Species traits and environmental conditions govern the relationship between biodiversity effects across trophic levels

Changing environments can have divergent effects on biodiversity–ecosystem function relationships at alternating trophic levels. Freshwater mussels fertilize stream foodwebs through nutrient excretion, and mussel species-specific excretion rates depend on environmental conditions. We asked how differences in mussel diversity in varying environments influence the dynamics between primary producers and consumers. We conducted field experiments manipulating mussel richness under summer (low flow, high temperature) and fall (moderate flow and temperature) conditions, measured nutrient limitation, algal biomass and grazing chironomid abundance, and analyzed the data with non-transgressive overyielding and tripartite biodiversity partitioning analyses. Algal biomass and chironomid abundance were best explained by trait-independent complementarity among mussel species, but the relationship between biodiversity effects across trophic levels (algae and grazers) depended on seasonal differences in mussel species’ trait expression (nutrient excretion and activity level). Both species identity and overall diversity effects were related to the magnitude of nutrient limitation. Our results demonstrate that biodiversity of a resource-provisioning (nutrients and habitat) group of species influences foodweb dynamics and that understanding species traits and environmental context are important for interpreting biodiversity experiments.     

Drought‐induced changes in flow regimes lead to long‐term losses in mussel‐provided ecosystem services

Extreme hydro‐meteorological events such as droughts are becoming more frequent, intense, and persistent. This is particularly true in the south central USA, where rapidly growing urban areas are running out of water and human‐engineered water storage and management are leading to broad‐scale changes in flow regimes. The Kiamichi River in southeastern Oklahoma, USA, has high fish and freshwater mussel biodiversity. However, water from this rural river is desired by multiple urban areas and other entities. Freshwater mussels are large, long‐lived filter feeders that provide important ecosystem services. We ask how observed changes in mussel biomass and community composition resulting from drought‐induced changes in flow regimes might lead to changes in river ecosystem services. We sampled mussel communities in this river over a 20‐year period that included two severe droughts. We then used laboratory‐derived physiological rates and river‐wide estimates of species‐specific mussel biomass to estimate three aggregate ecosystem services provided by mussels over this time period: biofiltration, nutrient recycling (nitrogen and phosphorus), and nutrient storage (nitrogen, phosphorus, and carbon). Mussel populations declined over 60%, and declines were directly linked to drought‐induced changes in flow regimes. All ecosystem services declined over time and mirrored biomass losses. Mussel declines were exacerbated by human water management, which has increased the magnitude and frequency of hydrologic drought in downstream reaches of the river. Freshwater mussels are globally imperiled and declining around the world. Summed across multiple streams and rivers, mussel losses similar to those we document here could have considerable consequences for downstream water quality although lost biofiltration and nutrient retention. While we cannot control the frequency and severity of climatological droughts, water releases from reservoirs could be used to augment stream flows and prevent compounded anthropogenic stressors.     

Understanding how nutrient cycles and freshwater mussels (Unionoida) affect one another

Nutrient loads and nutrient cycling, especially of phosphorus and nitrogen, are among the most important controls on the character of freshwater ecosystems and have been greatly affected by human actions. Despite the widespread importance of nutrients in freshwater ecosystems, the varied linkages between nutrient cycling and freshwater mussel populations have not been thoroughly described. Here, I explore three of these linkages. First, I suggest that nutrient loads are related to the well-being of mussel populations through several mechanisms, probably producing a nonlinear and non-monotonic relationship between nutrient loads and mussel populations. Second, I discuss the ability of mussels to spatially focus nutrients from the overlying water onto the sediments, which has not been fully appreciated, perhaps because nutrient cycling has been viewed chiefly from the viewpoint of the well-mixed water column rather than the patchy sediments. Third, I discuss the ability of mussel populations to accumulate and release nutrients, introducing time lags into nutrient dynamics and stoichiometry (“nutrient capacitance”). Finally, I propose a speculative analysis of the role of freshwater mussels in the nutrient cycles of pristine river systems, which must have been much greater than in modern rivers, with their high nutrient loads and depleted mussel populations.     

Dreissenid mussels enhance nutrient efflux, periphyton quantity and production in the shallow littoral zone of a large lake

Dreissenid mussels are notorious invasive organisms whose establishment is associated with large, ecosystem-scale changes to patterns of productivity in aquatic systems. We investigated how localized impacts of dreissenids affect the potential of littoral substrates to support primary and secondary production using in situ incubations and comparisons of natural mussel-colonized and mussel-free substrates in the littoral zone of a large, shallow lake. We compared dreissenid-colonized and dreissenid-free substrates in terms of nutrient balance, surface area, periphyton loads as well as benthic primary production and respiration rates. Dreissenid-colonized substrates acted as sources of dissolved nutrients to the water column, with mussel mass-specific rates of dissolved phosphorus and ammonia excretion averaging 7.2 ± 5.6 (mean ± SD), and 92.6 ± 64.7 μg/g mussel shell free dry mass/h, respectively. Mussel-colonized substrates also had higher surface area, and supported approximately double the amount of periphyton and organic matter loads compared to mussel-free substrates, as well as having higher rates of primary production and community respiration. We suggest that the localized effects of dreissenids can play an important role in changing whole-ecosystem production patterns, with the extent of dreissenid impacts strongly dependent on lake size and morphometry.     

Tracing Consumer-Derived Nitrogen in Riverine Food Webs

The flux of consumer-derived nutrients is recognized as an important ecosystem process, yet few studies have quantified the impact of these fluxes on freshwater ecosystems. The high abundance of bivalves in both marine and freshwater suggests that bivalves can exert large effects on aquatic food webs. The objective of our study was to determine the importance of unionid mussel-derived nitrogen (MDN) to the food web. We used a stable isotope tracer approach in conjunction with nutrient uptake and excretion experiments. We fed mussels (Lampsilis siliquiodea, n = 249) a ¹⁵N-enriched algal diet and placed them into a N-limited stream for 63 days. Mussel hemolymph was non-lethally sampled over the course of the experiment to measure tissue turnover of δ¹⁵N and excretion experiments were done to model the amount of N mussels provided in comparison to stream N uptake demand. Multiple food web pools were sampled twice prior and five times following the mussel addition to trace the ¹⁵N through the food web. Our mussel excretion rates in comparison to areal uptake demand suggested that mussel excretion can account for 40% of the total N demand in this stream. Our enrichment showed that MDN was entering the food web and supplied up to 19% of the N in specific compartments of the food web near the mussel bed. When scaled to a natural mussel aggregation, our results suggest up to 74% of N in the food web may be mussel-derived. Our results show that N supplied by mussels can be an important nutrient subsidy that provides food web support.     

Ecosystem Processes Performed by Unionid Mussels in Stream Mesocosms: Species Roles and Effects of Abundance

Unionid mussels are a guild of freshwater, sedentary filter-feeders experiencing a global decline in both species richness and abundance. To predict how these losses may impact stream ecosystems we need to quantify the effects of both overall mussel biomass and individual species on ecosystem processes. In this study we begin addressing these fundamental questions by comparing rates of ecosystem processes for two common mussel species, Amblema plicata and Actinonaias ligamentina, across a range of abundance levels and at two trophic states (low and high productivity) in stream mesocosms. At both low and high productivity, community respiration, water column ammonia, nitrate, and phosphorus concentrations, and algal clearance rates were all linearly related to overall mussel biomass. After removing the effects of biomass with ANCOVA, we found few differences between species. In a separate series of experiments, nutrient excretion (phosphorus, ammonia, and molar N:P) and biodeposition rates were only marginally different between species. For the species studied here, functional effects of unionids in streams were similar between species and linearly related to biomass, indicating the potential for strong effects when overall mussel biomass is high and hydrologic residence times are long.     

Ecosystem metabolism and nutrient dynamics in the main channel and backwaters of the Upper Mississippi River

相似文献   

Relationships between ecosystem metabolism, benthic macroinvertebrate densities, and environmental variables in a sub-arctic Alaskan river

Relationships between environmental variables, ecosystem metabolism, and benthos are not well understood in sub-arctic ecosystems. The goal of this study was to investigate environmental drivers of river ecosystem metabolism and macroinvertebrate density in a sub-arctic river. We estimated primary production and respiration rates, sampled benthic macroinvertebrates, and monitored light intensity, discharge rate, and nutrient concentrations in the Chena River, interior Alaska, over two summers. We employed Random Forests models to identify predictor variables for metabolism rates and benthic macroinvertebrate density and biomass, and calculated Spearman correlations between in-stream nutrient levels and metabolism rates. Models indicated that discharge and length of time between high water events were the most important factors measured for predicting metabolism rates. Discharge was the most important variable for predicting benthic macroinvertebrate density and biomass. Primary production rate peaked at intermediate discharge, respiration rate was lowest at the greatest time since last high water event, and benthic macroinvertebrate density was lowest at high discharge rates. The ratio of dissolved inorganic nitrogen to soluble reactive phosphorus ranged from 27:1 to 172:1. We found that discharge plays a key role in regulating stream ecosystem metabolism, but that low phosphorous levels also likely limit primary production in this sub-arctic stream.     

Nutrient loading associated with agriculture land use dampens the importance of consumer‐mediated niche construction

The linkages between biological communities and ecosystem function remain poorly understood along gradients of human‐induced stressors. We examined how resource provisioning (nutrient recycling), mediated by native freshwater mussels, influences the structure and function of benthic communities by combining observational data and a field experiment. We compared the following: (1) elemental and community composition (algal pigments and macroinvertebates) on live mussel shells and on nearby rocks across a gradient of catchment agriculture and (2) experimental colonisation of benthic communities on live vs. sham shells controlling for initial community composition and colonisation duration. We show that in near pristine systems, nutrient heterogeneity mediated by mussels relates to greater biodiversity of communities, which supports the notion that resource heterogeneity can foster biological diversity. However, with increased nutrients from the catchment, the relevance of mussel‐provisioned nutrients was nearly eliminated. While species can persist in disturbed systems, their functional relevance may be diminished or lost.     

Stoichiometry of regenerated nutrients differs between native and invasive freshwater mussels with implications for algal growth

相似文献   

Non-trophic Interactions Control Benthic Producers on Intertidal Flats

The importance of positive effects of ecosystem engineers on associated communities is predicted to increase with environmental stress. However, incorporating such non-trophic interactions into ecological theory is not trivial because facilitation of associated species is conditional on both the type of engineer and the type of abiotic stress. We tested the influence of two allogenic ecosystem engineers (lugworms, Arenicola marina L. and cockles, Cerastoderma edule L.) on the main primary producers (microphytobenthos) of the tidal flats, under different abiotic stresses controlled by reefs of blue mussels (Mytilus edulis L.). We added 25,000 cockles or 2,000 lugworms to 5 × 5 m plots, both in a muddy site with high sedimentation rates located coastward of a mussel bed, and in a sandy site without mussels and characterized by high hydrodynamic stress. After a year, cockles increased algal biomass in the sandy area, but not in the mussel bed site, where high values were measured in all plots. However, lugworms did not affect algal biomass in any of the sites. Field measurements suggest that cockles outweighed negative effects of water currents in the site without mussels by locally increasing sediment stability, whereas mussels overruled the effects of cockles in the wake of the reefs through hydrodynamic stress alleviation and/or biodeposition. Our results suggest that non-trophic interactions by ecosystem engineering bivalves control primary production of intertidal areas, and that the sediment-stabilizing effect of cockles plays a crucial role where the overruling effects of mussel beds are not present.     

The role of the blue mussel Mytilus edulis in the cycling of nutrients in the Oosterschelde estuary (The Netherlands)

The fluxes of particulate and dissolved material between bivalve beds and the water column in the Oosterschelde estuary have been measured in situ with a Benthic Ecosystem Tunnel. On mussel beds uptake of POC, PON and POP was observed. POC and PON fluxes showed a significant positive correlation, and the average C:N ratio of the fluxes was 9.4. There was a high release of phosphate, nitrate, ammonium and silicate from the mussel bed into the water column. The effluxes of dissolved inorganic nitrogen and phosphate showed a significant correlation, with an average N:P ratio of 16.5. A comparison of the in situ measurements with individual nutrient excretion rates showed that excretion by the mussels contributed 31–85% to the total phosphate flux from the mussel bed. Ammonium excretion by the mussels accounted for 17–94% of the ammonium flux from the mussel bed. The mussels did not excrete silicate or nitrate. Mineralization of biodeposition on the mussel bed was probably the main source of the regenerated nutrients.From the in situ observations net budgets of N, P and Si for the mussel bed were calculated. A comparison between the uptake of particulate organic N and the release of dissolved inorganic N (ammonium + nitrate) showed that little N is retained by the mussel bed, and suggested that denitrification is a minor process in the mussel bed sediment. On average, only 2/3 of the particulate organic P, taken up by the mussel bed, was recycled as phosphate. A net Si uptake was observed during phytoplankton blooms, and a net release dominated during autumn. It is concluded that mussel beds increase the mineralization rate of phytoplankton and affect nutrient ratios in the water column. A comparison of N regeneration by mussels in the central part of the Oosterschelde estuary with model estimates of total N remineralization showed that mussels play a major role in the recycling of nitrogen.     

Invasive mussels modify the cycling,storage and distribution of nutrients and carbon in a large lake

相似文献   

Temperature and the metabolic balance of streams

1. It is becoming increasingly clear that fresh waters play a major role in the global C cycle. Stream ecosystem respiration (ER) and gross primary productivity (GPP) exert a significant control on organic carbon fluxes in fluvial networks. However, little is known about how climate change will influence these fluxes. 2. Here, we used a ‘natural experiment’ to demonstrate the role of temperature and nutrient cycling in whole‐system metabolism (ER, GPP and net ecosystem production – NEP), in naturally heated geothermal (5–25 °C) Icelandic streams. 3. We calculated ER and GPP with a new, more accurate method, which enabled us to take into account the additional uncertainties owing to stream spatial heterogeneity in oxygen concentrations within a reach. ER ranged 1–25 g C m^?2 day^?1 and GPP 1–10 g C m^?2 day^?1. The median uncertainties (based on 1 SD) in ER and GPP were 50% and 20%, respectively. 4. Despite extremely low water nutrient concentrations, high metabolic rates in the warm streams were supported by fast cycling rates of nutrients, as revealed from inorganic nutrient (N, P) addition experiments. 5. ER exceeded GPP in all streams (with average GPP/ER = 0.6) and was more strongly related to temperature than GPP, resulting in elevated negative NEP with warming. We show that, as a first approximation based on summer investigations, global stream carbon emission to the atmosphere would nearly double from 0.12 Pg C year^?1 at 13 °C to 0.21 (0.15–0.33) Pg C year^?1 with a 5 °C warming. 6. Compared to previous studies from natural systems (including terrestrial ecosystems), the temperature dependence of stream metabolism was not confounded by latitude or altitude, seasonality, light and nutrient availability, water chemistry, space availability (water transient storage), and water availability. 7. Consequently, stream nutrient processing is likely to increase with warming, protecting downstream ecosystems (rivers, estuaries, coastal marine systems) during the summer low flows from nutrient enrichment, but at the cost of increased CO₂ flux back to the atmosphere.     

Experimental nitrogen and phosphorus enrichment stimulates multiple trophic levels of algal and detrital-based food webs: a global meta-analysis from streams and rivers

Anthropogenic increases in nitrogen (N) and phosphorus (P) concentrations can strongly influence the structure and function of ecosystems. Even though lotic ecosystems receive cumulative inputs of nutrients applied to and deposited on land, no comprehensive assessment has quantified nutrient-enrichment effects within streams and rivers. We conducted a meta-analysis of published studies that experimentally increased concentrations of N and/or P in streams and rivers to examine how enrichment alters ecosystem structure (state: primary producer and consumer biomass and abundance) and function (rate: primary production, leaf breakdown rates, metabolism) at multiple trophic levels (primary producer, microbial heterotroph, primary and secondary consumers, and integrated ecosystem). Our synthesis included 184 studies, 885 experiments, and 3497 biotic responses to nutrient enrichment. We documented widespread increases in organismal biomass and abundance (mean response = +48%) and rates of ecosystem processes (+54%) to enrichment across multiple trophic levels, with no large differences in responses among trophic levels or between autotrophic or heterotrophic food-web pathways. Responses to nutrient enrichment varied with the nutrient added (N, P, or both) depending on rate versus state variable and experiment type, and were greater in flume and whole-stream experiments than in experiments using nutrient-diffusing substrata. Generally, nutrient-enrichment effects also increased with water temperature and light, and decreased under elevated ambient concentrations of inorganic N and/or P. Overall, increased concentrations of N and/or P altered multiple food-web pathways and trophic levels in lotic ecosystems. Our results indicate that preservation or restoration of biodiversity and ecosystem functions of streams and rivers requires management of nutrient inputs and consideration of multiple trophic pathways.     

Aquatic Macrophytes Alter Metabolism and Nutrient Cycling in Lowland Streams

Macrophytes influence the physical, chemical, and biological characteristics of lowland streams, so may be critically important in stream management. We investigated the role of macrophytes in regulating metabolism and nutrient cycling in three lowland, agricultural streams. We measured stream metabolism over the growing season and following experimental macrophyte removal, and used short-term nutrient additions of phosphate (P) and ammonium to assess macrophyte influences on nutrient uptake. Primary production was closely correlated with macrophyte cover across all streams and dates, and decreased greatly with macrophyte removal, whereas ecosystem respiration was not correlated with macrophyte cover and was not altered by macrophyte removal. Phosphate uptake velocity was negatively related to primary production, suggesting that macrophyte activity actually slowed P uptake. Ammonium uptake was not correlated with macrophyte cover or metabolism metrics. Stream nitrate concentrations typically exceeded concentrations of incoming groundwater, suggesting little net nitrate retention in these macrophyte-dominated streams. Phosphorous demand by macrophytes was 10-fold lower than observed uptake rates, indicating that macrophyte P demand was much lower than that of other stream biota. Nitrogen demand by macrophytes was nearly equal to ammonium uptake and was not sufficient to affect the high nitrate flux. These results indicate that macrophytes drive ecosystem metabolism but have limited influence on water column nutrient concentrations because macrophyte demand is much lower than the supply available from the water column. Thus macrophytes in our streams had a large impact on stream trophic state, but offered little potential to influence nutrient removal via management.     

Mussels matter: postlarval dispersal dynamics altered by a spatially complex ecosystem engineer

This study investigated postlarval dispersal of soft-bottom macrofauna at a spatially complex intertidal mudflat comprising patches of bare sediment and an ecosystem engineer, the mussel Mytilus edulis. At each of four sites in Guard Point Cove, Maine, USA, we took core samples and deployed bedload traps in bare sediment and mussel bed habitats to estimate ambient densities, rates of sediment flux, and several measures of postlarval dispersal. Univariate and multivariate nonmetric multidimensional scaling (nMDS) results showed few significant site effects and no habitat×site interactions. In contrast, there were numerous significant habitat effects. Compared to the bare sediment, the mussel bed habitat had: fewer species; higher ambient density and proportional abundance of the oligochaete Tubificoides benedeni (the dominant species in both habitats); lower ambient densities and proportional abundances of major taxa and the nonoligochaetes as a group; and higher sediment flux and relative (i.e., per capita) dispersal of nonoligochaetes. Macrofauna species dispersed in relative proportions that were different from those in the ambient assemblage. Per capita T. benedeni transport rates were low in mussel beds compared to those for nonoligochaetes, consistent with the view that beds represent favorable habitat for oligochaetes. The number of total macrofauna individuals trap⁻¹ day⁻¹ was negatively correlated with ambient density and positively correlated with sediment flux in both habitats, but these relationships were significant only in the mussel bed. The results indicate that altered transport rates of sediment and postlarvae are important mechanisms by which mussels act as ecosystem engineers to modify soft-bottom habitats. Differential transport rates caused by aggregations of mussels and other foundation species must be considered in explanations of spatial pattern in soft-bottom communities.     

Evaluation of the Danish mussel fishery: suggestions for an ecosystem management approach

In Limfjorden, Denmark, an extensive mussel fishery exploits the wild stocks of Mytilus edulis with annual landings of 80,000–100,000 t of mussels. During the last 10 years the impact of mussel dredging on the ecosystem has been studied, including the effect of resuspension of sediment and nutrients and the impoverishment of in- and epi-fauna assemblages. Furthermore, dredging changes the physical structure and complexity of the seabed which affects mussel growth and interactions among zoobenthic species. The blue mussel constitutes the dominant fraction of the zoobenthic suspension feeders, and is important for the transport of material and energy from the pelagic to benthic systems and the control of phytoplankton biomass. In order to evaluate the impact on clearance capacity of a reduction in mussel densities due to mussel dredging, mussel filtration activity measured in situ has been related to the mixing of the water column and the amount of near-bed phytoplankton. Fishery practice for mussel dredging in Limfjorden is discussed in relation to its known impact on the ecosystem and the ecological role of the mussels, and modifications towards an ecosystem management approach and a more sustainable fishery are suggested. The suggested modifications include: a fishery practice where the mussel beds are thinned out when the mussels have attained good quality, and a transplantation practice of mussels from areas with a high mortality to areas with a high growth rate. Both practices intensify the production in a certain area, leaving other areas open for alternative production or for permanent closure for the benefit of the benthic flora and fauna. In addition, other shellfish species represent interesting new resources for fishing or aquaculture. Habitat restoration, such as the relaying of mussel shells from the mussel industry, is another important management tool that should be included in an ecosystem management approach of the mussel fishery. Electronic Publication     

Nutrient recycling by fish in streams along a gradient of agricultural land use

Nutrient recycling is an essential ecosystem process provided by animals. In many aquatic systems, fish have been identified as important in ecosystem nutrient recycling; however, this importance can vary widely between systems. The factors controlling intersystem variation in animal‐mediated nutrient cycling have rarely been examined and as such it remains unclear what impact human landscape changes will have upon these processes. Here we examined rates of nutrient recycling for temperate stream fish assemblages along a gradient of agricultural land use (proportion cropland in the watershed: 1–59%). We quantified nutrient excretion rates of both ammonium–N (NH₄⁺–N) and phosphate (as soluble reactive phosphate: SRP) for fish assemblages at eight streams in southern Ontario, Canada with species‐specific excretion measurements and quantitative assemblage sampling. For both nutrients, total assemblage excretion exhibited a strong positive relationship with riparian cropland. The distance required for fish assemblages to turn over ambient nutrient pools was shorter for cropland systems, indicating that the relative importance of excreted nutrients was higher in these systems. Based on measured uptake rates of NH₄⁺–N in two streams (one higher cropland and one low cropland) and on modeled uptake rates for all streams, the proportion of ecosystem demand that can be satisfied by excretion is generally higher in the more agricultural streams. These patterns appear to be driven largely by disproportionate increases in fish assemblage biomass with increasing stream nutrient concentrations.     

Ecosystem Function in Appalachian Headwater Streams during an Active Invasion by the Hemlock Woolly Adelgid

Forested ecosystems in the southeastern United States are currently undergoing an invasion by the hemlock woolly adelgid (HWA). Previous studies in this area have shown changes to forest structure, decreases in canopy cover, increases in organic matter, and changes to nutrient cycling on the forest floor and soil. Here, we were interested in how the effects of canopy loss and nutrient leakage from terrestrial areas would translate into functional changes in streams draining affected watersheds. We addressed these questions in HWA-infested watersheds at the Coweeta Hydrologic Laboratory in North Carolina. Specifically, we measured stream metabolism (gross primary production and ecosystem respiration) and nitrogen uptake from 2008 to 2011 in five streams across the Coweeta basin. Over the course of our study, we found no change to in-stream nutrient concentrations. While canopy cover decreased annually in these watersheds, this change in light penetration did not translate to higher rates of in-stream primary production during the summer months of our study. We found a trend towards greater heterotrophy within our watersheds, where in-stream respiration accounted for a much larger component of net ecosystem production than GPP. Additionally, increases in rhododendron cover may counteract changes in light and nutrient availability that occurred with hemlock loss. The variability in our metabolic and uptake parameters suggests an actively-infested ecosystem in transition between steady states.