Vulnerability of macronutrients to the concurrent effects of enhanced temperature and atmospheric pCO<Subscript>2</Subscript> in representative shelf sea sediment habitats

Fundamental changes in seawater carbonate chemistry and sea surface temperatures associated with the ocean uptake of anthropogenic CO₂ are accelerating, but investigations of the susceptibility of biogeochemical processes to the simultaneous occurrence of multiple components of climate change are uncommon. Here, we quantify how concurrent changes in enhanced temperature and atmospheric pCO₂, coupled with an associated shift in macrofaunal community structure and behavior (sediment particle reworking and bioirrigation), modify net carbon and nutrient concentrations (NH₄-N, NO_x-N, PO₄-P) in representative shelf sea sediment habitats (mud, sandy-mud, muddy-sand and sand) of the Celtic Sea. We show that net concentrations of organic carbon, nitrogen and phosphate are, irrespective of sediment type, largely unaffected by a simultaneous increase in temperature and atmospheric pCO₂. However, our analyses also reveal that a reduction in macrofaunal species richness and total abundance occurs under future environmental conditions, varies across a gradient of cohesive to non-cohesive sediments, and negatively moderates biogeochemical processes, in particular nitrification. Our findings indicate that future environmental conditions are unlikely to have strong direct effects on biogeochemical processes but, particularly in muddy sands, the abundance, activity, composition and functional role of invertebrate communities are likely to be altered in ways that will be sufficient to regulate the function of the microbial community and the availability of nutrients in shelf sea waters.     

Sequential bioavailability of sedimentary organic matter to heterotrophic bacteria

Aquatic sediments harbour diverse microbial communities that mediate organic matter degradation and influence biogeochemical cycles. The pool of bioavailable carbon continuously changes as a result of abiotic processes and microbial activity. It remains unclear how microbial communities respond to heterogeneous organic matrices and how this ultimately affects heterotrophic respiration. To explore the relationships between the degradation of mixed carbon substrates and microbial activity, we incubated batches of organic‐rich sediments in a novel bioreactor (IsoCaRB) that permitted continuous observations of CO₂ production rates, as well as sequential sampling of isotopic signatures (δ¹³C, Δ¹⁴C), microbial community structure and diversity, and extracellular enzyme activity. Our results indicated that lower molecular weight (MW), labile, phytoplankton‐derived compounds were degraded first, followed by petroleum‐derived exogenous pollutants, and finally by higher MW polymeric plant material. This shift in utilization coincided with a community succession and increased extracellular enzyme activities. Thus, sequential utilization of different carbon pools induced changes at both the community and cellular level, shifting community composition, enzyme activity, respiration rates, and residual organic matter reactivity. Our results provide novel insight into the accessibility of sedimentary organic matter and demonstrate how bioavailability of natural organic substrates may affect the function and composition of heterotrophic bacterial populations.     

Mediation of macronutrients and carbon by post-disturbance shelf sea sediment communities

Benthic communities play a major role in organic matter remineralisation and the mediation of many aspects of shelf sea biogeochemistry. Few studies have considered how changes in community structure associated with different levels of physical disturbance affect sediment macronutrients and carbon following the cessation of disturbance. Here, we investigate how faunal activity (sediment particle reworking and bioirrigation) in communities that have survived contrasting levels of bottom fishing affect sediment organic carbon content and macronutrient concentrations ([NH₄–N], [NO₂–N], [NO₃–N], [PO₄–P], [SiO₄–Si]). We find that organic carbon content and [NO₃–N] decline in cohesive sediment communities that have experienced an increased frequency of fishing, whilst [NH₄–N], [NO₂–N], [PO₄–P] and [SiO₄–Si] are not affected. [NH₄–N] increases in non-cohesive sediments that have experienced a higher frequency of fishing. Further analyses reveal that the way communities are restructured by physical disturbance differs between sediment type and with fishing frequency, but that changes in community structure do little to affect bioturbation and associated levels of organic carbon and nutrient concentrations. Our results suggest that in the presence of physical disturbance, irrespective of sediment type, the mediation of macronutrient and carbon cycling increasingly reflects the decoupling of organism-sediment relations. Indeed, it is the traits of the species that reside at the sediment–water interface, or that occupy deeper parts of the sediment profile, that are disproportionately expressed post-disturbance, that are most important for sustaining biogeochemical functioning.     

Sediment distribution and accumulation in lagoons of the Southern Mediterranean Region (the MELMARINA Project) with special reference to environmental change and aquatic ecosystems

Surface sediments and sediment cores were collected from coastal lagoons and lakes located in the Southern Mediterranean Region (SMR) as part of the MELMARINA Project which involved integrated eco-hydrological monitoring and modelling. This study uses surface sediments and sediment cores to infer spatial characteristics and temporal changes at the MELMARINA primary sites, Merja Zerga in Morocco, Ghar El Melh in Tunisia and Lake Manzala in Egypt. In addition, surface sediment sampling was undertaken at Egyptian Lake Bardawil and sediment cores were collected from the Lagune de Nador (Morocco). Sediment distribution patterns are investigated using GIS with georeferenced sample locations to facilitate display and resurvey. Major variations in sedimentary organic matter and, particularly, carbonate content, occur within and between sites. Local landscapes combined with hydrological and biogeochemical processes influence the distributions of sediment bulk components (carbonates, organic material and clastic matter) and molluscan shells and shell debris are an important source of sedimentary carbonate at all three primary sites. Sediment cores were dated using natural (²¹⁰Pb) and artificial (¹³⁷Cs) radionuclides, and sediment accumulation rate changes indicate that sources of sediment supply varied markedly through the twentieth century but have generally diminished after the mid-1960s. Sedimentary siliceous microfossils (diatoms) were generally poorly preserved, but mollusc shell remains were well represented. Sediment chronologies and sediment bulk composition allow discussion of some recent changes in bulk, minerogenic and biogenic sediment accumulation patterns in the SMR lagoons. Sediment accumulation rates also varied between sites and multiple cores from Lake Manzala indicated that rates showed considerable spatial variability. Low-level sediment contamination by fossil fuel combustion particulates and trace metals was demonstrated for Ghar El Melh and Lagune de Nador where Pb and Zn accumulation rates were highest in twentieth century sediment. It is emphasized that sediment quality and quantity have strong influences on lagoon ecosystem function and sedimentation is relevant to hydromorphology and to concepts of ecological quality. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Guest editors: J. R. Thompson & R. J. Flower Hydro-ecological Monitoring and Modelling of North African Coastal Lagoons     

Functional diversity of marine macrobenthic communities from sublittoral soft-sediment habitats off northern Chile

Benthic communities show changes in composition and structure across different environmental characteristics and habitats. However, incorporating species biological traits into the analysis can provide a better understanding of system functioning within habitats. We compare the functional diversity of macrobenthic communities from a contrasting shallow (15 m) and deep (50 m) sublittoral soft-sediment habitats in northern Chile, using biological traits analysis. Our aim was to highlight the biological characteristics responsible for differences between habitats and the implications for ecosystem functioning. Trait analysis showed that the deep habitat was restricted in providing functionally important biogenic structure and bioturbation and supports less diverse feeding-related energy pathways. The shallow habitat is characterized by more diverse energy pathways and a higher potential for matter exchange through bioturbation. We provide support to the predictions of transfer of energy from the benthos to upper trophic levels in the shallow, which is characterized mainly by normoxia and little organic matter content in the sediment. In the deep habitat, characterized by hypoxia and more organic matter, energy appears to be transferred to microbial components. We suggest that trait analysis should be added to the traditional approaches based on species diversity, because it provides indicators of ecosystem stress.     

Variable Importance of Macrofaunal Functional Biodiversity for Biogeochemical Cycling in Temperate Coastal Sediments

Coastal marine systems are currently subject to a variety of anthropogenic and climate-change-induced pressures. An important challenge is to predict how marine sediment communities and benthic biogeochemical cycling will be affected by these ongoing changes. To this end, it is of paramount importance to first better understand the natural variability in coastal benthic biogeochemical cycling and how this is influenced by local environmental conditions and faunal biodiversity. Here, we studied sedimentary biogeochemical cycling at ten coastal stations in the Southern North Sea on a monthly basis from February to October 2011. We explored the spatio-temporal variability in oxygen consumption, dissolved inorganic nitrogen and alkalinity fluxes, and estimated rates of nitrification and denitrification from a mass budget. In a next step, we statistically modeled their relation with environmental variables and structural and functional macrobenthic community characteristics. Our results show that the cohesive, muddy sediments were poor in functional macrobenthic diversity and displayed intermediate oxygen consumption rates, but the highest ammonium effluxes. These muddy sites also showed an elevated alkalinity release from the sediment, which can be explained by the elevated rate of anaerobic processes taking place. Fine sandy sediments were rich in functional macrobenthic diversity and had the maximum oxygen consumption and estimated denitrification rates. Permeable sediments were also poor in macrobenthic functional diversity and showed the lowest oxygen consumption rates and only small fluxes of ammonium and alkalinity. Macrobenthic functional biodiversity as estimated from bioturbation potential appeared a better variable than macrobenthic density in explaining oxygen consumption, ammonium and alkalinity fluxes, and estimated denitrification. However, this importance of functional biodiversity was manifested particularly in fine sandy sediments, to a lesser account in permeable sediments, but not in muddy sediments. The strong relationship between macrobenthic functional biodiversity and biogeochemical cycling in fine sandy sediments implies that a future loss of macrobenthic functional diversity will have important repercussions for benthic ecosystem functioning.     

Influence of ecosystem engineers on ecosystem processes is mediated by lake sediment properties

Invasive species may impact biotic community structure, ecosystem processes, and associated goods and services. Their impact may be especially strong when they also serve as ecosystem engineers (i.e. organisms affecting the physical habitat and resources for other species). Dreissenid mussels fill both these roles, having invaded the Laurentian Great Lakes in the late 1980s, and also serve as ecosystem engineers by altering nutrient fluxes and influencing the microbial food web through direct nutrient release and excretion of feces and pseudo‐feces at the water–sediment interface. We conducted laboratory experiments to investigate how the different functional traits of invasive quagga mussels (filtering activity and direct nutrient release) and native chironomid larvae (tube building and bioirrigation) interact with lake sediment of differing organic matter content to influence biogeochemical fluxes and water quality. Our results showed that sediment reworking and ventilation activities by chironomid larvae increased oxygen penetration in the sediment, affecting primarily pore water chemistry, whereas invasive mussels enhanced nutrient releases in the surface water. However, sediment organic matter modulated the effects of ecosystem engineers on system‐level processes; chironomids had a greater influence on sediment reworking and microbial‐mediated processes in organic‐rich sediments, whereas quagga mussels enhanced nutrient concentrations in the overlying water of organic‐poor sediments. These results have management implications, as the effects of invasive mussels on the biogeochemical functioning in the Great Lakes region and elsewhere can alter system bioenergetics and promote harmful algal blooms.     

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

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

Metagenomic and stable isotopic analyses of modern freshwater microbialites in Cuatro Ciénegas, Mexico

Ancient biologically mediated sedimentary carbonate deposits, including stromatolites and other microbialites, provide insight into environmental conditions on early Earth. The primary limitation to interpreting these records is our lack of understanding regarding microbial processes and the preservation of geochemical signatures in contemporary microbialite systems. Using a combination of metagenomic sequencing and isotopic analyses, this study describes the identity, metabolic potential and chemical processes of microbial communities from living microbialites from Cuatro Ciénegas, Mexico. Metagenomic sequencing revealed a diverse, redox-dependent microbial community associated with the microbialites. The microbialite community is distinct from other marine and freshwater microbial communities, and demonstrates extensive environmental adaptation. The microbialite metagenomes contain a large number of genes involved in the production of exopolymeric substances and the formation of biofilms, creating a complex, spatially structured environment. In addition to the spatial complexity of the biofilm, microbial activity is tightly controlled by sensory and regulatory systems, which allow for coordination of autotrophic and heterotrophic processes. Isotopic measurements of the intracrystalline organic matter demonstrate the importance of heterotrophic respiration of photoautotrophic biomass in the precipitation of calcium carbonate. The genomic and stable isotopic data presented here significantly enhance our evolving knowledge of contemporary biomineralization processes, and are directly applicable to studies of ancient microbialites.     

Preservation of overmature,ancient, sedimentary organic matter in carbonate concretions during outcrop weathering

Concretions are preferentially cemented zones within sediments and sedimentary rocks. Cementation can result from relatively early diagenetic processes that include degradation of sedimentary organic compounds or methane as indicated by significantly ¹³C‐depleted or enriched carbon isotope compositions. As minerals fill pore space, reduced permeability may promote preservation of sediment components from degradation during subsequent diagenesis, burial heating and outcrop weathering. Discrete and macroscopic organic remains, macro and microfossils, magnetic grains, and sedimentary structures can be preferentially preserved within concretions. Here, Cretaceous carbonate concretions of the Holz Shale are shown to contain relatively high carbonate‐free total organic carbon (TOC) contents (up to ~18.5 wt%) compared to the surrounding host rock (with <2.1 wt%). TOC increases with total inorganic carbon (TIC) content, a metric of the degree of cementation. Pyrite contents within concretions generally correlate with organic carbon contents. Concretion carbonate carbon isotope compositions (δ¹³C_carb) range from ?22.5 to ?3.4‰ (VPDB) and do not correlate strongly with TOC. Organic carbon isotope compositions (δ¹³C_org) of concretions and host rock are similar. Thermal maturity data indicate that both host and concretion organic matter are overmature and have evolved beyond the oil window maturity stage. Although the organic matter in general has experienced significant oxidative weathering, concretion interiors exhibit lower oxygen indices relative to the host. These results suggest that carbonate concretions can preferentially preserve overmature, ancient, sedimentary organic matter during outcrop weathering, despite evidence for organic matter degradation genetic mechanisms. As a result, concretions may provide an optimal proxy target for characterization of more primary organic carbon concentrations and chemical compositions. In addition, these findings indicate that concretions can promote delayed oxidative weathering of organic carbon in outcrop and therefore impact local chemical cycling.     

Impacts of Ocean Acidification on Sediment Processes in Shallow Waters of the Arctic Ocean

Despite the important roles of shallow-water sediments in global biogeochemical cycling, the effects of ocean acidification on sedimentary processes have received relatively little attention. As high-latitude cold waters can absorb more CO₂ and usually have a lower buffering capacity than warmer waters, acidification rates in these areas are faster than those in sub-tropical regions. The present study investigates the effects of ocean acidification on sediment composition, processes and sediment-water fluxes in an Arctic coastal system. Undisturbed sediment cores, exempt of large dwelling organisms, were collected, incubated for a period of 14 days, and subject to a gradient of pCO₂ covering the range of values projected for the end of the century. On five occasions during the experimental period, the sediment cores were isolated for flux measurements (oxygen, alkalinity, dissolved inorganic carbon, ammonium, nitrate, nitrite, phosphate and silicate). At the end of the experimental period, denitrification rates were measured and sediment samples were taken at several depth intervals for solid-phase analyses. Most of the parameters and processes (i.e. mineralization, denitrification) investigated showed no relationship with the overlying seawater pH, suggesting that ocean acidification will have limited impacts on the microbial activity and associated sediment-water fluxes on Arctic shelves, in the absence of active bio-irrigating organisms. Only following a pH decrease of 1 pH unit, not foreseen in the coming 300 years, significant enhancements of calcium carbonate dissolution and anammox rates were observed. Longer-term experiments on different sediment types are still required to confirm the limited impact of ocean acidification on shallow Arctic sediment processes as observed in this study.     

Biogeochemical processes in the groundwater discharge zone of urban streams

The influence of biogeochemical processes on nitrogen and organic matter transformation and transport was investigated for two urban streams receiving groundwater discharge during the dry summer baseflow period. A multiple lines of evidence approach involving catchment-, and stream reach-scale investigations were undertaken to describe the factors that influence pore water biogeochemical processes. At the catchment-scale gaining stream reaches were identified from water table mapping and groundwater discharge estimated to be between 0.1 and 0.8 m³ m^?2 d^?1 from baseflow analysis. Sediment temperature profiles also suggested that the high groundwater discharge limited stream water infiltration into the sediments. At the stream reach-scale, dissolved organic carbon (DOC) and dissolved organic nitrogen (DON) concentrations were higher in stream water than in groundwater. However, DOC and DON concentrations were greatest in sediment pore water. This suggests that biodegradation of sediment organic matter contributes dissolved organic matter (DOM) to the streams along with that delivered with groundwater flow. Pore water ammonium (NH₄ ⁺) was closely associated with areas of high pore water DOM concentrations and evidence of sulfate (SO₄ ^2?) reduction (low concentration and SO₄:Cl ratio). This indicates that anoxic DOM mineralization was occurring associated with SO₄ ^2? reduction. However the distribution of anoxic mineralization was limited to the center of the streambed, and was not constrained by the distribution of sediment organic matter which was higher along the banks. Lower sediment temperatures measured along the banks compared to the center suggests, at least qualitatively, that groundwater discharge is higher along the banks. Based on this evidence anoxic mineralization is influenced by groundwater residence time, and is only measurable along the center of the stream where groundwater flux rates are lower. This study therefore shows that the distribution of biogeochemical processes in stream sediments, such as anoxic mineralization, is strongly influenced by both the biogeochemical conditions and pore water residence time.     

Influence of hyporheic zone characteristics on the structure and activity of microbial assemblages

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Activity and community structures of sulfate-reducing microorganisms in polar,temperate and tropical marine sediments

Temperature has a fundamental impact on the metabolic rates of microorganisms and strongly influences microbial ecology and biogeochemical cycling in the environment. In this study, we examined the catabolic temperature response of natural communities of sulfate-reducing microorganisms (SRM) in polar, temperate and tropical marine sediments. In short-term sediment incubation experiments with ³⁵S-sulfate, we demonstrated how the cardinal temperatures for sulfate reduction correlate with mean annual sediment temperatures, indicating specific thermal adaptations of the dominant SRM in each of the investigated ecosystems. The community structure of putative SRM in the sediments, as revealed by pyrosequencing of bacterial 16S rRNA gene amplicons and phylogenetic assignment to known SRM taxa, consistently correlated with in situ temperatures, but not with sediment organic carbon concentrations or C:N ratios of organic matter. Additionally, several species-level SRM phylotypes of the class Deltaproteobacteria tended to co-occur at sites with similar mean annual temperatures, regardless of geographic distance. The observed temperature adaptations of SRM imply that environmental temperature is a major controlling variable for physiological selection and ecological and evolutionary differentiation of microbial communities.     

Major Role of Microbes in Carbon Fluxes during Austral Winter in the Southern Drake Passage

Carbon cycling in Southern Ocean is a major issue in climate change, hence the need to understand the role of biota in the regulation of carbon fixation and cycling. Southern Ocean is a heterogeneous system, characterized by a strong seasonality, due to long dark winter. Yet, currently little is known about biogeochemical dynamics during this season, particularly in the deeper part of the ocean. We studied bacterial communities and processes in summer and winter cruises in the southern Drake Passage. Here we show that in winter, when the primary production is greatly reduced, Bacteria and Archaea become the major producers of biogenic particles, at the expense of dissolved organic carbon drawdown. Heterotrophic production and chemoautotrophic CO₂ fixation rates were substantial, also in deep water, and bacterial populations were controlled by protists and viruses. A dynamic food web is also consistent with the observed temporal and spatial variations in archaeal and bacterial communities that might exploit various niches. Thus, Southern Ocean microbial loop may substantially maintain a wintertime food web and system respiration at the expense of summer produced DOC as well as regenerate nutrients and iron. Our findings have important implications for Southern Ocean ecosystem functioning and carbon cycle and its manipulation by iron enrichment to achieve net sequestration of atmospheric CO₂.     

Benthic Oxygen Uptake in the Arctic Ocean Margins - A Case Study at the Deep-Sea Observatory HAUSGARTEN (Fram Strait)

The past decades have seen remarkable changes in the Arctic, a hotspot for climate change. Nevertheless, impacts of such changes on the biogeochemical cycles and Arctic marine ecosystems are still largely unknown. During cruises to the deep-sea observatory HAUSGARTEN in July 2007 and 2008, we investigated the biogeochemical recycling of organic matter in Arctic margin sediments by performing shipboard measurements of oxygen profiles, bacterial activities and biogenic sediment compounds (pigment, protein, organic carbon, and phospholipid contents). Additional in situ oxygen profiles were performed at two sites. This study aims at characterizing benthic mineralization activity along local bathymetric and latitudinal transects. The spatial coverage of this study is unique since it focuses on the transition from shelf to Deep Ocean, and from close to the ice edge to more open waters. Biogeochemical recycling across the continental margin showed a classical bathymetric pattern with overall low fluxes except for the deepest station located in the Molloy Hole (5500 m), a seafloor depression acting as an organic matter depot center. A gradient in benthic mineralization rates arises along the latitudinal transect with clearly higher values at the southern stations (average diffusive oxygen uptake of 0.49 ± 0.18 mmol O₂ m^-2 d^-1) compared to the northern sites (0.22 ± 0.09 mmol O₂ m^-2 d^-1). The benthic mineralization activity at the HAUSGARTEN observatory thus increases southward and appears to reflect the amount of organic matter reaching the seafloor rather than its lability. Although organic matter content and potential bacterial activity clearly follow this gradient, sediment pigments and phospholipids exhibit no increase with latitude whereas satellite images of surface ocean chlorophyll a indicate local seasonal patterns of primary production. Our results suggest that predicted increases in primary production in the Arctic Ocean could induce a larger export of more refractory organic matter due to the longer production season and the extension of the ice-free zone.     

Sedimentary context controls the influence of ecosystem engineering by bioturbators on microbial processes in river sediments

By modifying the physical environment, ecosystem engineers can have inordinately large effects on surrounding communities and ecosystem functioning. However, the significance of engineering in ecosystems greatly depends on the physical characteristics of the engineered habitats. Mechanisms underlying such context‐dependent impact of engineers remain poorly understood even though they are crucial to establish general predictions concerning the contribution of engineers to ecosystem structure and function. The present study aimed to decrypt such mechanisms by determining how the environmental context modulates the effects of ecosystem engineers (bioturbators) on microorganisms in river sediments. To test the effects of environmental context on the role of bioturbators in sediments, we used mesocosms and recreated two sedimentary contexts in the laboratory by adding a layer of either fine or coarse sand at the top of a gravel‐sand matrix. For each sediment context, we examined how the sediment reworking activity of a bioturbating tubificid worm (Tubifex tubifex) generated changes in the physical (sediment structure and permeability) and abiotic environments (hydraulic discharge, water chemistry) of microorganisms. Microbial characteristics (abundances, activities) and leaf litter decomposition – a major microbially‐mediated ecological process – were measured to evaluate the impact of bioturbation on biotic compartment. Our results showed that the permeability, the availability of oxygen and the activities of microorganisms were reduced in sediments covered with fine sand, in comparison with sediments covered with coarse sand. Tubifex tubifex significantly increased permeability (by about six‐fold), restored aerobic conditions and ultimately stimulated microbial communities (resulting in a 30% increase in leaf litter breakdown rate) in sediments covered with fine sand. In contrast T. tubifex had low effects in sediments topped by coarse sand, where O₂ was already available for hyporheic microorganisms. Our study supports the idea that context dependency mainly modulates the effects of engineering by controlling the ability of engineers to create changes on abiotic (O₂ in the present study) factors that are limiting for surrounding communities.     

Species-Specific Effects on Ecosystem Functioning Can Be Altered by Interspecific Interactions

Biological assemblages are constantly undergoing change, with species being introduced, extirpated and experiencing shifts in their densities. Theory and experimentation suggest that the impacts of such change on ecosystem functioning should be predictable based on the biological traits of the species involved. However, interspecific interactions could alter how species affect functioning, with the strength and sign of interactions potentially depending on environmental context (e.g. homogenous vs. heterogeneous conditions) and the function considered. Here, we assessed how concurrent changes to the densities of two common marine benthic invertebrates, Corophium volutator and Hediste diversicolor, affected the ecological functions of organic matter consumption and benthic-pelagic nutrient flux. Complementary experiments were conducted within homogenous laboratory microcosms and naturally heterogeneous field plots. When the densities of the species were increased within microcosms, interspecific interactions enhanced effects on organic matter consumption and reduced effects on nutrient flux. Trait-based predictions of how each species would affect functioning were only consistently supported when the density of the other species was low. In field plots, increasing the density of either species had a positive effect on organic matter consumption (with no significant interspecific interactions) but no effect on nutrient flux. Our results indicate that species-specific effects on ecosystem functioning can be altered by interspecific interactions, which can be either facilitative (positive) or antagonistic (negative) depending on the function considered. The impacts of biodiversity change may therefore not be predictable based solely on the biological traits of the species involved. Possible explanations for why interactions were detected in microcosms but not in the field are discussed.     

Nematode biomass and allometric attributes as indicators of environmental quality in a Mediterranean harbour (Ligurian Sea,Italy)

Nematode biomass and allometric attributes (size spectra, body length and width, morphotypes) were measured and related to the environmental quality of sediments of the Genoa-Voltri commercial harbour (Ligurian Sea, NW Mediterranean) to investigate their possible use as ecological indicators. The sediment quality was defined by measuring the level of organic enrichment (quantity and biochemical composition of sedimentary organic matter) and oxygen stress (redox potential).Nematode biomass spectra (NBS) proved to be extremely valuable in determining differences in the environmental quality of sediments on a hundred-metre spatial scale. High peaks of the NBS were observed in the more organic-rich and oxygen-stressed stations probably in relation to a lower diversity of the nematode communities in these stations, with the predominance of tolerant genera such as Paracomesoma and Sabatieria. Among allometric variables, the length was found to be negatively correlated with oxygen concentrations and positively with TOM percentages, whilst the morphotype length/width ratio (L/W) resulted negatively related to oxygen concentrations and organic matter quality (protein:carbohydrate ratio), suggesting that these allometric attributes represent indicators of the functional adaptation of nematodes to the changing environmental conditions.We suggest that in contrast to time-consuming and expertise-requiring nematode taxonomic analysis, biomass and allometric attributes analysis can provide a simpler but comparable tool to assess sediment quality and environmental heterogeneity of harbour ecosystems.     

Benthic pH gradients across a range of shelf sea sediment types linked to sediment characteristics and seasonal variability

This study used microelectrodes to record pH profiles in fresh shelf sea sediment cores collected across a range of different sediment types within the Celtic Sea. Spatial and temporal variability was captured during repeated measurements in 2014 and 2015. Concurrently recorded oxygen microelectrode profiles and other sedimentary parameters provide a detailed context for interpretation of the pH data. Clear differences in profiles were observed between sediment type, location and season. Notably, very steep pH gradients exist within the surface sediments (10–20 mm), where decreases greater than 0.5 pH units were observed. Steep gradients were particularly apparent in fine cohesive sediments, less so in permeable sandier matrices. We hypothesise that the gradients are likely caused by aerobic organic matter respiration close to the sediment–water interface or oxidation of reduced species at the base of the oxic zone (NH₄ ⁺, Mn²⁺, Fe²⁺, S^?). Statistical analysis suggests the variability in the depth of the pH minima is controlled spatially by the oxygen penetration depth, and seasonally by the input and remineralisation of deposited organic phytodetritus. Below the pH minima the observed pH remained consistently low to maximum electrode penetration (ca. 60 mm), indicating an absence of sub-oxic processes generating H⁺ or balanced removal processes within this layer. Thus, a climatology of sediment surface porewater pH is provided against which to examine biogeochemical processes. This enhances our understanding of benthic pH processes, particularly in the context of human impacts, seabed integrity, and future climate changes, providing vital information for modelling benthic response under future climate scenarios.