of bankside management on chalk stream invertebrate communities

SUMMARY 1. Communities of aquatic macroinvertebrates and the terrestrial adult phases of aquatic insects were investigated from short stretches of English chalk streams with two different bankside vegetation types: simply structured grazed grass (grazed) and structurally complex herbaceous vegetation with scattered trees (ungrazed). Macroinvertebrates were sampled in spring, summer, autumn and winter 1996–97 from three aquatic habitats: mid-channel gravel, patches of the aquatic macrophyte Ranunculus and marginal emergent macrophytes. The terrestrial adult phases of aquatic insects were sampled in spring, summer and autumn from bankside vegetation.
2. Total macroinvertebrate abundance did not differ between stretches with different bankside vegetation. Taxon richness of mid-channel gravel was, however, significantly higher in ungrazed compared with grazed stretches and Shannon diversity ( H ') of mid-channel gravel and marginal vegetation was significantly higher in ungrazed compared with grazed stretches. Total abundance, taxon richness and Shannon diversity ( H ') of the terrestrial adult phases of aquatic insect were significantly higher from the bankside vegetation of ungrazed compared with grazed stretches.
3. Ordination of communities of aquatic macroinvertebrates and terrestrial adults demonstrated that individual families of both groups were generally more abundant in ungrazed stretches. Many more families were significantly associated with ungrazed stretches than with grazed stretches.
4. This investigation has shown that high structural diversity of bankside vegetation along lowland chalk streams is accompanied at the reach scale by increased diversity of both aquatic macroinvertebrates and the terrestrial adult phases of aquatic insects. The conservation potential of such streams may thus be lowered by management practices that result in the removal or simplification of bankside vegetation along extensive stream stretches.     

Controls over nitric oxide and ammonia emissions from Mojave Desert soils

Emissions of reactive N compounds produced during terrestrial N cycling can be an important N loss pathway from ecosystems. Most measurements of this process focus on NO and N(2)O efflux; however, in alkaline soils such as those in the Mojave Desert, NH(3) production can be an important component of N gas loss. We investigated patterns of NO and NH(3) emissions in the Mojave Desert and identified seasonal changes in temperature, precipitation and spatial heterogeneity in soil nutrients as primary controllers of soil efflux. Across all seasons, NH(3) dominated reactive N gas emissions with fluxes ranging from 0.9 to 10 ng N m(-2) s(-1) as compared to NO fluxes of 0.08-1.9 ng N m(-2) s(-1). Fluxes were higher in April and July than in October; however, a fall precipitation event yielded large increases in both NO and NH(3) efflux. To explore the mechanisms driving field observations, we combined NO and NH(3) soil flux measurements with laboratory manipulations of temperature, water and nutrient conditions. These experiments showed a large transient NH(3) pulse (~70-100 ng N m(-2) s(-1)) following water addition, presumably driven by an increase in soil NH(4) (+) concentrations. This was followed by an increase in NO production, with maximum NO flux rates of 34 ng N m(-2) s(-1). Our study suggests that immediately following water addition NH(3) volatilization proceeds at high rates due to the absence of microbial competition for NH(4) (+); during this period N gas loss is insensitive to changes in temperature and soil nutrients. Subsequently, NO emission increases and rates of both NO and NH(3) emission are sensitive to temperature and nutrient constraints on microbial activity. Addition of labile C reduces gaseous N losses, presumably by increasing microbial immobilization, whereas addition of NO(3) (-) stimulates NO and NH(3) efflux.     

Modeled impacts of farming practices and structural agricultural changes on nitrogen fluxes in the Netherlands

In the Netherlands, nutrient emissions from intensive animal husbandry have contributed to decreased species diversity in (semi) natural terrestrial and aquatic ecosystems, pollution of groundwater, and possibly global warming due to N2O emissions. This paper presents the results of a modelling study presenting the impacts of both structural measures and improved farming practices on major nitrogen (N) fluxes, including NH3 and N2O emission, uptake, leaching, and runoff, in the Netherlands, using input data for the year 2000. Average annual fluxes (Gg N year(-1)) for the year 2000 were estimated at 132 for NH3 emission (160 Gg NH 3 year(-1)), 28 for N2O emission, 50 for N inflow to groundwater, and 15 for N inflow to surface water at a total N input of 1046. At this input, nitrate (NO3) concentrations in groundwater often exceeded the target of 50 mg NO3 l(-1), specifically in well-drained sandy soils. The ammonia (NH3) emissions exceeded emission targets that were set to protect the biodiversity of nonagricultural land. Improved farming practices were calculated to lead to a significant reduction in NH3 emissions to the atmosphere and N leaching and runoff to groundwater and surface water, but these improvements were not enough to reach all the targets set for those fluxes. Only strong structural measures clearly improved the situation. The NH3 emission target of 30 Gg NH3 year(-1), suggested for the year 2030, could not be attained, however, unless pig and poultry farming is completely banned in the Netherlands and all cattle stay almost permanently in low emission stables.     

Land‐use effects on terrestrial consumers through changed size structure of aquatic insects

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Aquatic and terrestrial invertebrate drift in southern Appalachian Mountain streams: implications for trout food resources

1. We characterised aquatic and terrestrial invertebrate drift in six south‐western North Carolina streams and their implications for trout production. Streams of this region typically have low standing stock and production of trout because of low benthic productivity. However, little is known about the contribution of terrestrial invertebrates entering drift, the factors that affect these inputs (including season, diel period and riparian cover type), or the energetic contribution of drift to trout. 2. Eight sites were sampled in streams with four riparian cover types. Drift samples were collected at sunrise, midday and sunset; and in spring, early summer, late summer and autumn. The importance of drift for trout production was assessed using literature estimates of annual benthic production in the southern Appalachians, ecotrophic coefficients and food conversion efficiencies. 3. Abundance and biomass of terrestrial invertebrate inputs and drifting aquatic larvae were typically highest in spring and early summer. Aquatic larval abundances were greater than terrestrial invertebrates during these seasons and terrestrial invertebrate biomass was greater than aquatic larval biomass in the autumn. Drift rates of aquatic larval abundance and biomass were greatest at sunset. Inputs of terrestrial invertebrate biomass were greater than aquatic larvae at midday. Terrestrial invertebrate abundances were highest in streams with open canopies and streams adjacent to pasture with limited forest canopy. 4. We estimate the combination of benthic invertebrate production and terrestrial invertebrate inputs can support 3.3–18.2 g (wet weight) m⁻² year⁻¹ of trout, which is generally lower than values considered productive [10.0–30.0 g (wet weight) m⁻² year⁻¹]. 5. Our results indicate terrestrial invertebrates can be an important energy source for trout in these streams, but trout production is still low. Any management activities that attempt to increase trout production should assess trout food resources and ensure their availability.     

In vitro nitrogen fixation, phosphate solubilization, survival and nutrient release by Azotobacter strains in an aquatic system

In vitro nitrogen fixation, phosphate solubilization, survival and nutrient release by the three strains of Azotobacter (Mac-27, nitrogen fixer; PS-21, phosphate solubilizer and MSX-9, a phytohormone producer) in an aquatic system using three different organic substrates (cow-dung, piggery and poultry waste/excreta) were studied. It was shown that Azotobacter can grow and multiply in an aquatic system when fertilized either with cow-dung (100 mg 1(-1)), poultry (50 mg 1(-1)) or piggery (25 mg 1(-1)) waste/ excreta. Mac-27 and MSX-9 strains of Azotobacter fixed higher quantities of nitrogen (0.03-0.12 mg 1(-1) d(-1)) in aquaria water treated with cow-dung, while with poultry and piggery excreta, the three strains of Azotobacter fixed 0.03-0.09 mg 1(-1)d(-1) of nitrogen. O-PO4 concentration was also high in microbial-inoculated aquaria as compared to the controls which decreased with respect to time. Similarly, a decrease in total nitrogen concentration was also observed in the aquaria water after the maximum levels of fixation had been reached. Concomitant with the decrease in total nitrogen concentration, a simultaneous increase in NO3-N, NO2-N and NH4-N was observed. Maximum phytoplankton population was observed in MSX-9, while the population of zooplankton was maximum in Mac-27 inoculated aquaria. These studies indicated that Azotobacter can be used as a biofertilizer for enhancing nitrogen input and phosphate solubilization in fish ponds. Microbial inoculation along with organic substrate, however, has to be repeated at least at biweekly intervals.