Introduced bivalves in freshwater ecosystems: the impact of Corbicula on organic matter dynamics in a sandy stream

Previous research on Corbicula fluminea (a well-established, non-native bivalve) has clearly shown that this single species impacts ecosystem processes such as nutrient and dissolved organic carbon cycling in the water column of streams. Surprisingly, little was known about how Corbicula might influence similar processes in streambed sediments. Here, we used both laboratory and field experiments to determine how filter- and pedal-feeding by Corbicula impact organic matter dynamics in the sandy streambed (Goose Creek, Virginia). Corbicula consumed significant quantities of organic material in the streambed when conditions favored pedal-feeding but increased buried organic matter stores when filter-feeding promoted deposition of organic matter (by production of feces and pseudofeces). Corbicula contributed significantly to total benthic community respiration (and thus carbon dioxide production), and used pedal-feeding on benthic organic material to grow at a faster rate than that possible by filter-feeding alone. Corbicula should be an important coupler between benthic and pelagic processes because this bivalve uses organic matter from both the water column and the stream sediments. Given the widespread occurrence of this species, we speculate that the introduction of Corbicula may have had major implications for organic matter dynamics in this and many other streams in the United States. Received: 5 October 1998 / Accepted: 6 February 1999     

Effects of flow regime alteration on fluvial habitats and riparian quality in a semiarid Mediterranean basin

The Segura River Basin is one of the most arid and regulated zones in the Mediterranean as well as Europe that includes four hydrologic river types, according to their natural flow regime: main stem rivers, stable streams, seasonal streams and temporary streams. The relationships between flow regime and fluvial and riparian habitats were studied at reference and hydrologically altered sites for each of the four types. Flow regime alteration was assessed using two procedures: (1) an indirect index, derived from variables associated with the main hydrologic pressures in the basin, and (2) reference and altered flow series analyses using the Indicators of Hydrologic Alteration (IHA) and the Indicators of Hydrologic Alteration in Rivers (IAHRIS). Habitats were characterized using the River Habitat Survey (RHS) and its derived Habitat Quality Assessment (HQA) score, whereas riparian condition was assessed using the Riparian Quality Index (RQI) and an inventory of riparian native/exotic species. Flow stability and magnitude were identified as the main hydrologic drivers of the stream habitats in the Segura Basin. Hydrologic alterations were similar to those described in other Mediterranean arid and semiarid areas where dams have reduced flow magnitude and variability and produced the inversion of seasonal patterns. Additionally, the Segura Basin presented two general trends: an increase in flow torrentiality in main stems and an increase in temporality in seasonal and temporary streams. With the indirect alteration index, main stems presented the highest degree of hydrologic alteration, which resulted in larger channel dimensions and less macrophytes and mesohabitats. However, according to the hydrologic analyses, the seasonal streams presented the greatest alteration, which was supported by the numerous changes in habitat features. These changes were associated with a larger proportion of uniform banktop vegetation as well as reduced riparian native plant richness and mesohabitat density. Both stream types presented consequent reductions in habitat and riparian quality as the degree of alteration increased. However, stable streams, those least impacted in the basin, and temporary streams, which are subject to great hydrologic stress in reference conditions, showed fewer changes in physical habitat due to hydrologic alteration. This study clarifies the relationships between hydrologic regime and physical habitat in Mediterranean basins. The hydrologic and habitat indicators that respond to human pressures and the thresholds that imply relevant changes in habitat and riparian quality presented here will play a fundamental role in the use of holistic frameworks when developing environmental flows on a regional scale.     

降水变化和氮沉降影响森林叶根凋落物分解研究进展

全球环境变化通过改变凋落物质量和产量、土壤生物以及非生物因子调控森林凋落物分解,从而对森林生态系统物质和能量循环产生重要的影响。就森林凋落物分解对当前我国面临降水格局变化和大气氮沉降增加的响应进行了回顾和系统的分析,发现降水格局改变如降水减少可能降低凋落物质量从而减缓凋落物分解,而氮沉降增加通常提高凋落物质量从而促进凋落物分解（间接效应）;降水格局改变通过调节土壤含水量和溶解氧含量进而影响微生物参与的分解过程,或通过改变可溶性组分的淋溶量来影响凋落物分解的物理过程,而氮沉降增加主要通过提高外源氮素的有效性从而促进或抑制微生物参与的分解过程（直接效应）。现有研究大多是基于地上凋落物（例如叶凋落物）来理解和量化森林凋落物分解速率与环境因子之间的关系。但目前对降水格局变化及其与大气氮沉降增加的交互作用如何影响森林地上和地下凋落物分解,以及潜在的微生物学机制仍然缺乏统一和清晰的认识。从土壤性质、凋落物质量、微生物群落结构和功能3个方面构建了环境变化对森林地上和地下凋落物分解的概念框架,并进一步阐述未来研究的重点方向：（1）亟需查明地上和地下凋落物分解的驱动机制;（2）探明降水格局变化和氮添加单因子及两因子交互作用对凋落物分解和养分释放的影响及其生物化学调控机理;（3）阐明微生物群落结构和功能对降水格局变化和氮添加单因子及两因子交互的响应机制。以期为深入探讨全球环境变化对森林凋落物分解的影响,以及环境胁迫下森林土壤"碳库"维持机制的解释提供科学依据。     

Riparian plant species loss alters trophic dynamics in detritus-based stream ecosystems

Riparian vegetation is closely connected to stream food webs through input of leaf detritus as a primary energy supply, and therefore, any alteration of plant diversity may influence aquatic ecosystem functioning. We measured leaf litter breakdown rate and associated biological parameters in mesh bags in eight headwater streams bordered either with mixed deciduous forest or with beech forest. The variety of leaf litter types in mixed forest results in higher food quality for large-particle invertebrate detritivores (‘shredders’) than in beech forest, which is dominated by a single leaf species of low quality. Breakdown rate of low quality (oak) leaf litter in coarse mesh bags was lower in beech forest streams than in mixed forest streams, a consequence of lower shredder biomass. In contrast, high quality (alder) leaf litter broke down at similar rates in both stream categories as a result of similar shredder biomass in coarse mesh bags. Microbial breakdown rate of oak and alder leaves, determined in fine mesh bags, did not differ between the stream categories. We found however aquatic hyphomycete species richness on leaf litter to positively co-vary with riparian plant species richness. Fungal species richness may enhance leaf litter breakdown rate through positive effects on resource quality for shredders. A feeding experiment established a positive relationship between fungal species richness per se and leaf litter consumption rate by an amphipod shredder (Gammarus fossarum). Our results show therefore that plant species richness may indirectly govern ecosystem functioning through complex trophic interactions. Integrating microbial diversity and trophic dynamics would considerably improve the prediction of the consequences of species loss.     

Pacific salmon effects on stream ecosystems: a quantitative synthesis

Pacific salmon (Oncorhynchus spp.) disturb sediments and fertilize streams with marine-derived nutrients during their annual spawning runs, leading researchers to classify these fish as ecosystem engineers and providers of resource subsidies. While these processes strongly influence the structure and function of salmon streams, the magnitude of salmon influence varies widely across studies. Here, we use meta-analysis to evaluate potential sources of variability among studies in stream ecosystem responses to salmon. Results obtained from 37 publications that collectively included 79 streams revealed positive, but highly inconsistent, overall effects of salmon on dissolved nutrients, sediment biofilm, macroinvertebrates, resident fish, and isotopic enrichment. Variation in these response variables was commonly influenced by salmon biomass, stream discharge, sediment size, and whether studies used artificial carcass treatments or observed a natural spawning run. Dissolved nutrients were positively related to salmon biomass per unit discharge, and the slope of the relationship for natural runs was five to ten times higher than for carcass additions. Mean effects on ammonium and phosphorus were also greater for natural runs than carcass additions, an effect attributable to excretion by live salmon. In contrast, we observed larger positive effects on benthic macroinvertebrates for carcass additions than for natural runs, likely because disturbance by live salmon was absent. Furthermore, benthic macroinvertebrates and biofilm associated with small sediments (<32 mm) displayed a negative response to salmon while those associated with large sediments (>32 mm) showed a positive response. This comprehensive analysis is the first to quantitatively identify environmental and methodological variables that influence the observed effects of salmon. Identifying sources of variation in salmon–stream interactions is a critical step toward understanding why engineering and subsidy effects vary so dramatically over space and time, and toward developing management strategies that will preserve the ecological integrity of salmon streams. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The effects of alterations in temperature and flow regime on organic carbon dynamics in Mediterranean river networks

It is only recently that freshwaters have been identified as important quantitative components of the carbon (C) cycle at global and regional scales. To date there are no studies that quantitatively predict the effects of alterations in temperature and flow regimes, individually, or in concert, on organic C dynamics in streams. To address this need, we applied a mechanistic model to simulate organic C dynamics in Mediterranean river networks under 27 different scenarios of altered temperature and flow regimes. We predict that the organic C dynamics in freshwaters in the Mediterranean, as well as in other semiarid regions, will be highly sensitive to global climate change owing to major increases in the degree of intermittency as well as in flood frequency and magnitude. Results indicate that flow regime alterations increase C export rates, whereas temperature alterations increase instream metabolism of organic C. However, flow regime alterations exhibit a much greater influence on C dynamics than do changes in the temperature regime. Reservoirs partly counteract the effects of flow extremes on C export rates, and their role in the C dynamics increases with increasing flow variability. The present study is one of the first studies to quantify the complex interactions between the flow and the temperature regime on C dynamics, emphasizing the key role of extreme events such as dry periods and floods, compared with overall trend effects. This information is pivotal in understanding the impact of future climate change on global C dynamics.     

Potential impacts to freshwater ecosystems caused by flow regime alteration under changing climate conditions in Taiwan

Observed increases in the Earth’s surface temperature bring with them associated changes in precipitation and atmospheric moisture that consequentially alter river flow regimes. Climate-induced flow regime changes are examined using the Indicators of Hydrologic Alteration. This article uses observed daily streamflow data to examine the flow regime alteration and how these changes might potentially affect freshwater ecosystems. Flow data from 23 gauging stations throughout Taiwan show that the annual extreme water conditions (1-, 3-, 7-, 30-, 90-day annual minima or maxima) have increased alteration compared to baseline periods (1961–1990). Specifically, more severe flood and drought events occur in the period after 1991 than the period from 1961 to 1990. The frequency and duration of flood and drought events also show increased fluctuation. Flow regimes are currently being altered by stressors that will continue into the foreseeable future and it is also happing elsewhere in the world. Aquatic organisms not only need to defend themselves from anthropogenic damage to the river system, but also face the on-going threat from climate change-induced thermal and flow regime alteration. This article raises this issue so that water resources managers may identify precautionary measures that reduce the cumulative effects of both anthropogenic flow alteration and changing climate conditions.     

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.     

Groundwater influence on alpine stream ecosystems

1. Spatial and temporal variability of relative snow‐melt, glacier‐melt and groundwater contributions to streams play important roles in shaping alpine freshwater ecosystems. Although meltwater (particularly glacier‐fed) streams have received much attention in recent years, the influence of groundwater on alpine freshwater ecosystems remains poorly understood. 2. This study tested the hypotheses that increased groundwater contributions to meltwater‐dominated alpine streams would yield increases in water temperature, channel stability, electrical conductivity and particulate organic matter (POM) and decreases in suspended sediment concentration (SSC). These more favourable habitat conditions were hypothesised to result in increased macroinvertebrate abundance and diversity. 3. Groundwater contributions, physicochemical habitat variables and benthic macroinvertebrates were sampled throughout the 2002 and 2003 summer‐melt seasons in three streams in the French Pyrénées. 4. Increased groundwater contributions were significantly correlated with higher discharge, water temperature, electrical conductivity, POM and channel stability, but lower SSC. 5. Macroinvertebrate total abundance, taxonomic richness, number of Ephemeroptera, Plecoptera and Trichoptera genera, and per cent Plecoptera all increased significantly with greater groundwater contributions to streamflow. However, beta diversity and Trichoptera relative abundance decreased. 6. Abundance of most macroinvertebrate taxa was highest under groundwater‐dominated conditions but a gradient of optimum groundwater preferences was evident across all taxa. Some taxa were found only where groundwater contributions were low (i.e. in predominantly meltwater‐fed streams). 7. This study provides evidence that water source, physicochemical habitat and stream biota are strongly linked. Therefore, an interdisciplinary approach is necessary for future studies aiming to develop conservation strategies or predict the response of alpine river ecosystems to global climate change.     

Dynamics of dissolved and particulate organic carbon in a saline and semiarid stream of southeast Spain (Chicamo stream)

Annual variations in the concentration of dissolved (DOC) and particulate organic carbon (CPOC = Coarse; FPOC = Fine; UPOC = Ultrafine) were studied in a 100 m-reach of the Chicamo stream, an intermittent saline stream in southeast Spain. DOC represented the most important fraction of organic carbon flowing in the Chicamo stream (>98%), with concentrations of about 1.7 mgC l^–1 during most of the year, reaching 2.5 mgC l^–1 in summer. One high flow episode during a rain storm in winter was characterized by a considerably increased concentration of DOC (9.4 mgC l^–1). CPOC was the dominant POC fraction. Positive and significant correlations were found for DOC and discharge, which support the idea of allochthonous inputs due to floods. There was no significant correlation between POC and discharge. No significant correlations were found for DOC or POC with the physico-chemical parameters measured, while a negative significant correlation was found between DOC and temperature. The export of total organic carbon from the drainage basin of the Chicamo stream was low (6.2 × 10^–4 gC m^–2 yr^–1) and typical of streams in arid and semi-arid regions. The results of a Principal Component Analysis defined three different phases. The first consisted of short periods, during which floods provide pulses of allochthonous organic carbon and nutrients, the second a dry phase (summer), defined by biotic interactions, during which the stream could acts as a `sink' of organic matter, and the third and final phase which is characterised by hydrological stability.     

Seasonal dynamics of macroinvertebrate communities in a semiarid saline spring stream with contrasting environmental conditions

Semiarid saline streams are rare aquatic ecosystem types. Their constituent biota is expected to have adapted evolutionarily to strong hydrological variability and salinity stress; however, their ecology is not well known. In this study, we quantify the seasonal changes in the structure of the macroinvertebrate community in the Reventón Rambla (south-eastern Spain), a permanent saline spring stream which is included in a drainage system consisting of ephemeral dry channels (so-called “ramblas”). Seasonal patterns of community structure were studied in two reaches with contrasting environmental regimes using univariate and multivariate statistics. The upstream site showed more stable environmental conditions than the downstream site, and both sites also differed with regard to species richness, and structural and functional group attributes. On a seasonal basis, community dissimilarity was high during periods when both sites were isolated during summer droughts but dissimilarity decreased when both sites were connected through surface flow. Furthermore, the communities tended to show cyclical trajectories in multivariate ordination space. Rather than being related to salinity stress, these patterns seemed to track the hydrological disturbance regime of this rambla system. Spates tended to disrupt communities, while signs of recovery were evident during low-flow periods. Results suggest that salinity fluctuation does not pose a severe abiotic constraint to these adapted macroinvertebrate communities. Their suits of functional properties provide them with the necessary traits to recover quickly from natural disturbance. While human-caused salinization of streams severely impacts communities eventually reducing their recovery potential, our results suggest that communities in natural saline streams may show similar responses to hydrological disturbance as communities from non-saline streams.     

Sentinel systems on the razor's edge: effects of warming on Arctic geothermal stream ecosystems

The Earth is experiencing historically unprecedented rates of warming, with surface temperatures projected to increase by 3–5 °C globally, and up to 7.5 °C in high latitudes, within the next century. Knowledge of how this will affect biological systems is still largely restricted to the lower levels of organization (e.g. species range shifts), rather than at the community, food web or ecosystem level, where responses cannot be predicted from studying single species in isolation. Further, many correlational studies are confounded with time and/or space, whereas experiments have been mostly confined to laboratory microcosms that cannot capture the true complexity of natural ecosystems. We used a ‘natural experiment’ in an attempt to circumvent these shortcomings, by characterizing community structure and trophic interactions in 15 geothermal Icelandic streams ranging in temperature from 5 °C to 45 °C. Even modest temperature increases had dramatic effects across multiple levels of organization, from changes in the mean body size of the top predators, to unimodal responses of species populations, turnover in community composition, and lengthening of food chains. Our results reveal that the rates of warming predicted for the next century have serious implications for the structure and functioning of these fragile ‘sentinel’ ecosystems across multiple levels of organization.     

Evolutionary regime shifts in simulated ecosystems

Catastrophic regime shifts in ecosystems occur when the system is tipped into a new attractor state under some external forcing. Here we consider whether evolutionary adaptations within ecosystems can trigger similar transitions. We use an individual‐based, evolutionary model of interconnected ecosystems to analyze nonlinear changes in global state resulting from local adaptations. Transitions between periods of stability occur when new traits arise that allow exploitation of under‐utilized resources. Subsequent rapid growth of the population carrying the new trait causes abrupt environmental change that drives incumbent species extinct. We call these transitions ‘evolutionary regime shifts’. These internally generated perturbations can result in ecosystem collapse, followed by recovery to an alternate stable state, or occasionally system‐wide extinction. While these disruptions may have a negative impact on ecosystem productivity in individual simulation runs, mean results over many simulations show a trend for increasing ecosystem productivity and stability over time. Feedback between life and the abiotic environment in the model creates a ‘long‐tailed’ distribution of extinction sizes without any external trigger for large extinction events.     

景观对河流生态系统的影响

从景观的视角研究河流生态系统,是目前河流生态学中受到广泛关注且发展迅速的内容之一。流域内多尺度景观强烈影响河流的理化及生物特征,已成为共识,但有关量度、整合景观与河流生态系统二者之间联系的理论体系与方法的建立尚处于起步阶段。对景观组成与空间格局影响河流生态系统的机制与途径进行了系统总结,提出了该领域研究中的主要难题,即如何识别景观中人为因素和自然因素的协变现象,如何衡量多空间尺度景观对河流生态系统的交互影响,如何理解景观阈值的不确定性。为克服当前研究面临的困难与挑战,填补已有知识的不足,提出今后研究的重点方向:河流景观分类系统的改进;更具代表性时空数据的采集;新型景观指标的开发与应用;微观尺度数据与宏观尺度数据的整合等。     

Gradual regime shifts in spatially extended ecosystems

Ecosystem regime shifts are regarded as abrupt global transitions from one stable state to an alternative stable state, induced by slow environmental changes or by global disturbances. Spatially extended ecosystems, however, can also respond to local disturbances by the formation of small domains of the alternative state. Such a response can lead to gradual regime shifts involving front propagation and the coalescence of alternative-state domains. When one of the states is spatially patterned, a multitude of intermediate stable states appears, giving rise to step-like gradual shifts with extended pauses at these states. Using a minimal model, we study gradual state transitions and show that they precede abrupt transitions. We propose indicators to probe gradual regime shifts, and suggest that a combination of abrupt-shift indicators and gradual-shift indicators might be needed to unambiguously identify regime shifts. Our results are particularly relevant to desertification in drylands where transitions to bare soil take place from spotted vegetation, and the degradation process appears to involve step-like events of local vegetation mortality caused by repeated droughts.     

Trophic‐level dependent effects on CO2 emissions from experimental stream ecosystems

Concern over accelerating rates of species invasions and losses have initiated investigations into how local and global changes to predator abundance mediate trophic cascades that influence CO₂ fluxes of aquatic ecosystems. However, to date, no studies have investigated how species additions or losses at other consumer trophic levels influence the CO₂ flux of aquatic ecosystems. In this study, we added a large predatory stonefly, detritivorous stonefly, or grazer tadpole to experimental stream food webs and over a 70‐day period quantified their effects on community composition, leaf litter decomposition, chlorophyll‐a concentrations, and stream CO₂ emissions. In general, streams where the large grazer or large detritivore were added showed no change in total invertebrate biomass, leaf litter loss, chlorophyll‐a concentrations, or stream CO₂ emissions compared with controls; although we did observe a spike in CO₂ emissions in the large grazer treatment following a substantial reduction in chlorophyll‐a concentrations on day 28. However, the large grazer and large detritivore altered the community composition of streams by reducing the densities of other grazer and detritivore taxa, respectively, compared with controls. Conversely, the addition of the large predator created trophic cascades that reduced total invertebrate biomass and increased primary producer biomass. The cascading effects of the predator additions on the food web ultimately led to decreased CO₂ emissions from stream channels by up to 95%. Our results suggest that stream ecosystem processes were more influenced by changes in large predator abundance than large grazer or detritivore abundance, because of a lack of functionally similar large predators. Our study demonstrates that the presence/absence of species with unique functional roles may have consequences for the exchange of CO₂ between the ecosystem and the atmosphere.     

Potential effects of elevated atmospheric carbon dioxide on benthic autotrophs and consumers in stream ecosystems: a test using experimental stream mesocosms

Elevated atmospheric carbon dioxide (eCO₂) has been shown to have a variety of ecosystem‐level effects in terrestrial systems, but few studies have examined how eCO₂ might affect aquatic habitats. This limits broad generalizations about the effects of a changing climate across biomes. To broaden this generalization, we used free air CO₂ enrichment to compare effects of eCO₂ (i.e., double ambient ～720 ppm) relative to ambient CO₂ (aCO₂～360 ppm) on several ecosystem properties and functions in large, outdoor, experimental mesocosms that mimicked shallow sand‐bottom prairie streams. In general, we showed that eCO₂ had strong bottom‐up effects on stream autotrophs, which moved through the food web and indirectly affected consumer trophic levels. These general effects were likely mediated by differential CO₂ limitation between the eCO₂ and aCO₂ treatments. For example, we found that eCO₂ decreased water‐column pH and increased dissolved CO₂ in the mesocosms, reducing CO₂‐limitation at times of intense primary production (PP). At these times, PP of benthic algae was about two times greater in the eCO₂ treatment than aCO₂ treatment. Elevated PP enhanced the rate of carbon assimilation relative to nutrient uptake, which reduced algae quality in the eCO₂ treatment. We predicted that reduced algae quality would negatively affect benthic invertebrates. However, density, biomass and average individual size of benthic invertebrates increased in the eCO₂ treatment relative to aCO₂ treatment. This suggested that total PP was a more important regulator of secondary production than food quality in our experiment. This study broadens generalizations about ecosystem‐level effects of a changing climate by providing some of the first evidence that the global increase in atmospheric CO₂ might affect autotrophs and consumers in small stream ecosystems throughout the southern Great Plains and Gulf Coastal slope of North America.     

Precipitation pulses and carbon fluxes in semiarid and arid ecosystems

In the arid and semiarid regions of North America, discrete precipitation pulses are important triggers for biological activity. The timing and magnitude of these pulses may differentially affect the activity of plants and microbes, combining to influence the C balance of desert ecosystems. Here, we evaluate how a pulse of water influences physiological activity in plants, soils and ecosystems, and how characteristics, such as precipitation pulse size and frequency are important controllers of biological and physical processes in arid land ecosystems. We show that pulse size regulates C balance by determining the temporal duration of activity for different components of the biota. Microbial respiration responds to very small events, but the relationship between pulse size and duration of activity likely saturates at moderate event sizes. Photosynthetic activity of vascular plants generally increases following relatively larger pulses or a series of small pulses. In this case, the duration of physiological activity is an increasing function of pulse size up to events that are infrequent in these hydroclimatological regions. This differential responsiveness of photosynthesis and respiration results in arid ecosystems acting as immediate C sources to the atmosphere following rainfall, with subsequent periods of C accumulation should pulse size be sufficient to initiate vascular plant activity. Using the average pulse size distributions in the North American deserts, a simple modeling exercise shows that net ecosystem exchange of CO₂ is sensitive to changes in the event size distribution representative of wet and dry years. An important regulator of the pulse response is initial soil and canopy conditions and the physical structuring of bare soil and beneath canopy patches on the landscape. Initial condition influences responses to pulses of varying magnitude, while bare soil/beneath canopy patches interact to introduce nonlinearity in the relationship between pulse size and soil water response. Building on this conceptual framework and developing a greater understanding of the complexities of these eco-hydrologic systems may enhance our ability to describe the ecology of desert ecosystems and their sensitivity to global change.