Surface-subsurface interactions in stream ecosystems

Stream ecologists have recently recognized that sediments below streams play an important role in lotic ecosystems. Water flows not only across the surface of stream channels, but also through sediment interstices; consequently, surface and subsurface biogeochemical processes are linked. Recent attempts to understand the influence of subsurface processes on stream ecosystems have tried to resolve the surface-subsurface hydrologic interactions, and to gain knowledge of the ecology of subsurface organisms.     

The boundaries of river systems: the metazoan perspective

1. This overview of metazoans associated with the riparian/groundwater interface focuses on the fauna inhabiting substratum interstices within the stream bed and in alluvial aquifers beneath the floodplain. The objective is to integrate knowledge of habitat conditions and ecology of the interstitial fauna into a broad spatiotemporal perspective of lotic ecosystems. 2. Most aquatic metazoans of terrestrial ancestry, secondarily aquatic forms including insects and water mites (Hydracarina), are largely confined to surface waters (epigean), most of the time penetrating only the superficial interstices of the stream bed. 3. Primary aquatic metazoans include crustaceans and other groups whose entire evolutionary histories took place in water. Some species are epigean, whereas other members of the primary aquatic fauna are true subterranean forms (hypogean ) , residing deep within the stream bed and in alluvial aquifers some distance laterally from the channel. 4. The hypogean/epigean affinities of interstitial animals are reflected in repetitive gradients of species distribution patterns along vertical (depth within the stream bed), longitudinal (riffle/pool), and lateral (across the floodplain) spatial dimensions, as well as along recovery trajectories following floods (temporal dimension). 5. Fluvial dynamics and sediment characteristics interact to determine hydraulic conductivity, oxygen levels, pore space, particle size heterogeneity, organic content and other habitat conditions within the interstitial milieu. 6. Multidimensional environmental gradients occur at various scales across riparian/groundwater boundary zones. The spatiotemporal variability of hydrogeomorphological processes plays an important role in determining habitat heterogeneity, habitat stability, and connectivity between habitat patches, thereby structuring biodiversity patterns across the riverine landscape. 7. The erosive action of flooding maintains a diversity of hydrarch and riparian successional stages in alluvial floodplains. The patchy distribution patterns of interstitial communities at the floodplain scale reflect, in part, the spatial heterogeneity engendered by successional processes. 8. Interstitial metazoans engage in passive and active movements between surface waters and ground waters, between aquatic and riparian habitats, and between different habitat types within the lotic system. Some of these are extensive migrations that involve significant exchange of organic matter and energy between ecosystem compartments. 9. The generally high resilience of lotic ecosystems to disturbance is attributable, in part, to high spatiotemporal heterogeneity. Habitat patches less affected by a particular perturbation may serve as ’refugia ‘; from which survivors recolonize more severely affected areas. Mechanisms of refugium use may also occur within habitats, as, for example, through ontogenetic shifts in microhabitat use. Rigorous investigations of interstitial habitats as refugia should lead to a clearer understanding of the roles of disturbance and stochasticity in lotic ecosystems. 10. Development of realistic ’whole river ‘; food webs have been constrained by the exclusion of interstitial metazoans, which may in fact contribute the majority of energy flow in lotic ecosystems. A related problem is failure to include groundwater/riparian habitats as integral components of alluvial rivers. A conceptual model is presented that integrates groundwater and riparian systems into riverine food webs and that reflects the spatiotemporal complexity of the physical system and connectivity between different components. 11. Interstitial metazoans also serve as ’ecosystem engineers, ‘; by influencing the availability of resouces to other species and by modifying habitat conditions within the sediment. For example, by grazing on biofilm, interstitial animals may markedly stimulate bacterial growth rates and nutrient dynamics. 12. Although there has been a recent surge of interest in the role of interstitial animals in running waters, the knowledge gaps are vast. For example, basic environmental requirements of the majority of groundwater metazoans remain uninvestigated. Virtually nothing is known regarding the role of biotic interactions in structuring faunal distribution patterns across groundwater/riparian boundary zones. Interstitial metazoans may contribute significantly to the total productivity and energy flow of the biosphere, but such data are not available. Nor are sufficient data available to determine the contribution of groundwater animals to estimates of global biodiversity. 13. Effective ecosystem management must include groundwater/riparian ecotones and interstitial metazoans in monitoring and restoration efforts. Evidence suggests that a ’connected ‘; groundwater/riparian system provides natural pollution control, prevents clogging of sediment interstices and maintains high levels of habitat heterogeneity and successional stage diversity. River protection and restoration should maintain or re-establish at least a portion of the natural fluvial dynamics that sustains the ecological integrity of the entire riverine–floodplain–aquifer ecosystem. Keywords: groundwater/riparian ecotones, hyporheic habitat, epigean, hypogean, interstitial fauna, biodiversity, food webs     

Dynamic modeling of nitrogen losses in river networks unravels the coupled effects of hydrological and biogeochemical processes

The importance of lotic systems as sinks for nitrogen inputs is well recognized. A fraction of nitrogen in streamflow is removed to the atmosphere via denitrification with the remainder exported in streamflow as nitrogen loads. At the watershed scale, there is a keen interest in understanding the factors that control the fate of nitrogen throughout the stream channel network, with particular attention to the processes that deliver large nitrogen loads to sensitive coastal ecosystems. We use a dynamic stream transport model to assess biogeochemical (nitrate loadings, concentration, temperature) and hydrological (discharge, depth, velocity) effects on reach-scale denitrification and nitrate removal in the river networks of two watersheds having widely differing levels of nitrate enrichment but nearly identical discharges. Stream denitrification is estimated by regression as a nonlinear function of nitrate concentration, streamflow, and temperature, using more than 300 published measurements from a variety of US streams. These relations are used in the stream transport model to characterize nitrate dynamics related to denitrification at a monthly time scale in the stream reaches of the two watersheds. Results indicate that the nitrate removal efficiency of streams, as measured by the percentage of the stream nitrate flux removed via denitrification per unit length of channel, is appreciably reduced during months with high discharge and nitrate flux and increases during months of low-discharge and flux. Biogeochemical factors, including land use, nitrate inputs, and stream concentrations, are a major control on reach-scale denitrification, evidenced by the disproportionately lower nitrate removal efficiency in streams of the highly nitrate-enriched watershed as compared with that in similarly sized streams in the less nitrate-enriched watershed. Sensitivity analyses reveal that these important biogeochemical factors and physical hydrological factors contribute nearly equally to seasonal and stream-size related variations in the percentage of the stream nitrate flux removed in each watershed.     

Integrative freshwater ecology and biodiversity conservation

Freshwater ecosystems provide goods and services of critical importance to human societies, yet they are among the most heavily altered ecosystems with an overproportional loss of biodiversity. Major threats to freshwater biodiversity include overexploitation, water pollution, fragmentation, destruction or degradation of habitat, and invasions by non-native species. Alterations of natural flow regimes by man-made dams, land-use changes, river impoundments, and water abstraction often have profound impacts on lotic communities. An understanding of the functional interactions and processes in freshwater ecosystems presents a major challenge for scientists, but is crucial for effective and sustainable restoration. Most conservation approaches to date have considered single species or single level strategies. In contrast, the concept of ‘Integrative Freshwater Ecology and Biodiversity Conservation’ (IFEBC) proposed herein addresses the interactions between abiotic and biotic factors on different levels of organization qualitatively and quantitatively. It consequently results in a more holistic understanding of biodiversity functioning and management. Core questions include modeling of the processes in aquatic key habitats and their functionality based on the identification and quantification of factors which control the spatial and temporal distribution of biodiversity and productivity in aquatic ecosystems. The context and importance of research into IFEBC is illustrated using case studies from three major areas of research: (i) aquatic habitat quality and restoration ecology, (ii) the genetic and evolutionary potential of aquatic species, and (iii) the detection of stress and toxic effects in aquatic ecosystems using biomarkers. In conclusion, our understanding of the functioning of aquatic ecosystems and conservation management can greatly benefit from the methodological combination of molecular and ecological tools.     

Parallels and contrasts between intermittently freezing and drying streams: From individual adaptations to biodiversity variation

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Putting the meio- into stream ecology: current findings and future directions for lotic meiofaunal research

• 1 There is a paucity of research on epigean freshwater lotic meiofauna. This may result from a previous emphasis on interstitial (groundwater and hyporheic) meiofauna and/or a reliance on sampling methodologies in lotic systems which are inappropriate for meiofauna.
• 2 Meiofauna contribute much to the diversity of lotic ecosystems. Species lists for seven streams reveal that meiofauna contribute 58–82% of total species numbers, with rotifers and chironomids dominating most systems. The absence of taxonomic keys for most meiofaunal taxa in large areas of the world precludes a wider analysis of their contribution to lotic diversity and an assessment of biogeographical patterns and processes.
• 3 The trophic and functional role of meiofauna in lotic ecosystems is unclear. There are few estimates of meiofaunal production in freshwaters and biomass spectra have produced conflicting results for lotic meiofauna. Present static estimates suggest that the contribution of meiofauna to lotic productivity and biomass is small to moderate, but further studies incorporating a temporal component may provide a more realistic picture of the total contribution of meiofauna to biomass size spectra.
• 4 Meiofauna differ from macroinvertebrates in several respects apart from size and conceptual models for lotic ecosystems should include all metazoans if they are to be truly representative.
• 5 Information on the basic ecology of certain lotic meiofauna (i.e. nematodes, tardigrades, microturbellarians) is urgently required. For those groups whose distributional patterns are better understood (e.g. microcrustaceans), the mechanisms underpinning these patterns should be explored. It is essential that the importance of meiofauna is recognised by lotic ecologists; the only realistic way forward is for greater collaboration among meiofaunal ecologists and taxonomists and other lotic scientists.

     

Seasonal biogeochemical hotspots in the streambed around restoration structures

Rapid exchange of stream water and groundwater in streambeds creates hotspots of biogeochemical cycling of redox-sensitive solutes. Although stream–groundwater interaction can be increased through stream restoration, there are few detailed studies of the increased heterogeneity of water and solute fluxes through the streambed and associated patterns of biogeochemical processes around stream restoration structures. In this study, we examined the seasonal patterns of water and solute fluxes through the streambed around a stream restoration structure to relate patterns of water flux through the streambed to morphology of the channel and biogeochemical processes occurring in the bed. We characterized different biogeochemical zones in the streambed using principal component analysis (PCA) and examined the change in spatial patterns of these zones during different seasons. The PCA results show that two principal components summarized 83% of the variance in the original data set. Streambed pore water was characterized as oxic (indicating production of nitrate), anoxic (indicating sulfate, iron and manganese reduction), or stream-like (indicating there was minimal change in the stream water chemistry in the bed). Regardless of season of the year, anoxic zones were predominantly located upstream of the structure, in a low-velocity pool, and oxic zones were predominantly located downstream of the structure, in a turbulent riffle. We expect structures that span the full channel, are impermeable, and permanent, such as those installed in natural channel design restoration will similarly impact biogeochemical processing in the streambed. The installation of these types of restoration structures may be a way to increase the degree of biogeochemical cycling in stream ecosystems.     

Effects of road salts on groundwater and surface water dynamics of sodium and chloride in an urban restored stream

Road salts are a growing environmental concern in urban watersheds. We examined groundwater (GW) and surface water (SW) dynamics of Na⁺ and Cl^? in Minebank Run (MBR), an urban stream in Maryland, USA. We observed an increasing salinity trend in this restored stream. Current baseflow salinity does not exceed water quality recommendations, but rapid “first flush” storm flow was approximately one-third that of seawater. Comparisons between the upstream and downstream study reaches suggest that a major interstate highway is the primary road salt source. A heavily used road parallels most of MBR and was an additional source to GW concentrations, especially the downstream right bank. A baseflow synoptic survey identified zones of increased salinity. Downstream piezometer wells exhibited increases in salt concentrations and there was evidence that Na⁺ is exchanging Ca²⁺ and Mg²⁺ on soils. SW salt concentrations were generally elevated above GW concentrations. Salinity levels persisted at MBR throughout the year and were above background levels at Bynum Run, a nearby reference stream not bisected by a major highway, suggesting that GW is a long-term reservoir for accumulating road salts. Chronic salinity levels may be high enough to damage vegetation and salinity peaks could impact other biota. Beneficial uses and green infrastructure investments may be at risk from salinity driven degradation. Therefore, road salt may represent an environmental risk that could affect aquatic biota and limit the effectiveness of costly resource management and restoration efforts.     

溪流粗木质残体的生态学研究进展

粗木质残体（CWD）是森林或溪流生态系统中残存的超过一定直径大小的站杆、倒木、枝桠及根系等死木质物的总称,溪流CWD对于溪流生态系统的稳定,水生生物多样性,河槽形态及其变化过程有着重要的作用。对溪流CWD的产生和分类,溪流CWD对于溪流生态系统的稳定,水生生物多样性,河槽形态及其变化过程有着重要的作用。对溪流CWD的产生和分类,溪流CWD贮量,分布和动态,以及溪流CWD的功能和管理分别进行了总结,并指出应尽快在国内开展溪流CWD的研究和管理。     

Critical landscape attributes that influence fish population dynamics in headwater streams

Three landscape attributes are likely to have strong effects on the rate-dependent processes determining fish population dynamics in headwater streams: (1) functional interactions at terrestrial-aquatic ecotones and their influence on temporal and spatial variation in resource supply and predator-prey interactions, (2) large-scale spatial habitat relationships and their effect on resource use and fish movement, and (3) presence of refugia from harsh environmental conditions and their influence on fish survival and emigration/immigration rates. Elucidating how these factors interact over a range of temporal and spatial scales should be a major goal of lotic fish ecologists.     

Decadal-scale Dynamics of Water, Carbon and Nitrogen in a California Chaparral Ecosystem: DAYCENT Modeling Results

The Mediterranean climate, with its characteristic of dry summers and wet winters, influences the hydrologic and microbial processes that control carbon (C) and nitrogen (N) biogeochemical processes in chaparral ecosystems. These biogeochemical processes in turn determine N cycling under chronic N deposition. In order to examine connections between climate and N dynamics, we quantified decadal-scale water, C and N states and fluxes at annual, monthly and daily time steps for a California chaparral ecosystem in the Sierra Nevada using the DAYCENT model. The daily output simulations of net mineralization, stream flow and stream nitrate (NO₃⁻) export were developed for DAYCENT in order to simulate the N dynamics most appropriate for the abrupt rewetting events characteristic of Mediterranean chaparral ecosystems. Overall, the magnitude of annual modeled net N mineralization, soil and plant biomass C and N, nitrate export and gaseous N emission agreed with those of observations. Gaseous N emission was a major N loss pathway in chaparral ecosystems, in which nitric oxide (NO) is the dominant species. The modeled C and N fluxes of net primary production (NPP), N uptake and N mineralization, NO₃⁻ export and gaseous N emission showed both high inter-annual and intra-annual variability. Our simulations also showed dramatic fire effects on NPP, N uptake, N mineralization and gaseous N emission for three years of postfire. The decease in simulated soil organic C and N storages was not dramatic, but lasted a longer time. For the seasonal pattern, the predicted C and N fluxes were greatest during December to March, and lowest in the summer. The model predictions suggested that an increase in the N deposition rate would increase N losses through gaseous N emission and stream N export in the chaparral ecosystems of the Sierra Nevada due to changes in N saturation status. The model predictions could not capture stream NO₃⁻ export during most rewetting events suggesting that a dry-rewetting mechanism representing the increase in N mineralization following soil wetting needs to be incorporated into biogeochemical models of semi-arid ecosystems.     

Manipulation of the Dissolved Organic Carbon Pool in an Agricultural Stream: Responses in Microbial Community Structure, Denitrification, and Assimilatory Nitrogen Uptake

Carbon (C) and nitrogen (N) are strongly coupled across ecosystems due to stoichiometrically balanced assimilatory demand as well as dissimilatory processes such as denitrification. Microorganisms mediate these biogeochemical cycles, but how microbial communities respond to environmental changes, such as dissolved organic carbon (DOC) availability, and how those responses impact coupled biogeochemical cycles in streams is not clear. We enriched a stream in central Indiana with labile DOC for 5?days to investigate coupled C and N cycling. Before, and on day 5 of the enrichment, we examined assimilatory uptake and denitrification using whole-stream ¹⁵N-nitrate tracer additions and short-term nitrate releases. Concurrently, we measured bacterial and denitrifier abundance and community structure. We predicted N assimilation and denitrification would be stimulated by the addition of labile C and would be mediated by increases in bacterial activity, abundance, and a shift in community structure. In response to the twofold increase in DOC concentrations in the water column, N assimilation increased throughout the enrichment. Community respiration doubled during the enrichment and was associated with a change in bacterial community structure (based on terminal restriction fragment length polymorphisms of the 16S rRNA gene). In contrast, there was little response in denitrification or denitrifier community structure, likely because labile C was assimilated by heterotrophic communities on the stream bed prior to reaching denitrifiers within the sediments. Our results suggest that coupling between C and N in streams involves potentially complex interactions with sediment texture and organic matter, microbial community structure, and possibly indirect biogeochemical pathways.     

Use of Shields stress to reconstruct and forecast changes in river metabolism

1. Discharge patterns of streams and rivers may be substantially affected by changes in water management, land use, or climate. Such hydrological alterations are likely to influence biotic processes, including overall ecosystem metabolism (photosynthesis and respiration). One regulator of aquatic ecosystem metabolism directly tied to hydrology is movement of bed sediments. 2. We propose that ecosystem metabolism can be reconstructed or predicted for any suite of hydrological conditions through the use of quantitative relationships between discharge, bed movement and metabolism. We tested this concept on a plains reach of the South Platte River in Colorado. 3. Movement of bed sediments was predicted from river discharge and the Shields stress, a ratio of velocity‐induced stress to sediment grain size. Quantitative relationships were established empirically between metabolic response to bed movement and recovery from bed movement, thus linking metabolism to hydrology. 4. The linkage of metabolism to hydrology allowed us to reconstruct daily photosynthesis and respiration over the 70‐year period for which discharge is known at our study site on the South Platte River. The reconstruction shows major ecological change caused by hydrological manipulation: the river has lost two‐thirds of its photosynthetic potential, and the ratio of photosynthesis to respiration is now much lower than it was prior to 1960. 5. The same approach could be used to anticipate ecological responses to proposed hydrological manipulations, to quantify benefits of hydrological restoration, or to illustrate potential effects of change in climate or land use on flowing‐water ecosystems.     

蜉蝣稚虫形态多样性及其适应性变化

根据蜉蝣稚虫的生活环境,可简单地将其分为静水区种类和流水区种类。静水区种类可生活于水体中、底质表面和底质中三种不同的小栖境,流水区种类可生活于水体中、底质表面和底质缝隙间三类小栖境。形态各异的不同种类蜉蝣稚虫生活在不同的小环境,表明种类演化和形态变化与环境有密切关系。     

A distinct urban biogeochemistry?

Most of the global human population lives in urban areas where biogeochemical cycles are controlled by complex interactions between society and the environment. Urban ecology is an emerging discipline that seeks to understand these interactions, and one of the grand challenges for urban ecologists is to develop models that encompass the myriad influences of people on biogeochemistry. We suggest here that existing models, developed primarily in unmanaged and agricultural ecosystems, work poorly in urban ecosystems because they do not include human biogeochemical controls such as impervious surface proliferation, engineered aqueous flow paths, landscaping choices, and human demographic trends. Incorporating these human controls into biogeochemical models will advance urban ecology and will require enhanced collaborations with engineers and social scientists.     

Implications of seasonal variation in nitrate export from forested ecosystems: a review from the hydrological perspective of ecosystem dynamics

In recent decades, the seasonal variation in NO₃ ^? discharge from forest ecosystems has been used increasingly by ecologists and hydrologists as a diagnostic indicator of the nutrient status of ecosystems. Major factors underlying the seasonal patterns of stream NO₃ ^? concentration include seasonal variations in (1) ecosystem nutrient demand, (2) solute transport capability of the hydrological condition, and (3) in-stream nutrient usage and supply. In this article I show how case studies have helped elucidate the dominant controlling factors by comparing data from Japanese catchments with previously compiled data from studies in North America and Europe. Moreover, I explain the different influences that hydrological and biogeochemical controls exert in regions with rainy (Japan) and dry (northeastern United States and Europe) summers. The seasonal variation in hydrological conditions is the predominant controlling factor in Japanese forests, whereas nutrient demand may predominate in the northeastern United States and Europe. I emphasize the importance of comparing seasonal patterns among regions with differing climates to obtain more universal explanations of seasonal variations in NO₃ ^?-related biogeochemical and hydrological dynamics in a forest ecosystem. In addition, multi-scale investigations will be needed to provide insight into the relative contributions of hillslope biogeochemical effects and in-stream biological activities.     

Predictive models in spatially and temporally variable freshwater systems

Abstract This paper discusses the relationships between scaling and predictability in ecosystems. The logical basis of ecosystem modelling is explored using ideas first developed in complexity theory and analogies with the behaviour of complex adaptive systems. Any ecological model is a scale-dependent entity and both empirical and dynamic models of freshwater systems have their strengths and weaknesses. The logical basis of modelling using functional groups is explored. I conclude that such an approach can be justified and that such models have predictive power. Any predictive model of freshwater systems must take the major scales of external (atmospheric and catchment) forcing into account as well as the scales of key processes in the ecosystem itself. The importance of so-called ‘pink noise’ spectra, which arise both from external forcing and the internal dynamics of dynamic systems, is noted. The key scales of pattern and process in freshwater ecosystems are discussed in relation to the properties of the major functional groups. In order to have predictive power, I conclude that models of freshwater systems must include sediment exchanges and the properties of aquatic macrophytes as well as water column interactions and the pelagic components. When viewed at the scale of functional groups and the major biogeochemical processes, freshwater ecosystems may not be as complex as is often assumed.