Oxygenation of anoxic sediments triggers hatching of zooplankton eggs

Many coastal marine systems have extensive areas with anoxic sediments and it is not well known how these conditions affect the benthic–pelagic coupling. Zooplankton lay their eggs in the pelagic zone, and some sink and lie dormant in the sediment, before hatched zooplankton return to the water column. In this study, we investigated how oxygenation of long-term anoxic sediments affects the hatching frequency of dormant zooplankton eggs. Anoxic sediments from the brackish Baltic Sea were sampled and incubated for 26 days with constant aeration whereby, the sediment surface and the overlying water were turned oxic. Newly hatched rotifers and copepod nauplii (juveniles) were observed after 5 and 8 days, respectively. Approximately 1.5 × 10⁵ nauplii m⁻² emerged from sediment turned oxic compared with 0.02 × 10⁵ m⁻² from controls maintained anoxic. This study demonstrated that re-oxygenation of anoxic sediments activated a large pool of buried zooplankton eggs, strengthening the benthic–pelagic coupling of the system. Modelling of the studied anoxic zone suggested that a substantial part of the pelagic copepod population can derive from hatching of dormant eggs. We suggest that this process should be included in future studies to understand population dynamics and carbon flows in marine pelagic systems.     

Branchial morphology and ion–water transport proteins in Antarctic teleosts with different modes of life

In the Antarctic Ocean salt concentration differs from the bottom to the surface owing to the seasonal forming and melting of sea ice. Antarctic teleosts present different lifestyle from benthic to pelagic. While benthic animals face a constant seawater salinity, benthic–pelagic animals have to face different salt concentration. Branchial morphology and ion–water transport proteins were compared in animals with different lifestyle. The ultrastructure of the gills was investigated by scanning electron microscopy (SEM). Na⁺/K⁺/ATPase, Na⁺/K⁺/Cl⁻ cotransport protein NKCC₁ and Aquaporin 3 (AQP3), were investigated by immunohistochemistry. The immunoreactivity for the ion transporter proteins were more intense in the active benthic–pelagic animals and in the icefishes than in the sluggish benthic ones. Conversely, AQP immunoreactivity was stronger in the animals with sedentary lifestyles. The SEM showed the secondary lamellae in the benthic–pelagic animals more densely packed with the exception of the haemoglobin free teleosts.     

Sediment Resuspension Effects on the Benthic Microbial Loop in Experimental Microcosms

Sediment resuspension induced by anthropogenic disturbance is becoming a major threat of marine coastal ecosystems worldwide. The effects of sediment resuspension on the pelagic domain and on macro- and meiobenthos are well documented in the literature, whereas the effects on the benthic microbial components are nearly neglected. We have investigated the effects of sediment resuspension at two different disturbance levels on benthic bacterial abundance, biomass, and activities and on heterotrophic nanobenthos abundance in experimental microcosms. The results of our experiments pointed out that, independently from the amount of involved energy, sediment resuspension determined a general decrease of all benthic microbial components. The main effects consisted of a decreased abundance of the metabolically active bacterial fraction and of heterotrophic nanobenthos abundance. However, the amount of energy involved in sediment resuspension had differential effects on the structure and functioning of the benthic microbial loop, but only in the short term (i.e., within 36 h). Sediment resuspension had a stimulatory effect on activities of surviving bacterial cells, which, in turn, resulted in increased sediment organic C turnover rates. We hypothesize that such an effect, enhancing nutrient availability, might have relevant consequences on the trophic state of coastal marine ecosystem.     

L’Explosion cambrienne ou l’émergence des écosystèmes modernes

Marine ecosystems with complex trophic structure and dominated by animals started to build up in the Early Cambrian. Fossil evidence from exceptional fossil localities such as the Chengjiang Lagerstätte from South China indicate a high level of biological interactivity (e.g. prey–predator relationships) and the colonization of a wide range of pelagic and benthic niches by predators, scavengers, and detritus and suspension feeders. Swimmers are numerous, but there is no evidence for the extensive occupation of the water column by the Early Cambrian. On the contrary, animal life may have concentrated in hyperbenthic environments, close to the sea bottom. This would have been the initial step towards the colonization of the whole pelagic realm and the building-up of pelagic food chains. A chain of biotic innovations and events seems to have catalyzed both the animal diversification and the build-up of a completely new type of ecosytem, with: (1) the achievement of complex nervous systems, visual organs and motor functions; (2) the introduction of new selective pressure (e.g. predation and feed-back effects); and (3) the colonization of new niches. The role of environmental factors (e.g. oxygen, water chemistry, climate) may have been important in the early stages of metazoan evolution, but was probably negligible in the ecological turnover itself that takes place in the Early Cambrian. Close resemblances between the trophic structure of present-day ecosystems and that of Cambrian ones are confirmed by fossil data and recent mathematical models. This unprecedented increase of interdependence between animal species and trophic levels probably increased the general stability of marine ecosystems, but made them for the first time in their history, highly vulnerable to environmental perturbations. This will largely influence the post-Cambrian evolution of the marine world.     

Global patterns of bacterial beta-diversity in seafloor and seawater ecosystems

Background

Marine microbial communities have been essential contributors to global biomass, nutrient cycling, and biodiversity since the early history of Earth, but so far their community distribution patterns remain unknown in most marine ecosystems.

Methodology/Principal Findings

The synthesis of 9.6 million bacterial V6-rRNA amplicons for 509 samples that span the global ocean''s surface to the deep-sea floor shows that pelagic and benthic communities greatly differ, at all taxonomic levels, and share <10% bacterial types defined at 3% sequence similarity level. Surface and deep water, coastal and open ocean, and anoxic and oxic ecosystems host distinct communities that reflect productivity, land influences and other environmental constraints such as oxygen availability. The high variability of bacterial community composition specific to vent and coastal ecosystems reflects the heterogeneity and dynamic nature of these habitats. Both pelagic and benthic bacterial community distributions correlate with surface water productivity, reflecting the coupling between both realms by particle export. Also, differences in physical mixing may play a fundamental role in the distribution patterns of marine bacteria, as benthic communities showed a higher dissimilarity with increasing distance than pelagic communities.

Conclusions/Significance

This first synthesis of global bacterial distribution across different ecosystems of the World''s oceans shows remarkable horizontal and vertical large-scale patterns in bacterial communities. This opens interesting perspectives for the definition of biogeographical biomes for bacteria of ocean waters and the seabed.     

The importance of benthic–pelagic coupling for marine ecosystem functioning in a changing world

Benthic–pelagic coupling is manifested as the exchange of energy, mass, or nutrients between benthic and pelagic habitats. It plays a prominent role in aquatic ecosystems, and it is crucial to functions from nutrient cycling to energy transfer in food webs. Coastal and estuarine ecosystem structure and function are strongly affected by anthropogenic pressures; however, there are large gaps in our understanding of the responses of inorganic nutrient and organic matter fluxes between benthic habitats and the water column. We illustrate the varied nature of physical and biological benthic–pelagic coupling processes and their potential sensitivity to three anthropogenic pressures – climate change, nutrient loading, and fishing – using the Baltic Sea as a case study and summarize current knowledge on the exchange of inorganic nutrients and organic material between habitats. Traditionally measured benthic–pelagic coupling processes (e.g., nutrient exchange and sedimentation of organic material) are to some extent quantifiable, but the magnitude and variability of biological processes are rarely assessed, preventing quantitative comparisons. Changing oxygen conditions will continue to have widespread effects on the processes that govern inorganic and organic matter exchange among habitats while climate change and nutrient load reductions may have large effects on organic matter sedimentation. Many biological processes (predation, bioturbation) are expected to be sensitive to anthropogenic drivers, but the outcomes for ecosystem function are largely unknown. We emphasize how improved empirical and experimental understanding of benthic–pelagic coupling processes and their variability are necessary to inform models that can quantify the feedbacks among processes and ecosystem responses to a changing world.     

Altered Sea Ice Thickness and Permanence Affects Benthic Ecosystem Functioning in Coastal Antarctica

Antarctic sea ice and the cold waters surrounding the continent are key elements of the global climate system, influencing heat redistribution, oceanic circulation and the absorption of carbon dioxide from the atmosphere. However, the Southern Ocean is predicted to warm by 1–6°C over the next century, altering sea ice extent, thickness and permanence. To better understand the connections between coastal sea ice conditions and the functioning of Antarctica’s unique marine benthic ecosystems, we performed manipulative experiments on the seafloor at two southwestern Ross Sea sites with contrasting sea ice conditions. Benthic systems at both study sites were net heterotrophic during the study period (early November), with primary production most likely limited by light availability rather than nutrients. There was five times more fresh algal detrital material in benthic sediments at the site with the thinner, snow-free, annually formed sea ice, relative to the site with thicker, multiyear sea ice. This elevated quantity and quality of algal detrital matter corresponded with a significantly greater rate of sediment oxygen utilization by the benthos and an altered pathway of nitrogen regeneration (tighter coupling between nitrification and denitrification). Large benthic animals (brittle stars, Ophionotus victoriae) enhanced the efflux of dissolved inorganic nutrients from the sediment to the water column and played a greater role in nutrient regeneration at the site with more food. Although changes in sea ice characteristics in the Western Ross Sea are difficult to predict at present, large benthic organisms can be expected to have an expanded role in mediating the effects of elevated coastal productivity and detritus supply on ecosystem dynamics in this part of Antarctica.     

Nutrient reduction and climate change cause a potential shift from pelagic to benthic pathways in a eutrophic marine ecosystem

The degree to which marine ecosystems may support the pelagic or benthic food chain has been shown to vary across natural and anthropogenic gradients for e.g., in temperature and nutrient availability. Moreover, such external forcing may not only affect the flux of organic matter but could trigger large and abrupt changes, i.e., trophic cascades and ecological regime shifts, which once having occurred may prove potentially irreversible. In this study, we investigate the state and regulatory pathways of the Kattegat; a eutrophied and heavily exploited marine ecosystem, specifically testing for the occurrence of regime shifts and the relative importance of multiple drivers, e.g., climate change, eutrophication and commercial fishing on ecosystem dynamics and trophic pathways. Using multivariate statistics and nonlinear regression on a comprehensive data set, covering abiotic factors and biotic variables across all trophic levels, we here propose a potential regime shift from pelagic to benthic regulatory pathways; a possible first sign of recovery from eutrophication likely triggered by drastic nutrient reductions (involving both nitrogen and phosphorus), in combination with climate‐driven changes in local environmental conditions (e.g., temperature and oxygen concentrations).     

Spatial variability of benthic-pelagic coupling in an estuary ecosystem: consequences for microphytobenthos resuspension phenomenon

The high degree of physical factors in intertidal estuarine ecosystem increases material processing between benthic and pelagic compartments. In these ecosystems, microphytobenthos resuspension is a major phenomenon since its contribution to higher trophic levels can be highly significant. Understanding the sediment and associated microphytobenthos resuspension and its fate in the water column is indispensable for measuring the food available to benthic and pelagic food webs. To identify and hierarchize the physical/biological factors potentially involved in MPB resuspension, the entire intertidal area and surrounding water column of an estuarine ecosystem, the Bay des Veys, was sampled during ebb tide. A wide range of physical parameters (hydrodynamic regime, grain size of the sediment, and suspended matter) and biological parameters (flora and fauna assemblages, chlorophyll) were analyzed to characterize benthic-pelagic coupling at the bay scale. Samples were collected in two contrasted periods, spring and late summer, to assess the impact of forcing variables on benthic-pelagic coupling. A mapping approach using kriging interpolation enabled us to overlay benthic and pelagic maps of physical and biological variables, for both hydrological conditions and trophic indicators. Pelagic Chl a concentration was the best predictor explaining the suspension-feeders spatial distribution. Our results also suggest a perennial spatio-temporal structure of both benthic and pelagic compartments in the ecosystem, at least when the system is not imposed to intense wind, with MPB distribution controlled by both grain size and bathymetry. The benthic component appeared to control the pelagic one via resuspension phenomena at the scale of the bay. Co-inertia analysis showed closer benthic-pelagic coupling between the variables in spring. The higher MPB biomass observed in summer suggests a higher contribution to filter-feeders diets, indicating a higher resuspension effect in summer than in spring, in turn suggesting an important role of macrofauna bioturbation and filter feeding (Cerastoderma edule).     

A trait-based approach to species’ roles in stream ecosystems: climate change,community structure,and material cycling

The sustained decline in habitat quality and community integrity highlights the importance of understanding how communities and environmental variation interactively contribute to ecosystem services. We performed a laboratory experiment manipulating effects of acclimation temperature (5, 15, 25, and 35°C) on resource acquisition, assimilation and subsequent ecosystem services provided by eight freshwater mussel species. Our results suggest that although freshwater mussels are broadly categorized as filter feeders, there are distinct nested functional guilds (thermally tolerant and sensitive) associated with their thermal performance. At 35°C, thermally tolerant species have increased resource assimilation and higher rates of contributed ecosystem services (nutrient excretion, benthic–pelagic coupling). Conversely, thermally sensitive species have decreased assimilation rates and display an array of functional responses including increased/decreased benthic–pelagic coupling and nutrient excretion. Although thermally sensitive species may be in poorer physiological condition at warmer temperatures, their physiological responses can have positive effects on ecosystem services. We extrapolated these results to real mussel beds varying in species composition to address how shifts in community composition coupled with climate change may shift their contributed ecological services. Comparative field data indicate that two co-existing, abundant species with opposing thermal performance (Actinonaias ligamentina, Amblema plicata) differentially dominate community biomass. Additionally, communities varying in the relative proportion of these species differentially influence the magnitude (benthic–pelagic coupling) and quality (N:P excretion) of ecosystem services. As species are increasingly threatened by climate change, greater emphasis should be placed on understanding the contribution of physiological stress to the integrity and functioning of ecosystems. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Ecosystem functioning of two marine food webs in the North‐Western Ionian Sea (Central Mediterranean Sea)

The ecosystem functioning of two marine food webs covering the north‐eastern (Salento) and south‐western (Calabria) sectors of the North‐Western Ionian Sea (NWIS) (Central Mediterranean Sea) was investigated through a food‐web model. Data inputs covered a wide set of ecological information applied to 58 functional groups (FGs). The sum of consumption and the mean predation mortality rate were calculated for benthic, demersal, and pelagic subsystems indicating the predator and prey roles of the FGs. A complex system of energy and biomass exchanges characterized the investigated food webs indicating an important benthic‐pelagic coupling. In the food webs of both areas, the regulation of flows between the benthic‐pelagic coupling seems to occur through the benthopelagic shrimps and the small pelagics due to their wasp‐waist control role. Differences were observed concerning the top predators. Odontocetes play this keystone role in the Salento food web. Anglers, bathyal squids, and sharks assume this functional role in Calabria. The geomorphology and hydrography in the NWIS could affect the biomass and energy exchanges in this coupling. The higher flows of consumption of the benthic system observed in the Calabria food web could be influenced by a widespread presence of canyons along the continental edge which increase the benthic productivity. In contrast, the flows of consumption in the Salento food web seem to be driven by the planktonic productivity supporting the pelagic, benthopelagic, and demersal compartments. This condition could be favored by the large extension of the shelf break zone. The food‐web models realized for the NWIS represent ideal platforms for the development of analysis with dynamic simulations. The comparative analysis of the two food webs by means of the FGs and their functional traits allowed the general pattern of ecosystem structure and functioning in the NWIS to be identified, making it an interesting approach to investigate the marine ecosystem.     

Multiple Interception Pathways for Resource Utilisation and Increased Ecosystem Resilience

Natural ecosystems consist of multiple pathways for resource capture and energy flow. As landscapes become impacted, the diversity of these pathways is reduced and ecosystem health suffers. Efficient resource processing is essential for successful ecosystem restoration and yet little information is available relating the two issues. Disturbances to terrestrial, marine, and freshwater environments have decreased the complexity of such ecosystems thus reducing their resilience to increases in resource loading from anthropogenic sources. The effects of a reduction in the number of pathways are observed in terminal water bodies where increased levels of nutrients are not successfully intercepted beforehand, and conditions often favour a single phytoplankton pathway, resulting in algal blooms. If a suitable number of pathways is restored, significant amounts of resources may be retained, reducing the impacts on downstream ecosystems. The restoration of functional diversity is likely to provide process redundancy and therefore improve ecosystem resilience to further disturbance. This concept provides a more holistic approach to management strategies that may enhance resource interception, while restoring habitat and biological diversity.     

Habitat coupling in lake ecosystems

Lakes are complex ecosystems composed of distinct habitats coupled by biological, physical and chemical processes. While the ecological and evolutionary characteristics of aquatic organisms reflect habitat coupling in lakes, aquatic ecology has largely studied pelagic, benthic and riparian habitats in isolation from each other. Here, we summarize several ecological and evolutionary patterns that highlight the importance of habitat coupling and discuss their implications for understanding ecosystem processes in lakes. We pay special attention to fishes because they play particularly important roles as habitat couplers as a result of their high mobility and flexible foraging tactics that lead to inter-habitat omnivory. Habitat coupling has important consequences for nutrient cycling, predator-prey interactions, and food web structure and stability. For example, nutrient excretion by benthivorous consumers can account for a substantial fraction of inputs to pelagic nutrient cycles. Benthic resources also subsidize carnivore populations that have important predatory effects on plankton communities. These benthic subsidies stabilize population dynamics of pelagic carnivores and intensify the strength of their interactions with planktonic food webs. Furthermore, anthropogenic disturbances such as eutrophication, habitat modification, and exotic species introductions may severely alter habitat connections and, therefore, the fundamental flows of nutrients and energy in lake ecosystems.     

The importance of physical and biotic scaling to the experimental simulation of a coastal marine ecosystem

A marine microcosm, consisting of a pelagic phase interacting with a benthic phase, is described. Variations in water turnover, turbulence, incident radiation and ratio of pelagic volume to benthic surface area are shown to have significant effects on the behavior of these microcosms. It is argued that the inclusion and accurate simulation of appropriate levels of these variables is important in microcosm studies designed to study the dynamics of natural systems.     

Contribution of benthic microalgae to the temporal variation in phytoplankton assemblages in a macrotidal system

Suspended marine benthic microalgae in the water column reflect the close relationship between the benthic and pelagic components of coastal ecosystems. In this study, a 12‐year phytoplankton time‐series was used to investigate the contribution of benthic microalgae to the pelagic system at a site along the French‐Atlantic coast. Furthermore, all taxa identified were allocated into different growth forms in order to study their seasonal patterns. The highest contribution of benthic microalgae was observed during the winter period, reaching up to 60% of the carbon biomass in the water column. The haptobenthic growth form showed the highest contribution in terms of biomass, dominant in the fall‐winter period when the turbidity and the river flow were high. The epipelic growth form did not follow any seasonal pattern. The epiphytic diatom Licmophora was most commonly found during summer. As benthic microalgae were found in the water column throughout the year, the temporal variation detected in the structure of pelagic assemblages in a macrotidal ecosystem was partly derived from the differentiated contribution of several benthic growth forms.     

Terrestrial subsidies to lake food webs: an experimental approach

Cross-ecosystem movements of material and energy are ubiquitous. Aquatic ecosystems typically receive material that also includes organic matter from the surrounding catchment. Terrestrial-derived (allochthonous) organic matter can enter aquatic ecosystems in dissolved or particulate form. Several studies have highlighted the importance of dissolved organic carbon to aquatic consumers, but less is known about allochthonous particulate organic carbon (POC). Similarly, most studies showing the effects of allochthonous organic carbon (OC) on aquatic consumers have investigated pelagic habitats; the effects of allochthonous OC on benthic communities are less well studied. Allochthonous inputs might further decrease primary production through light reduction, thereby potentially affecting autotrophic resource availability to consumers. Here, an enclosure experiment was carried out to test the importance of POC input and light availability on the resource use in a benthic food web of a clear-water lake. Corn starch (a C(4) plant) was used as a POC source due to its insoluble nature and its distinct carbon stable isotope value (δ(13)C). The starch carbon was closely dispersed over the bottom of the enclosures to study the fate of a POC source exclusively available to sediment biota. The addition of starch carbon resulted in a clear shift in the isotopic signature of surface-dwelling herbivorous and predatory invertebrates. Although the starch carbon was added solely to the sediment surface, the carbon originating from the starch reached zooplankton. We suggest that allochthonous POC can subsidize benthic food webs directly and can be further transferred to pelagic systems, thereby highlighting the importance of benthic pathways for pelagic habitats.     

Stability of complex food webs: resilience, resistance and the average interaction strength

In the face of stochastic climatic perturbations, the overall stability of an ecosystem will be determined by the balance between its resilience and its resistance, but their relative importance is still unknown. Using aquatic food web models we study ecosystem stability as a function of food web complexity. We measured three dynamical stability properties: resilience, resistance, and variability. Specifically, we evaluate how a decrease in the strength of predator-prey interactions with food web complexity, reflecting a decrease in predation efficiency with the number of prey per predator, affects the overall stability of the ecosystem. We find that in mass conservative ecosystems, a lower interaction strength slows down the mass cycling rate in the system and this increases its resistance to perturbations of the growth rate of primary producers. Furthermore, we show that the overall stability of the food webs is mostly given by their resistance, and not by their resilience. Resilience and resistance display opposite trends, although they are shown not to be simply opposite concepts but rather independent properties. The ecological implication is that weaker predator-prey interactions in closed ecosystems can stabilize food web dynamics by increasing its resistance to climatic perturbations.     

Lake morphometry predicts the degree of habitat coupling by a mobile predator

Habitat coupling is an ecosystem process whereby semi-discontinuous habitats are connected through the movement of energy and nutrients by chemical, physical or biological processes. One oft-cited example is that of littoral–pelagic coupling in lakes. Theory has argued that such habitat coupling may be critical to food web dynamics, yet there have been few empirical studies that have quantified ecological factors that affect the degree of habitat coupling in ecosystems. Specifically, the degree to which habitat coupling occurs across important physical gradients has largely been ignored. To address this, we investigate the degree of littoral habitat coupling (i.e. the degree to which a top predator lake trout, Salvelinus namaycush, derives energy from the littoral zone) along a gradient of lake shape, where lake shape modifies the relative quantity of coupled epilimnetic benthic and pelagic habitats within each lake. Herein we demonstrate that littoral habitat coupling is intensified in simple circular lakes compared to their reticulate counterparts in seven Canadian Shield lakes. Although the more reticulate lakes had larger areas of epilimnetic benthic habitat, littoral food sources comprised 11% compared to 24% of lake trout diet in reticulate and circular lakes, respectively. This heightened interaction in circular lakes also appears to translate into increased omnivory in more circular lakes compared to reticulate lakes such that lake trout of circular lakes have a significantly lower trophic position than lake trout of reticulate lakes (F_1,5=6.71 p=0.05). These results suggest that it is the accessibility of littoral production via thermal refugia, and not the amount of littoral production, that determines the degree to which lake trout couple littoral and pelagic habitats in lakes.     

Nitrogen cycling in coastal marine ecosystems

It is generally considered that nitrogen availability is one of the major factors regulating primary production in temperate coastal marine environments. Coastal regions often receive large anthropogenic inputs of nitrogen that cause eutrophication. The impact of these nitrogen additions has a profound effect in estuaries and coastal lagoons where water exchange is limited. Such increased nutrient loading promotes the growth of phytoplankton and fast growing pelagic macroalgae while rooted plants (sea-grasses) and benthic are suppressed due to reduced light availability. This shift from benthic to pelagic primary production introduces large diurnal variations in oxygen concentrations in the water column. In addition oxygen consumption in the surface sediments increases due to the deposition of readily degradable biomass. In this review the physico-chemical and biological factors regulating nitrogen cycling in coastal marine ecosystems are considered in relation to developing effective management programmes to rehabilitate seagrass communities in lagoons currently dominated by pelagic macroalgae and/or cyanobacteria.     

Towards the trophic structure of the Bouvet Island marine ecosystem

Although Bouvet Island is of considerable importance for Southern Ocean species conservation, information on the marine community species inventory and trophic functioning is scarce. Our combined study of stable isotopes and feeding relationships shows that (1) the marine system conforms to the trophic pattern described for other Antarctic systems within the Antarctic circumpolar current (ACC); (2) both the benthic and the pelagic subsystem are almost exclusively linked via suspended particulate organic matter (SPOM); and (3) there is no evidence of a subsystem driven by macroalgae. Bouvet Island can therefore be characterized as a benthic “oasis” within a self-sustaining open ocean pelagic system.