Direct evidence of a prey depletion "halo" surrounding a pelagic predator colony.

Colonially breeding birds and mammals form some of the largest gatherings of apex predators in the natural world and have provided model systems for studying mechanisms of population regulation in animals. According to one influential hypothesis, intense competition for food among large numbers of spatially constrained foragers should result in a zone of prey depletion surrounding such colonies, ultimately limiting their size. However, while indirect and theoretical support for this phenomenon, known as "Ashmole's halo," has steadily accumulated, direct evidence remains exceptionally scarce. Using a combination of vessel-based surveys and Global Positioning System tracking, we show that pelagic seabirds breeding at the tropical island that first inspired Ashmole's hypothesis do indeed deplete their primary prey species (flying fish; Exocoetidae spp.) over a considerable area, with reduced prey density detectable >150 km from the colony. The observed prey gradient was mirrored by an opposing trend in seabird foraging effort, could not be explained by confounding environmental variability, and can be approximated using a mechanistic consumption-dispersion model, incorporating realistic rates of seabird predation and random prey dispersal. Our results provide a rare view of the resource footprint of a pelagic seabird colony and reveal how aggregations of these central-place foraging, marine top predators profoundly influence the oceans that surround them.

Mind the Exposure Gaps-Modeling Chemical Transport in Sediment Toxicity Tests.

Chemical exposure in flow-through sediment toxicity tests can vary in time, between pore and overlying water, and amid free and bound states, complicating the link between toxicity and observable concentrations such as free pore (Cfree,pore), free overlying (Cfree,over), or the corresponding dissolved concentrations (Cdiss, free + bound to dissolved organic carbon, DOC). We introduce a numerical model that describes the desorption from sediments to pore water, diffusion through pores and the sediment-water boundary, DOC-mediated transport, and mixing in and outflow from overlying water. The model explained both the experimentally measured gap between Cfree,over and Cfree,pore and the continuous decrease in overlying Cdiss. Spatially resolved modeling suggested a steep concentration gradient present in the upper millimeter of the sediment due to slow chemical diffusion in sediment pores and fast outflux from the overlying water. In contrast to continuous decrease in overlying Cdiss expected for any chemical, Cfree,over of highly hydrophobic chemicals was kept relatively constant following desorption from DOC, a mechanism comparable to passive dosing. Our mechanistic analyses emphasize that exposure will depend on the chemical's hydrophobicity, the test organism habitat and uptake of bound chemicals, and the properties of sediment components, including DOC. The model can help to re-evaluate existing toxicity data, optimize experimental setups, and extrapolate laboratory toxicity data to field exposure.

Ocean Alkalinity, Buffering and Biogeochemical Processes.

Alkalinity, the excess of proton acceptors over donors, plays a major role in ocean chemistry, in buffering and in calcium carbonate precipitation and dissolution. Understanding alkalinity dynamics is pivotal to quantify ocean carbon dioxide uptake during times of global change. Here we review ocean alkalinity and its role in ocean buffering as well as the biogeochemical processes governing alkalinity and pH in the ocean. We show that it is important to distinguish between measurable titration alkalinity and charge balance alkalinity that is used to quantify calcification and carbonate dissolution and needed to understand the impact of biogeochemical processes on components of the carbon dioxide system. A general treatment of ocean buffering and quantification via sensitivity factors is presented and used to link existing buffer and sensitivity factors. The impact of individual biogeochemical processes on ocean alkalinity and pH is discussed and quantified using these sensitivity factors. Processes governing ocean alkalinity on longer time scales such as carbonate compensation, (reversed) silicate weathering, and anaerobic mineralization are discussed and used to derive a close-to-balance ocean alkalinity budget for the modern ocean.

Biogeochemical dynamics in a marine storm demonstrates differences between natural and anthropogenic impacts.

This study explores the impact of a wind storm on sediment resuspension and marine biogeochemical dynamics. Additionally, the storm took place during an expedition researching bottom trawling, enabling the direct comparison of certain natural and fisheries-related disturbances. The storm was initiated by a decline in atmospheric pressure and a 2 h period of gale force winds, which was followed by over 40 h of elevated bottom currents. Storm induced turbidity, potentially a cumulative post-fishing impact, was remarkably higher compared to what was observed in a recent trawling event. Storm-induced mixing and movement of water masses led to decreased silicate and increased phosphate concentrations in the water column, accompanied by lower salinity and higher fluorescence. The erosion depth of the seabed averaged around 0.3 cm during the peak turbidity period. Trawl-induced erosion in the area has been measured at over twice that depth, and has been linked to intermittent reductions in near-bed oxygen levels. In contrast, storm-induced turbidity coincided with increased oxygen due to wave mixing, suggesting inherent differences in how trawling and storms can oxidize reduced substances. These findings suggest that storms have a greater regional impact, whereas the local impacts of bottom trawling on biogeochemistry can be more significant.

Partial recovery of macrozoobenthos on the northwestern shelf of the Black Sea.

The northwestern shelf of the Black Sea has been affected by eutrophication and bottom hypoxia since the sixties. Consequently, the macrozoobenthos has suffered a well-established decline in biodiversity. However, the nature of the current benthic communities remains questionable. From 1995 to 2017, we compiled species and abiotic data for 138 sites over the shelf. Through an appropriate multivariate analytical approach, we identified benthic community changes solely due to organic pollution variations. Our results show signs of recovery with an increase in biodiversity and proportion of species vulnerable to organic enrichment. These changes were related to a decrease in riverine loads and subsequent eutrophication. However, some long-lived species typical of the area still did not exhibit noticeable recovery, which suggests that either the recovery process has not yet been achieved or some environmental conditions are still not met to warrant a sea floor ecosystem state substantially healthy.

The relative roles of multiple drivers on benthic ciliate communities in an intertidal zone.

Intertidal biodiversity is being severely disrupted as a result of increased anthropogenic activity. However, our knowledge about how natural gradients, human induced disturbance and biotic interactions affect biodiversity is limited. So, we investigated how three facets of alpha diversity and community composition of benthic ciliates responded to environmental and biological gradients in the intertidal zone of Zhejiang, China. The key determinants and their relative effects on ciliate communities were identified using structural equation modeling, distance-based redundancy analysis and variation partitioning analysis. Our results revealed that sediment grain size was the most important factor affecting alpha diversity and community composition. Human induced eutrophication had significant effects on phylogenetic alpha diversity and community composition. However, the effects of biotic interactions on ciliate communities were relatively small. Moreover, we found community composition was more sensitive to human disturbance than alpha diversity, thus, more suitable for indicating human-induced eutrophication.

Polymetallic nodules are essential for food-web integrity of a prospective deep-seabed mining area in Pacific abyssal plains.

Polymetallic nodule fields provide hard substrate for sessile organisms on the abyssal seafloor between 3000 and 6000 m water depth. Deep-seabed mining targets these mineral-rich nodules and will likely modify the consumer-resource (trophic) and substrate-providing (non-trophic) interactions within the abyssal food web. However, the importance of nodules and their associated sessile fauna in supporting food-web integrity remains unclear. Here, we use seafloor imagery and published literature to develop highly-resolved trophic and non-trophic interaction webs for the Clarion-Clipperton Fracture Zone (CCZ, central Pacific Ocean) and the Peru Basin (PB, South-East Pacific Ocean) and to assess how nodule removal may modify these networks. The CCZ interaction web included 1028 compartments connected with 59,793 links and the PB interaction web consisted of 342 compartments and 8044 links. We show that knock-down effects of nodule removal resulted in a 17.9% (CCZ) to 20.8% (PB) loss of all taxa and 22.8% (PB) to 30.6% (CCZ) loss of network links. Subsequent analysis identified stalked glass sponges living attached to the nodules as key structural species that supported a high diversity of associated fauna. We conclude that polymetallic nodules are critical for food-web integrity and that their absence will likely result in reduced local benthic biodiversity.

Faunal and environmental drivers of carbon and nitrogen cycling along a permeability gradient in shallow North Sea sediments.

Ecosystem functions are driven by abiotic and biotic factors, but due to high collinearity of both, it is often difficult to disentangle the drivers of these ecosystem functions. We studied sedimentological and faunal controls of benthic organic matter mineralization, a crucial ecosystem process provided for by sediments of shelf seas. Subtidal benthic habitats representative of the wide permeability gradient found in the Belgian Part of the North Sea (Northeast Atlantic Shelf) were characterized in terms of sediment descriptors, macrofauna, and sediment biogeochemistry was estimated. Our results confirmed a strong correlation between sediment characteristics and macrofauna, and estimated sediment biogeochemical process rates were clearly linked to both. Results of variance partitioning and statistical modelling showed that oxic mineralization and nitrification were mainly regulated by faunal activities whereas anoxic mineralization was regulated by sediment properties, with permeability as a decisive factor. Both biotic and abiotic factors were needed to explain variability in oxygen consumption and total mineralization estimates, suggesting that macrofaunal activities have different effects across habitats. The statistical models were a useful tool to interpret the impact of anthropogenic activities in the study area and represent a step towards predicting the effects of human activities on crucial ecosystem functions.

Controlling CaCO3 Particle Size with {Ca2+}:{CO3 2-} Ratios in Aqueous Environments.

The effect of stoichiometry on the new formation and subsequent growth of CaCO3 was investigated over a large range of solution stoichiometries (10-4 < r aq < 104, where r aq = {Ca2+}:{CO3 2-}) at various, initially constant degrees of supersaturation (30 < Ωcal < 200, where Ωcal = {Ca2+}{CO3 2-}/K sp), pH of 10.5 ± 0.27, and ambient temperature and pressure. At r aq = 1 and Ωcal < 150, dynamic light scattering (DLS) showed that ion adsorption onto nuclei (1-10 nm) was the dominant mechanism. At higher supersaturation levels, no continuum of particle sizes is observed with time, suggesting aggregation of prenucleation clusters into larger particles as the dominant growth mechanism. At r aq ≠ 1 (Ωcal = 100), prenucleation particles remained smaller than 10 nm for up to 15 h. Cross-polarized light in optical light microscopy was used to measure the time needed for new particle formation and growth to at least 20 µm. This precipitation time depends strongly and asymmetrically on r aq. Complementary molecular dynamics (MD) simulations confirm that r aq affects CaCO3 nanoparticle formation substantially. At r aq = 1 and Ωcal ≫ 1000, the largest nanoparticle in the system had a 21-68% larger gyration radius after 20 ns of simulation time than in nonstoichiometric systems. Our results imply that, besides Ωcal, stoichiometry affects particle size, persistence, growth time, and ripening time toward micrometer-sized crystals. Our results may help us to improve the understanding, prediction, and formation of CaCO3 in geological, industrial, and geo-engineering settings.

Ecological significance of hazardous concentrations in a planktonic food web.

Species sensitivity distributions (SSDs) are statistical distributions that are used to estimate the potentially affected fraction (PAF) of species at a given toxicant concentration, the hazardous concentration for that fraction of species (HC(PAF)). Here, we use an aquatic food web model that includes 14 phytoplankton and 6 zooplankton species to estimate the number of species experiencing a biomass reduction when the food web is exposed to the HC(PAF) and this for 1000 hypothetical toxicants and for PAF=5-30%. When choosing a 20% decrease as a cut-off to categorize a species' biomass as affected, 0-1 and 2-5 out of the 20 species were affected at the HC(5) and HC(30), respectively. From this, it can be concluded that the PAF is a relatively good estimator of the number of affected species. However, when phytoplankton species experiencing >or=20% biomass increase were also classified as affected, the number of affected species predicted by the food web model varied strongly among toxicants for PAF >5, with 2-16 out of 20 species affected at the HC(30). Phytoplankton species with extreme (both high and low) values for uptake rates and light limitation constants experienced smaller effects on their biomass than phytoplankton species with more average parameter values. We conclude that, next to measures of toxicity, ecological characteristics of species may help understanding ecological effects occurring in ecosystems also.

Seasonal variability in ecosystem functioning across estuarine gradients: The role of sediment communities and ecosystem processes.

Functional trait approaches advance the understanding of biodiversity-ecosystem function (BDEF) relationships and its control by the environmental context. Application of these insights into management remains constrained due to lack of evidence from real-world ecosystems that capture the natural spatial and temporal gradients at which biodiversity and environmental conditions operate. In this study we measured macrofauna community traits, ecosystem processes and abiotic properties at 9 locations during 4 months, spanning a wide gradient in sedimentary habitats and salinity in the Scheldt estuary, and quantified the (a)biotic contribution to sediment community oxygen consumption, as a measure of ecosystem function. We found that functional attributes of the macrofauna community and its effect on bio-irrigation can predict ecosystem function, but especially during the colder period of the year. This result highlights that generalizations about BDEF relationships, and biodiversity loss on ecosystem functions, are limited whenever this temporal component is not acknowledged.

Experimental bottom trawling finds resilience in large-bodied infauna but vulnerability for epifauna and juveniles in the Frisian Front.

In this study, we analysed the benthic effects of two in situ fisheries disturbance experiments using a combination of side-scan sonar, high definition underwater video, sediment profile imagery, and box core sampling techniques after conventional beam trawling and box core sampling after electric pulse trawling in a southern North Sea habitat. Acoustic and optical methods visualised the morphological changes induced by experimental beam trawling, showing the flattening and homogenisation of surface sediments. Video transects found a 94% decrease in epibenthos in beam trawled sediments compared to an untrawled control site and a 74% decrease in untrawled sediments of the same transect. Box core samples taken 5.5 h, 29 h and 75 h after trawling detected a downward trend in infaunal densities and species richness that continued after the initial impact with small-bodied and juvenile taxa being especially prone to depletion. Data from shallow sediment samples showed trawl resilience in large mud shrimps and evidence of their upward movement towards the sediment surface after disturbance. Both trawl gears induced significant changes to infaunal communities, with no differential effect between the two gears. Our results suggest that in the Frisian Front, trawling may favour the survival of deep burrowers while removing surficial macrofauna.

The BenBioDen database, a global database for meio-, macro- and megabenthic biomass and densities.

Benthic fauna refers to all fauna that live in or on the seafloor, which researchers typically divide into size classes meiobenthos (32/64 µm-0.5/1 mm), macrobenthos (250 µm-1 cm), and megabenthos (>1 cm). Benthic fauna play important roles in bioturbation activity, mineralization of organic matter, and in marine food webs. Evaluating their role in these ecosystem functions requires knowledge of their global distribution and biomass. We therefore established the BenBioDen database, the largest open-access database for marine benthic biomass and density data compiled so far. In total, it includes 11,792 georeferenced benthic biomass and 51,559 benthic density records from 384 and 600 studies, respectively. We selected all references following the procedure for systematic reviews and meta-analyses, and report biomass records as grams of wet mass, dry mass, or ash-free dry mass, or carbon per m2 and as abundance records as individuals per m2. This database provides a point of reference for future studies on the distribution and biomass of benthic fauna.

The SCOC database, a large, open, and global database with sediment community oxygen consumption rates.

Sediment community oxygen consumption (SCOC) rates provide important information about biogeochemical processes in marine sediments and the activity of benthic microorganisms and fauna. Therefore, several databases of SCOC data have been compiled since the mid-1990s. However, these earlier databases contained much less data records and were not freely available. Additionally, the databases were not transparent in their selection procedure, so that other researchers could not assess the quality of the data. Here, we present the largest, best documented, and freely available database of SCOC data compiled to date. The database is comprised of 3,540 georeferenced SCOC records from 230 studies that were selected following the procedure for systematic reviews and meta-analyses. Each data record states whether the oxygen consumption was measured ex situ or in situ, as total oxygen uptake, diffusive or advective oxygen uptake, and which measurement device was used. The database will be curated and updated annually to secure and maintain an up-to-date global database of SCOC data.

Global Carbon Cycling on a Heterogeneous Seafloor.

Diverse biological communities mediate the transformation, transport, and storage of elements fundamental to life on Earth, including carbon, nitrogen, and oxygen. However, global biogeochemical model outcomes can vary by orders of magnitude, compromising capacity to project realistic ecosystem responses to planetary changes, including ocean productivity and climate. Here, we compare global carbon turnover rates estimated using models grounded in biological versus geochemical theory and argue that the turnover estimates based on each perspective yield divergent outcomes. Importantly, empirical studies that include sedimentary biological activity vary less than those that ignore it. Improving the relevance of model projections and reducing uncertainty associated with the anticipated consequences of global change requires reconciliation of these perspectives, enabling better societal decisions on mitigation and adaptation.

Ecosystem engineering creates a direct nutritional link between 600-m deep cold-water coral mounds and surface productivity.

Cold-water corals (CWCs) form large mounds on the seafloor that are hotspots of biodiversity in the deep sea, but it remains enigmatic how CWCs can thrive in this food-limited environment. Here, we infer from model simulations that the interaction between tidal currents and CWC-formed mounds induces downwelling events of surface water that brings organic matter to 600-m deep CWCs. This positive feedback between CWC growth on carbonate mounds and enhanced food supply is essential for their sustenance in the deep sea and represents an example of ecosystem engineering of unparalleled magnitude. This 'topographically-enhanced carbon pump' leaks organic matter that settles at greater depths. The ubiquitous presence of biogenic and geological topographies along ocean margins suggests that carbon sequestration through this pump is of global importance. These results indicate that enhanced stratification and lower surface productivity, both expected consequences of climate change, may negatively impact the energy balance of CWCs.

Temporal dynamics in a shallow coastal benthic food web: Insights from fatty acid biomarkers and their stable isotopes.

We investigated the temporal variation of pelagic and benthic food sources in the diet of benthic taxa at a depositional site in the Southern Bight of the North Sea by means of fatty acid (FA) biomarkers and compound-specific stable isotope analysis (CSIA). The taxa were the non-selective deposit feeding nematodes (Sabatieria spp. and 'other nematodes'), and three dominant macrobenthic species: two true suspension-deposit feeders (the bivalve Abra alba and the tube dwelling polychaete Owenia fusiformis) and the suspected predatory mud-dwelling anemone Sagartia sp. These species make up on average 16% (Abra alba), 17% (Sagartia sp.) and 20% (Owenia fusiformis) of the biomass in the Abra alba-Kurtiella bidentata community in this area. Phytoplankton dynamics in the suspended particulate matter of the water column as inferred from cell counts, chlorophyll-a and organic carbon content were clearly visible in sediment and animal FA abundance as well, whereas phytodetritus dynamics in the sediment FA composition were less clear, probably due to patchy distribution or stripping of FA by macrofauna. Nematodes appeared to assimilate mainly Polyunsaturated Fatty Acids (PUFAs) from their sedimentary environment and were further non-selectively accumulating more (Sabatieria spp.) or less ('other nematodes') FA from the deposited phytodetritus. In contrast, Abra alba FA composition was consistent with a diatom-dominated diet and consumption of Phaeocystis was observed in Owenia fusiformis, whereas Sagartia sp. showed evidence of a predatory behaviour. While the total FA content in Owenia fusiformis remained constant throughout the year, Sagartia sp. doubled and Abra alba increased its FA level more than 10-fold in response to the organic matter deposition from the phytoplankton bloom. This leads to the conclusion that there is no resource partitioning between non-selective deposit feeding nematodes and the suspension-deposit feeding macrobenthic organisms, suggesting they belong to separate parts of the benthic food web.

Empirical evidence reveals seasonally dependent reduction in nitrification in coastal sediments subjected to near future ocean acidification.

Research so far has provided little evidence that benthic biogeochemical cycling is affected by ocean acidification under realistic climate change scenarios. We measured nutrient exchange and sediment community oxygen consumption (SCOC) rates to estimate nitrification in natural coastal permeable and fine sandy sediments under pre-phytoplankton bloom and bloom conditions. Ocean acidification, as mimicked in the laboratory by a realistic pH decrease of 0.3, significantly reduced SCOC on average by 60% and benthic nitrification rates on average by 94% in both sediment types in February (pre-bloom period), but not in April (bloom period). No changes in macrofauna functional community (density, structural and functional diversity) were observed between ambient and acidified conditions, suggesting that changes in benthic biogeochemical cycling were predominantly mediated by changes in the activity of the microbial community during the short-term incubations (14 days), rather than by changes in engineering effects of bioturbating and bio-irrigating macrofauna. As benthic nitrification makes up the gross of ocean nitrification, a slowdown of this nitrogen cycling pathway in both permeable and fine sediments in winter, could therefore have global impacts on coupled nitrification-denitrification and hence eventually on pelagic nutrient availability.

Trophic dynamics of deep-sea megabenthos are mediated by surface productivity.

Most deep-sea benthic ecosystems are food limited and, in the majority of cases, are driven by the organic matter falling from the surface or advected downslope. Species may adapt to this scarceness by applying a wide variety of responses, such as feeding specialisation, niche width variation, and reduction in metabolic rates. The Mediterranean Sea hosts a gradient of food availability at the deep seafloor over its wide longitudinal transect. In the Mediterranean, broad regional studies on trophic habits are almost absent, and the response of deep-sea benthos to different trophic conditions is still speculative. Here, we show that both primary and secondary production processes taking place at surface layers are key drivers of deep-sea food web structuring. By employing an innovative statistical tool, we interpreted bulk-tissue δ(13)C and δ(15)N isotope ratios in benthic megafauna, and associated surface and mesopelagic components from the 3 basins of the Mediterranean Sea at 3 different depths (1200, 2000, and 3000 m). The trophic niche width and the amplitude of primary carbon sources were positively correlated with both primary and secondary surface production indicators. Moreover, mesopelagic organic matter utilization processes showed an intermediate position between surface and deep benthic components. These results shed light on the understanding of deep-sea ecosystems functioning and, at the same time, they demand further investigation.

Meiofauna metabolism in suboxic sediments: currently overestimated.

Oxygen is recognized as a structuring factor of metazoan communities in marine sediments. The importance of oxygen as a controlling factor on meiofauna (32 µm-1 mm in size) respiration rates is however less clear. Typically, respiration rates are measured under oxic conditions, after which these rates are used in food web studies to quantify the role of meiofauna in sediment carbon turnover. Sediment oxygen concentration ([O(2)]) is generally far from saturated, implying that (1) current estimates of the role of meiofauna in carbon cycling may be biased and (2) meiofaunal organisms need strategies to survive in oxygen-stressed environments. Two main survival strategies are often hypothesized: 1) frequent migration to oxic layers and 2) morphological adaptation. To evaluate these hypotheses, we (1) used a model of oxygen turnover in the meiofauna body as a function of ambient [O(2)], and (2) performed respiration measurements at a range of [O(2)] conditions. The oxygen turnover model predicts a tight coupling between ambient [O(2)] and meiofauna body [O(2)] with oxygen within the body being consumed in seconds. This fast turnover favors long and slender organisms in sediments with low ambient [O(2)] but even then frequent migration between suboxic and oxic layers is for most organisms not a viable strategy to alleviate oxygen limitation. Respiration rates of all measured meiofauna organisms slowed down in response to decreasing ambient [O(2)], with Nematoda displaying the highest metabolic sensitivity for declining [O(2)] followed by Foraminifera and juvenile Gastropoda. Ostracoda showed a behavioral stress response when ambient [O(2)] reached a critical level. Reduced respiration at low ambient [O(2)] implies that meiofauna in natural, i.e. suboxic, sediments must have a lower metabolism than inferred from earlier respiration rates conducted under oxic conditions. The implications of these findings are discussed for the contribution of meiofauna to carbon cycling in marine sediments.