Top-down versus bottom-up limitation in benthic infaunal communities: Direct and indirect effects

Top-down effects of predators and bottom-up effects related to resource availability can be important in determining community structure and function through both direct and indirect processes. Their relative influence may vary among habitats. We examined the effects of nutrient enhancement and predation in southeastern North Carolina to determine relative effects on benthic macrofaunal communities. Short-term nutrient additions and predator exclusions were conducted in two estuaries to examine main and interactive effects on benthic microalgae and infauna. This experimental approach was complemented by comparisons of microalgal biomass, infaunal abundance and composition, predator abundance and predator exclusion among four estuarine systems that varied in background nutrient levels. In the short-term experiments, nutrient enhancement induced increased microalgal biomass but had limited effects on abundances or sizes of infauna. Predator exclusion increased the density of sedentary and near-surface dwelling fauna, but we did not observe interactions between predation and responses to nutrient additions as might be predicted from a simple cascade model. General patterns of abundance were explained to a larger extent by interannual and amongestuary pattems. These results indicate a lack of simple trophic cascade responses for this community over a short time scale and little evidence for local interactive effects. The lack of interactive effects may reflect the opportunistic nature of the dominant infaunal species and potentially different time and spatial scales for the effects of predation and resource controls.     

Multiple stressors in an estuarine system: Effects of nutrients, trace elements, and trophic complexity on benthic photosynthesis and respiration

The effects of nutrients, trace elements, and trophic complexity on benthic photosynthesis and respriation were studied in the Paxtuxent River estuary near St. Leonard, Maryland. Experiments were conducted over three years (1995–1997) in mesocosms containing riverine sediment and water. The experimental design was 2×2×3 factorial with two levels of nutrients (ambient and + nutrients), two of trace elements (ambient and + trace elements) and three of trophic complexity (plankton, plankton + fish, and plankton + fish + benthos). Trace elements included arsenic (As), copper (Cu), and cadmium (Cd). The experiment was conducted three times in 1995 and 1997 and four times in 1996. In 1995 and 1996, sediments were muddy, while in the final year sediments were sandy. In mesocoms with sandy sediments, nutrient additions increased benthic photosynthesis overall, while trace element additions increased benthic photosynthesis in two of three experimental runs. Benthic photosynthesis in these mesocosms appeared to be related to nitrogen loading. Benthic respiration increased in nutrient and trace element amended mesocosms with sandy sediments, apparently in response to higher benthic photosynthesis. Increasing trophic complexity, particularly the presence of benthic organisms, also increased benthic respiration in mesocosms with sandy sediments. There were no effects of nutrient or trace element additions on benthic photosynthesis and respiration when the sediments were muddy. The lack of consistent responses to nutrient additions was surprising given that benthic respiration rates (and presumably nutrient regeneration) were similar in all three years, regardless of sediment type. Muddy, sediments did not mask, the effects of nutrient addition by supplying more nutrients to benthic microalgae than sandy sediments. In 1996, the presence of filter feeding bivalves increased the relative heterotrophy of sediments, measured as production: respiration. Consistent with increased heterotrophy, effluxes of ammonium and soluble reactive phosphorus from sediments were greater in mesocosms containing benthic organisms. Anthropogenically-induced changes in estuaries, such as loading of nutrients and trace elements or reduced trophic complexity, can have important effects on benthic processes and potentially pelagic processes through feedback mechanisms.     

The responses of Patuxent River upper trophic levels to nutrient and trace element induced changes in the lower food web

As a result of human activities, coastal waters can be exposed to multiple stressors that affect primary producers and their interactions with higher trophic levels. Mesocosm experiments were conducted during spring and summer 1996–1998 to investigate the responses of natural populations of primary producers to multiple stressors and the potential for these responses to be transmitted to higher trophic levels (i.e., copepods, bivalves, anemones, and fish). The effects of two stressors, elevated nutrient and trace element loadings, were examined individually and in combination. Nutrient additions had a positive effect on biomass, productivity, and abundance of primary producers (Breitburg et al. 1999; Riedel et al. 2003). Growth or abundance of consumers increased with nutrient additions, but the magnitude of the response was reduced relative to that of their prey. Responses to trace element additions varied seasonally and among taxa. The responses of zooplankton and bivalves to stressor additions were affected by the biomass and changes in species composition of phytoplankton assemblages. The presence of fish predators did not alter zooplankton responses to stressor additions. These results suggest that the extent to which nutrient and trace element effects are transmitted from primary producers to higher trophic levels depends on the capacity of consumers to respond to stressor-induced changes in abundance and species composition of prey, on the absolute abundance of prey, and on the ability of predators to feed on alternative prey. The magnitude of the effects of stressors on estuarine food webs may depend on seasonal variability in species composition of phytoplankton assemblages, whether sensitive species dominate, and whether these species are important prey for secondary consumers. Because spatial and temporal patterns in nutrient and trace element loadings to the estuary can affect species composition of primary producers, it is critically important to examine the magnitude, timing, and spatial relationships of loadings of multiple stressors to coastal waters in order to understand the impacts of these stressors on higher trophic levels.     

Patterns of invertebrate distribution and abundance in the intertidal salt marsh: Causes and questions

The intertidal marsh community comprises both benthic and natant faunal components. The benthic components are primarily small invertebrates residing within or on the soft sediments of the vegetated marsh surface. The natant components include larger, fully aquatic organisms (e.g., fish and shrimp) that inhabit the shallow waters adjacent to the marsh at low tide but interact with the benthic components of the community when the marsh is tidally inundated. In this structurally complex and often expansive intertidal environment, patterns of invertebrate distribution and abundance are not apparent to the casual observer. Benthic core samples taken along an intertidal marsh transect on Sapelo Island, Georgia, USA show that many of the inconspicuous infaunal organisms, which numerically dominate the macrofaunal elements of this soft-substrate community, exhibit zonal distribution patterns along a tidal gradient. Patterns of invertebrate distribution in the intertidal salt marsh are often attributed to the activities of aquatic predators. The results of most predator exclusion experiments have left little doubt that predation/disturbance can be an important determinant of invertebrate abundance in soft-substrate communities; but a growing number of experiments, in both freshwater and marine environments, have produced results that apparently conflict with this, general tenet. Dismissed by some as “failed” experiments, these investigations have exposed our lack of knowledge about the effects of specific predators and the importance of complex interactions which involve more than two trophic levels. Although the importance of predation has been stressed in many recent experimental investigations, there are many other factors that, alone or in combination, may also influence the structure of salt marsh invertebrate assemblages. Included among these are: (1) various density-dependent processes (e.g., adult-larval interactions, agonistic behavior, interspecific competition), (2) selective larval settlement or mortality, (3) the influence of physical factors expressed through habitat preferences, and (4) unpredictable or cyclic physical disturbances. Many questions concerning the spatial and temporal patterns of invertebrate distribution and abundance in the salt marsh are unresolved and remain as challenges to our understanding of soft-substrate community dynamics.     

Experimental studies of predation on polychaetes associated with seagrass beds

Predation on benthic polychaetes associated with seagrass beds was examined in laboratory and field predator inclusion experiments with one fish and three invertebrate predators. Predation had differential effects on polychaete taxa which depended on their microhabitat utilization patterns. The magnitude of predation effects strongly depended on the predator species, with the shrimpPenaeus duorarum having the greatest impact and the pinfishLagodon rhomboides the least. Abundance of spionid polychaetes, which are located near the sediment-water interface, was reduced most often by predation. Nereid and capitellid polychaete abundances were reduced by some predators and not by others. The results emphasize the need for more detailed natural history information on polychaete species in order to improve the interpretation of results from predation experiments.     

The primacy of top-down effects in shallow benthic ecosystems

Individual scientists, scientific organizations, and government agencies have all concluded that eutrophication is among the most detrimental of all human activities in coastal ecosystems; very large amounts of funding have been earmarked to study the negative consequences of nutrient pollution. Most studies of eutrophication have been conducted long after the numbers and diversity of larger marine consumers were dramatically reduced by centuries of intense harvesting. It is now understood that these once abundant predators played pivotal roles in regulating ecosystem structure and function, and that the widespread overharvesting of large consumers can trigger indirect effects that alter species compositions in ways that are very similar to those reported to result from eutrophication. All of this suggests that we should reevaluate whether the many negative effects attributed to eutrophication are actually a result of nutrient additions or whether they may be the result of the indirect effects of dramatically altered coastal food webs. In this essay, we review experimental assessments of the degree to which changes in consumer abundances have indirectly altered the structure of benthic ecosystems in coastal waters, and on the relative importance of top-down and bottom-up effects on coral reefs, rocky shores, and seagrass meadows. We find that the evidence clearly indicates that indirect consumer effects are the primary drivers of coastal benthic ecosystem structure and function.     

Ecological effects from the balance between new and recycled nitrogen in Texas coastal waters

Benthic nutrient recycling is a significant source of dissolved nitrogen for south Texas coastal waters in the region of the Corpus Christi Bay estuary. Studies indicate that 90% of the dissolved nitrogen supply for phytoplankton production is derived from sediments in the upper-estuary, whereas benthic regeneration supplies only 33% of the dissolved nitrogen required for primary production outside the barrier island in coastal waters (15 m depth). In the upper-estuary relationships were observed between fluvial flow, water-column dissolved nitrogen, and phytoplankton productivity. In the middle-estuary relationships were observed between sediment recycling rates and water-column dissolved nitrogen. Beyond the barrier island, relationships were observed between fluvial flow and water-column dissolved nitrogen during high flow periods, while benthic regeneration appeared to be the major nutrient source during low flow periods. We suggest that combined effects from new and recycled nutrient sources buffer south Texas coastal productivity against long periods of low nutrient input from fluvial flow. The comparison of biological responses at several trophic levels to temporal variability in nitrogen recycling and fluvial flow indicated the importance of freshwater nitrogen inputs in stimulating primary production. Freshwater nitrogen inputs also appeared to sustain long-term productivity by replacing nutrients lost from the system by extended reliance upon recycling.     

“A strikingly rich zone”—Nutrient enrichment and secondary production in coastal marine ecosystems

Despite a recent review concluding that there is little or no reason to expect that the production of fish and other animals will increase with nutrient enrichment or eutrophication, there is a variety of evidence that anthropogenic nutrients can stimulate secondary production in marine ecosystems. Unique multiple-year fertilization experiments were carried out over fifty years ago in Scottish sea lochs that showed dramatic increases in the abundance of benthic infauna and greatly enhanced growth of fish as a result of inorganic nitrogen (N) and phosphorus (P) additions. These experiments appear to have provided a good qualitative model for the responses of the Baltic Sea to nutrient enrichment and resulting eutrophication. Historical comparisons by others have shown that the weight of benthic animals per unit area above the halocline in the Baltic is now up to 10 or 20 times greater than it was in the early 1920s and that the total fish biomass in the system may have increased 8 fold between the early part of the 1900s and the 1970s. While there are no similar data for the highly enriched central and southern North Sea, there is convincing evidence that the growth rates of plaice, sole, and other species have increased there since the 1960s or 1970s. Cross-system comparisons have also shown that there are strong correlations between primary production and the production and yield of fish and the standing crop and production of benthic macrofauma in phytoplankton-dominated marine ecosystems. Concerns over the growing nutrient (especially N) enrichment of coastal marine waters are clearly valid and deserve the attention of scientists and managers, but the recent demonizing of N ignores the fact that nutrients are a fundamental requirement for producing biomass. Decisions regarding the amount of N or P that will be allowed to enter marine ecosystems should be made with the full knowledge that there may be tradeoffs between increases in water clarity and dissolved oxygen and the abundance of oysters, clams, fish, and other animals we desire.     

Vertical Distribution and Migration of Decapod Larvae in Relation to Light and Tides in Willapa Bay,Washington

We investigated the occurrence of behaviors that maximize predator avoidance and seaward transport in estuarine decapod zoeae by collecting larvae from discrete depths in a partially mixed estuary, Willapa Bay, Washington, USA, and relating their abundance and vertical distribution to a suite of environmental variables. Abundances of first zoeae of Neotrypaea californiensis and Pinnotheridae were associated with tidal phase, diel phase, and water height. Both taxa were most abundant during ebb tides, and abundances increased with water height, suggesting behaviors that enhanced seaward transport. Additionally, N. californiensis were both shallower and more abundant at night, indicative of behaviors to avoid visual predators. Our results suggest that both tidal transport and predator avoidance are important and sometimes interactive selective forces shaping larval decapod behavior.     

Late Glacial–Holocene record of benthic foraminiferal morphogroups from the eastern Arabian Sea OMZ: Paleoenvironmental implications

The Arabian Sea is characterized today by a well-developed and perennial oxygen minimum zone (OMZ) at mid-water depths. The Indian margin where the OMZ impinges provides sediment records ideal to study past changes in the OMZ intensity and its vertical extent in response to the changes of monsoon-driven primary productivity and intermediate water ventilation. Benthic foraminifera, depending upon their adaptation capabilities to variation in sea floor environment and microhabitat preferences, develop various functional morphologies that can be potentially used in paleoenvironmental reconstruction. In this study, we analysed benthic foraminiferal morphogroups in assemblage records of the last 30 ka in a sediment core collected from the lower OMZ of the Indian margin (off Goa). In total, nine morphogroups within two broadly classified epifaunal and infaunal microhabitat categories are identified. The abundance of morphogroups varies significantly during the late Glacial, Deglacial and Holocene. It appears that monsoon wind driven organic matter flux, and water column ventilation governing the OMZ intensity and sea-bottom oxygen condition, have profound influence on structuring the benthic foraminiferal morphogroups. We found a few morphogroups showing major changes in their abundances during the periods corresponding to the northern hemisphere climatic events. Benthic foraminifera with planoconvex tests are abundant during the cold Heinrich events, when the sea bottom was oxygenated due to a better ventilated, weak OMZ; whereas, those having tapered/cylindrical tests dominate during the last glacial maximum and the Holocene between 5 and 8 ka BP, when the OMZ was intensified and poorly ventilated, leading to oxygen-depleted benthic environment. Characteristically, increased abundance of taxa with milioline tests during the Heinrich 1 further suggests enhanced ventilation attributed probably to the influence of oxygen-rich Antarctic Intermediate Water (AAIW).     

Dynamics of epiphytic photoautotrophs and heterotrophs inZostera marina (eelgrass) microcosms: Responses to nutrient enrichment and grazing

The combined effects of nutrient enrichment and grazing by isopods and amphipods on abundances of seagrass epiphytes were tested inZostera marina L. (eelgrass) microcosms. Using epifluorescence microscopy, densities of epiphytic diatoms, cyanobacteria, heterotrophic flagellates, and heterotrophic bacteria were enumerated after 1 mo and 2 mo of treatment. In general, numbers of diatoms decreased, in the presence of grazers and showed little response to nutrient enrichment, whereas numbers of cyanobacteria increased with nutrient enrichment and showed little response to grazing. Thus, macrofaunal grazing maintained a photoautotrophic community domainated by cyanobacteria, particularly under nutrient enriched conditions. Following 2 mo of treatment, dense macroalgal growth under nutrient-enriched conditins with grazers absent appeared to limit populations of both epiphytic autotrophs. Patterns of abundance of heterotrophic bacteria suggested that the original bacteria population was nutrient limited. Bacteria populations may have been limited by organic carbon supplies at the end of the experiment. Abundances of heterotrophic flagellates and bacteria were strongly correlated on both sampling dates. Results suggest that heterotrophic flagellates might serve as a link between heterotrophic bacterial production and higher trophic levels in seagrass epiphyte food webs.     

Trophic relationships of three sunfishes (Lepomis spp.) in an estuarine bayou

We examined ontogenetic, interspecific, and seasonal trophic patterns among sympatric sunfish species, redspotted sunfish,Lepomis miniatus; redear sunfish,Lepomis microlophus; and bluegill,Lepomis macrochirus, in an estuarine bayou. In particular we studied these feeding patterns in relation to relative abundances of prey from different benthic feeding habitats. All three sunfishes showed ontogenetic divergence in their trophic niches, reflecting different ecomorphological specializations. Small fishes depended on zooplankton, whereas larger fishes of all three species shifted their diets to benthic macrofauna. A potential for trophic resource partitioning was reflected by dietary differences among the three sunfish species. One impalied mechanism for resource partitioning was feeding habitat, as redear sunfish frequently used sediment-associated prey, while bluegill showed greater use of water-column-associated prey, and redspotted sunfish often used SAV-associated prey. However, all three sunfishes apparently used each feeding habitat to some degree; and, trophic differences were more clearly based on prey type than on feeding habitat. Redear sunfish, which can crush hard-shelled prey, exhibited the most distinctive diet. An apparent seasonal shift in feeding habitat occurred in autumn/winter, as indicated by increased overlap between diets and SAV. This shift was facilitated by changes in the relative abundances of several common prey types between benthic habitats. The relative abuandance and use of freshwater and estuarine-derived prey also varied seasonally, suggesting a possible trophic benefit of consistent prey availability in the estuarine bayou.     

Microphytobenthos: The ecological role of the “secret garden” of unvegetated,shallow-water marine habitats. II. role in sediment stability and shallow-water food webs

The microphytobenthos form an important component of all shallow-water ecosystems where enough light reaches the sediment surface to support appreciable primary production. Although less conspicuous than macroalgae or vascular plants, the microphytobenthos can contribute significantly to primary production and can modify habitat characteristics. The microphytobenthos alter sediment properties (e.g., erodibility) both directly, in the extreme forming a mat or scum on the sediment surface, and indirectly by modifying the activities of benthic infauna (e.g., pelletization, burrowing, tube building, and sediment tracking). Carbon dioxide fixed by the microphytobenthos supports higher, grazing trophic levels. These include deposit-feeding and suspension-feeding macrofauna as well as meiofauna and microfauna. Quantitative relations between the feeding and growth rates of macrofauna and the abundance of microphytobenthos and suspended organic matter (i.e., functional responses) are reviewed. Given the current state of knowledge of the direct and indirect interactions involving trophic dynamics, sediment properties, and benthic microalgae, we argue for reductionist studies of particular interactions as distinct entities. This is a prerequisite for the emergence of a comprehensive picture of unvegetated ecosystems and the ability to predict their responses to man’s activities. *** DIRECT SUPPORT *** A01BY074 00005     

Seasonal variability in response of estuarine phytoplankton communities to stress: Linkages between toxic trace elements and nutrient enrichment

We examined individual and interactive effects of two stressors—nutrients (nitrogen [N] and phosphorus [P]) and trace elements (a mix of arsenic [As], copper [Cu], and cadmium [Cd], and in a second experiment also zinc [Zn] and nickel [Ni])—on phytoplankton of the mesohaline Patuxent River, a tributary of Chesapeake Bay. Experiments were conducted in twenty 1-m³ mesocosms. Four mesocosm runs used two levels of nutrient loadings (0.7–1.0 × ambient N loading and enriched to 1.3–1.6 × ambient N loading) crossed with two levels of trace elements (ambient and enriched approximately 2–5 × higher than ambient concentrations) crossed with five progressive levels of ecosystem complexity. To examine seasonal patterns of responses to stressors, data from these experiments were combined with results of a similar experiment conducted during 1996 (Breitburg et al. 1999a). A second mesocosm experiment examined effects of individual and mixed trace elements, both alone and in combination with nutrients, to further examine which nutrient-trace element interactions were important. Nutrients consistently increased phytoplankton productivity and biomass. Most of the increased biomass was created by large centric diatoms, which increased the mean cell size of the phytoplankton community. Trace element additions decreased phytoplankton productivity and biomass, as well as the contribution of large centric diatoms to phytoplankton biomass. When both trace elements and nutrients were added, trace elements reduced nutrient stimulation. Although the magnitude of the response to nutrient additions tended to be somewhat greater in spring, the seasonal patterns of trace element effects, and nutrient-trace element interactions were far more striking with significant responses restricted to spring mesocosm runs. The second experiment indicated that both As and Cu were more inhibitory to phytoplankton in spring than in summer, but As was more inhibitory in the low nutrient treatments and Cu was more inhibitory in the nutrient enrichment treatments. The potential for strong seasonal patterns and high temporal variability in stressor effects and multiple stressor interactiosn will require close attention in the design and interpretation of management-relevant research and monitoring and may indicate the need for seasonally varying management strategies.     

Oyster-sea nettle interdependence and altered control within the Chesapeake Bay ecosystem

Research on the effects of declining abundances of the Eastern oyster (Crassostrea virginica) in Chesapeake Bay and other estuaries has primarily focused on the role of oysters in filtration and nutrient dynamics, and as habitat for fish or fish prey. Oysters also play a key role in providing substrate for the overwintering polyp stage of the scyphomedusa sea nettle,Chrysaora quinquecirrha, which is an important consumer of zooplankton, ctenophores, and icthyoplankton. Temporal trends in sea nettle abundances in visual counts from the dock at Chesapeake Biological Laboratory, trawls conducted in the mesohaline portion of the Patuxent River, and published data from the mainstem Chesapeake Bay indicate that sea nettles declined in the mid 1980s when overfishing and increased disease mortality led to sharp decreases in oyster landings and abundance. Climate trends, previously associated with interannual variation in sea nettle abundances, do not explain the sharp decline. A potentially important consequence of declining sea nettle abundances may be an increase in their ctenophore prey (Mnemiopsis leidyi), and a resultant increase in predation on icthyoplankton and oyster larvae. Increased predation on oyster larvae by ctenophores may inhibit recovery of oyster populations and reinforce the current low abundance of oysters in Chesapeake Bay.     

Natural Disturbance Shapes Benthic Intertidal Macroinvertebrate Communities of High Latitude River Deltas

Unlike lower latitude coastlines, the estuarine nearshore zones of the Alaskan Beaufort Sea are icebound and frozen up to 9 months annually. This annual freezing event represents a dramatic physical disturbance to fauna living within intertidal sediments. The main objectives of this study were to describe the benthic communities of Beaufort Sea deltas, including temporal changes and trophic structure. Understanding benthic invertebrate communities provided a baseline for concurrent research on shorebird foraging ecology at these sites. We found that despite continuous year-to-year episodes of annual freezing, these estuarine deltas are populated by a range of invertebrates that represent both marine and freshwater assemblages. Freshwater organisms like Diptera and Oligochaeta not only survive this extreme event, but a marine invasion of infaunal organisms such as Amphipoda and Polychaeta rapidly recolonizes the delta mudflats following ice ablation. These delta sediments of sand, silt, and clay are fine in structure compared to sediments of other Beaufort Sea coastal intertidal habitats. The relatively depauperate invertebrate community that ultimately develops is composed of marine and freshwater benthic invertebrates. The composition of the infauna also reflects two strategies that make life on Beaufort Sea deltas possible: a migration of marine organisms from deeper lagoons to the intertidal and freshwater biota that survive the 9-month ice-covered period in frozen sediments. Stable isotopic analyses reveal that both infaunal assemblages assimilate marine and terrestrial sources of organic carbon. These results provide some of the first quantitative information on the infaunal food resources of shallow arctic estuarine systems and the long-term persistence of these invertebrate assemblages. Our data help explain the presence of large numbers of shorebirds in these habitats during the brief summer open-water period and their trophic importance to migrating waterfowl and nearshore populations of estuarine fishes that are the basis of subsistence lifestyles by native inhabitants of the Beaufort Sea coast.     

Trophic ecology of the dominant fishes in Elkhorn Slough,California, 1974–1980

Food habits of the dominant fishes collected from 1974 to 1980 at eight locations in Elkhorn Slough, California, and the adjacent ocean were investigated. Epifaunal crustacea was the major prey group identified from stomach contents of more than 2,000 fishes, followed by epifaunal and infaunal worms, and molluscs. Overall, 18 fish species consumed 263 different prey taxa, ranging from 10 taxa to 125 taxa per fish species and including 99 crustacean, 56 polychaete, and 39 molluscan taxa. Mean prey richness was greatest at stations near the ocean and lowest at inshore stations. Detailed dietary data for all prey taxa were summarized as trophic spectra for each fish species. Trophic spectra represented functional groups of prey and were used for comparisons of dietary similarity. Cluster analyses, based on trophic spectra, resulted in four feeding guilds of fishes. Of 18 fish species, seven (Amphistichus argenteus, Leptocottus armatus, Embiotoca jacksoni, Clevelandia ios, Gillichthys mirabilis, Cymatogaster aggregata, andCitharichthys stimaeus) fed principally on epifaunal crustacea. Four species (Pleuronectes vetulus, Platichthys stellatus, Phanerodon furcatus, andMyliobatus californica) consumed mostly molluscs and infaunal worms. Two species (Psettichthys melanostictus andTriakis semifasciata) fed on mobile crustacea, and five species (Hyperprosopon anale, Engraulis mordax, Clupea pallasi, Atherinopsis californiensis, andAtherinops affinis) fed largely on zooplankton and plant material. Our results suggest that high food availability enhances the nursery function of imshore habitats, and emphasize the importance of invertebrate prey populations and the indirect linkage of plant production to the ichthyofaunal assemblarly marine immigrant species that are likely ‘estuarine dependent’.     

Insights into estuarine benthic dissolved organic carbon (DOC) dynamics using δC-DOC values, phospholipid fatty acids and dissolved organic nutrient fluxes

Benthic dissolved organic carbon (DOC) flux rates and changes in DOC isotope ratios, along with nutrient fluxes, phospholipid fatty acids concentration and carbon isotope ratios were measured in productive estuarine sediments over a diel cycle to determine the mechanisms driving benthic-pelagic coupling of DOC. There was uptake of DOC during the dark and efflux during the light at all sites. DOC uptake rates were related to benthic respiration (dark O₂ uptake) and effluxes were coupled to the trophic status (ratio of production to respiration) of the sediments. Highest uptake and efflux rates were observed at two high nutrient concentration sites. The DOC:DON ratio of water column dissolved organic matter (DOM) decreased during the dark and increased during the light indicating preferential uptake and release of carbon rich dissolved organic matter. The calculated carbon isotope ratio of the DOC taken up by the benthos was significantly more depleted than the bulk water column DOC pool, suggesting preferential uptake of selected components of the water column DOC pool. Generally the isotope ratio of the DOC released during the light was more enriched than that taken up during the dark, which suggests that the benthos has the potential to significantly alter the estuarine DOC pool. Uptake and efflux were coupled to respiration and algal grazing/mineralization, therefore increased nutrient loading may shift the composition of the estuarine DOC pool through changes in the magnitude of benthic DOC fluxes. A combination of biological (diel shifts in DOC production and consumption) and abiotic processes (flocculation) appear to be driving the observed benthic DOC dynamics at the study sites. This study was the first to measure carbon isotopic changes in the water column DOC pool due to benthic processes, and shows that the benthos can alter the estuarine DOC pool through diel differences in DOC uptake and efflux.     

Influence of Organic Enrichment and <Emphasis Type="Italic">Spisula subtruncata</Emphasis> (da Costa, 1778) on Oxygen and Nutrient Fluxes in Fine Sand Sediments

The role of labile organic material and macrofaunal activity in benthic respiration and nutrient regeneration have been tested in sublittoral fine sand sediments from the Gulf of Valencia (northwestern Mediterranean Sea). Three experimental setups were made using benthic chambers. One experiment was performed in-situ through the annual cycle in a well-sorted fine sand community. The remaining experiments were carried out with mesocosms under laboratory conditions: one with different concentrations of organic enrichment (mussel meat and concentrated diatoms culture), and the other adding two different densities of the endofaunal bivalve Spisula subtruncata. Biochemical variables in surface sediment and changes in oxygen consumption and nutrient fluxes throughout incubation period were studied in each experiment. In the in situ incubations, dissolved oxygen (DO) fluxes showed a strong correlation with sedimentary biopolymeric fraction of organic carbon. Organic enrichment in the laboratory experiments was responsible for increased benthic respiration. However, sediment response (expressed as DO uptake and dissolved inorganic nitrogen—DIN—release) between oligotrophic and eutrophic conditions was more intense than between eutrophic and hypertrophic conditions. S. subtruncata abundances close to 400 and 850 ind m^?2 also intensified benthic metabolism. DO uptake and DIN production in mesocosms with added fauna were between 60 and 75 % and 65–100 % higher than in the control treatment respectively. The results of these three experiments suggest that the macrobenthic community may increase the benthic respiration by roughly a factor of two in these bottoms, where S. subtruncata is one of the dominant species. Both organic enrichment and macrobenthic community in general, and S. subtruncata in particular, did not seem to have a relevant role in P and Si cycles in these sediments.     

Spatial and Temporal Variability of Benthic Oxygen Demand and Nutrient Regeneration in an Anthropogenically Impacted New England Estuary

Strong benthic–pelagic coupling is an important characteristic of shallow coastal marine ecosystems. Building upon a rich history of benthic metabolism data, we measured oxygen uptake and nutrient fluxes across the sediment–water interface along a gradient of water column primary production in Narragansett Bay, RI (USA). Despite the strong gradients seen in water column production, sediment oxygen demand (SOD) and benthic nutrient fluxes did not exhibit a clear spatial pattern. Some of our sites had been studied in the 1970s and 1980s and thus allowed historical comparison. At these sites, we found that SOD and benthic fluxes have not changed uniformly throughout Narragansett Bay. In the uppermost portion of the bay, the Providence River Estuary, we observed a significant decrease in dissolved inorganic phosphorus fluxes which we attribute to management interventions. At another upper bay site, we observed significant declines in SOD and dissolved inorganic nitrogen fluxes which may be linked to climate-induced decreases in water column primary production and shifts in bloom phenology. In the 1970s, benthic nutrient regeneration supplied 50% to over 200% of the N and P needed to support primary production by phytoplankton. Summer nutrient regeneration in the Providence River Estuary and Upper bay now may only supply some 5–30% of the N and 3–20% of the P phytoplankton demand.