Biodiversity of benthic invertebrates and organic matter processing in shallow marine sediments: an experimental study

The main objective of this study was to measure the impact of benthic invertebrate diversity on processes occurring at the water-sediment interface. We analyzed the effects of interactions between three shallow water species (Cerastoderma edule, Corophium volutator, and Nereis diversicolor). The impacts of different species richness treatments were measured on sediment reworking, bacterial characteristics, and biogeochemical processes (bromide fluxes, O₂ uptake, nutrient fluxes, and porewater chemistry) in sediment cores. The results showed that the three species exhibited different bioturbation activities in the experimental system: C. edule acted as a biodiffusor, mixing particles in the top 2 cm of the sediments; C. volutator produced and irrigated U-shaped tubes in the top 2 cm of the sediments; and N. diversicolor produced and irrigated burrow galleries in the whole sediment cores. C. edule had minor effects on biogeochemical processes, whereas the other species, through their irrigation of the burrows, increased the solute exchange between the water column and the sediment two-fold. These impacts on sediment structure and solute transport increased the O₂ consumption and the release of nutrients from sediments. As N. diversicolor burrowed deeper in the sediment than C. volutator, it irrigated a greater volume of sediments, with great impact on the sediment cores.Most treatments with a mixture of species indicated that observed values were often lower than predicted values from the addition of the individual effects of each species, demonstrating a negative interaction among species. This type of negative interaction measured between species on ecosystem processes certainly resulted from an overlap of bioturbation activities among the three species which lived and foraged in the same habitat (water-sediment interface). All treatments with N. diversicolor (in isolation and in mixture) produced similar effect on sediment reworking, water fluxes, nutrient releases, porewater chemistry, and bacterial characteristics. Whichever species associated with N. diversicolor, the bioturbation activities of the worm hid the effect of the other species. The results suggest that, in the presence of several species that use and modify the same sediment space, impact of invertebrates on ecosystem processes was essentially due to the most efficient bioturbator of the community (N. diversicolor). In consequence, the functional traits (mode of bioturbation, depth of burrowing, feeding behaviour) of an individual species in a community could be more important than species richness for some ecosystem processes.     

Fish Reliance on Littoral–Benthic Resources and the Distribution of Primary Production in Lakes

Pelagic, littoral, and terrestrial resources can all play a role in supporting consumers in lakes. The role of benthic algal-derived food web pathways in lakes is perhaps the least understood because limnologists have historically focused on pelagic (open-water) production and processes. We compiled carbon stable isotope data from 546 fish populations (75 lakes), and used a two end-member mixing model to calculate littoral–benthic reliance for each fish species in each lake. Fish littoral–benthic reliance values were averaged by lake to assess overall fish species benthic reliance for each lake. Lake-specific mean littoral reliance (BR_L; fish species not weighted according to production or biomass) averaged 57% and was independent of lake morphological and limnological attributes. For these same lakes, water column nutrients, light, and morphometry data were used to estimate whole-lake benthic algal and phytoplankton primary production. On average, benthic algae comprised 36% of whole-lake primary production (BP_f = 0.36). BP_f and BR_L were weakly correlated: BR_L tends to be high even in large/deep lakes in which benthic algae is a minor contributor to whole-lake primary production. The high littoral–benthic contribution to individual fish species appears to reflect the high concentration of fish species diversity in the littoral zone. Our work cannot be extrapolated to whole-lake fish production. However, the result is consistent with other work indicating that most fish species inhabit the littoral zone, whereas relatively few exclusively inhabit the pelagic. Our results suggest that it takes less primary production to support a single fish species in the littoral zone than is required to support a species in the pelagic.     

Ecological consequences of invasion across the freshwater–marine transition in a warming world

The freshwater–marine transition that characterizes an estuarine system can provide multiple entry options for invading species, yet the relative importance of this gradient in determining the functional contribution of invading species has received little attention. The ecological consequences of species invasion are routinely evaluated within a freshwater versus marine context, even though many invasive species can inhabit a wide range of salinities. We investigate the functional consequences of different sizes of Corbicula fluminea—an invasive species able to adapt to a wide range of temperatures and salinity—across the freshwater–marine transition in the presence versus absence of warming. Specifically, we characterize how C. fluminea affect fluid and particle transport, important processes in mediating nutrient cycling (NH₄‐N, NO₃‐N, PO₄‐P). Results showed that sediment particle reworking (bioturbation) tends to be influenced by size and to a lesser extent, temperature and salinity; nutrient concentrations are influenced by different interactions between all variables (salinity, temperature, and size class). Our findings demonstrate the highly context‐dependent nature of the ecosystem consequences of invasion and highlight the potential for species to simultaneously occupy multiple components of an ecosystem. Recognizing of this aspect of invasibility is fundamental to management and conservation efforts, particularly as freshwater and marine systems tend to be compartmentalized rather than be treated as a contiguous unit. We conclude that more comprehensive appreciation of the distribution of invasive species across adjacent habitats and different seasons is urgently needed to allow the true extent of biological introductions, and their ecological consequences, to be fully realized.     

Rapid reworking of subtidal sediments by burrowing spatangoid urchins

The mixing and displacement of sediment by benthic macrofauna (bioturbation) has major biogeochemical implications, and can control rates of organic matter degradation and carbon burial. Large, abundant, mobile macrofauna often dominate sediment bioturbation, and heart urchins of the genus Echinocardium are regarded as key sediment bioturbators in marine systems throughout the world. To better understand the bioturbation potential and functional role of Echinocardium, we developed a mathematical model and parameterized it with field data from six locations in northern New Zealand in order to estimate bioturbation rates in these places. Although urchin sizes and densities were measured in consecutive years at all six locations, we obtained a third model parameter, urchin movement rate, from one time and place only (Site OB5). Because confidence in model output was greatest at OB5, and since OB5 had the highest sediment reworking rate of all sites, our model yielded a good upper bound estimate for the bioturbation potential of Echinocardium in the areas examined. The volume of sediment displaced by Echinocardium populations reached 20,000 cm³ m⁻² d⁻¹ at OB5, suggesting that surface sediment is reworked about every 3 days at sites where Echinocardium is abundant. Experimental work with a fluorescent tracer at OB5 suggested limited downward particle movement as a result of Echinocardium bioturbation, though vertical profiles of chlorophyll a and organic matter content indicated well mixed sediment. The loss of Echinocardium because of broad-scale anthropogenic disturbance to the seabed could have major consequences on marine ecosystem functioning.     

A cost-effective method to quantify biological surface sediment reworking

We propose a simple and inexpensive method to determine the rate and pattern of surface sediment reworking by benthic organisms. Unlike many existing methods commonly used in bioturbation studies, which usually require sediment sampling, our approach is fully non-destructive and is well suited for investigating non-cohesive fine sediments in streams and rivers. Optical tracer (e.g. luminophores or coloured sand) disappearance or appearance is assessed through time based on optical quantification of surfaces occupied by tracers. Data are used to calculate surface sediment reworking (SSR) coefficients depicting bioturbation intensities. Using this method, we evaluated reworking activity of stream organisms (three benthic invertebrates and a fish) in laboratory microcosms mimicking pool habitats or directly in the field within arenas set in depositional zones. Our method was sensitive enough to measure SSR as low as 0.2 cm² day^?1, such as triggered by intermediate density (774 m^?2) of Gammarus fossarum (Amphipoda) in microcosms. In contrast, complex invertebrate community in the field and a fish (Barbatula barabatula) in laboratory microcosms were found to yield to excessively high SSR (>60 cm² day^?1). Lastly, we suggest that images acquired during experiments can be used for qualitative evaluation of species-specific effects on sediment distribution.     

Influence of benthic macrofauna on microbial production of methylmercury in sediments on the New England continental shelf

Microbial methylation processes in sediment are an important source of toxic monomethylmercury (MMHg) to aquatic ecosystems. Although bioturbation activities (feeding, digging of galleries, excavations, bioirrigation) by benthic fauna are known to affect many biogeochemical processes, their influence on benthic MMHg production is poorly understood. We investigated the effect of benthic fauna on the microbial production of MMHg in sediments on the continental shelf of the northwest Atlantic Ocean in September 2009. Replicate cores of sieved (control) and unaltered sediment containing native macrofauna were incubated to examine the influence of benthic macrofauna on net MMHg production, potential gross rates of Hg methylation, sediment reworking, dissolved oxygen and organic carbon concentrations, and microbial metabolic activities. The presence of macrofauna stimulated aerobic microbial respiration and net MMHg production, but had no observed effect on short-term gross rates of Hg methylation. This suggests that bioturbation may promote net MMHg production by inhibiting demethylating microorganisms, although overall community metabolism was increased. Results from this work emphasize the need to enhance our knowledge and understanding of the interactions among benthic fauna, microorganisms, and geochemistry in affecting MMHg production.     

The effect of bioturbation by polychaetes (Opheliidae) on benthic foraminiferal assemblages and test preservation

Biological activity such as burrowing can alter benthic foraminiferal shell preservation and may also modify benthic foraminiferal assemblages by vertical mixing, inducing sediment homogenization. Here, we analyse benthic foraminiferal assemblages and taphonomy of upper Miocene marine deposits from Conil de la Frontera (Cádiz, south‐western Spain). The deposits consist of marls displaying a pervasive alternation of intensively bioturbated beds dominated by Macaronichnus segregatis traces (ichnofabric index 4–5) and non‐bioturbated beds. Benthic foraminiferal assemblages are dominated by Cibicidoides mundulus and Cibicides refulgens, indicating that the marls were deposited on an oligotrophic, well‐oxygenated upper slope. The impact of burrowing on the preservation of benthic foraminiferal tests was tested using Q‐mode cluster analysis, which found two well‐differentiated groups of samples, one including the non‐bioturbated beds and the other encompassing the bioturbated ones. Fragmentation and recrystallization account for the differentiation of these groups, both being higher in the bioturbated sediments. Aggressive chemical digestion by the Macaronichnus trace‐makers, assumed to be a polychaete worm of the family Opheliidae, etched the microfossil shells, making them more vulnerable to fragmentation. Intense bioturbation favoured the circulation of pore fluids, encouraging recrystallization. Pervasive burrowing resulted in significant vertical reworking of microfossils. As a consequence, benthic foraminiferal assemblages in the bioturbated beds were homogenized in the mixed layer; that is, the uppermost layer of the substrate totally burrowed. The alternation of bioturbated and non‐bioturbated beds reflects episodic transfer of food particles down slope from shallower parts of the shelf as well as from the continent due to storms under otherwise homogeneous oligotrophic marine conditions.     

Classification of Reynolds phytoplankton functional groups using individual traits and machine learning techniques

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Breaking the boundary — The key to bottom recovery? The role of mysid crustaceans in oxygenizing bottom sediments

Large areas of the Baltic Sea bottoms suffer from low oxygen conditions and anoxia, impoverishing the benthic macrofauna. The important macrofaunal function bioturbation, which improves the transport of oxygen into the sediment does not occur in an absence of benthic macrofauna. The objective of this study was to investigate if a semi-pelagic species, like the mysid crustacean Mysis relicta, is able to improve the oxygen conditions of the sediment and thereby acts as a facilitator for re-colonization of azoic sediments by benthic species. We also wanted to study the potential of M. relicta in breaking the diffusive boundary layer under varying degrees of oxygen deficiency. Three types of sediment qualities were used to mimic the severity of oxygen deficiency. Under normoxia, moderate hypoxia (40% O₂) and hypoxia, (20% O₂) M. relicta's bioturbation activity was studied by recording oxygen profiles in sediments with and without mysids. In normoxia the mysids were able to oxygenize the sediment independent of sediment quality. The results show that mysids are able to bioturbate the sediment to some extent in hypoxia independent of the sediment quality. In all treatments with mysids the diffusive boundary layer was more or less completely broken down. In normoxia treatment with sediment of very low quality the mysids prevented growth of the sulphur bacteria Beggiatoa spp. which usually occurs on anoxic bottoms. The ability of this semi-pelagic species to improve benthic oxygen conditions can be seen as an important first step in re-colonization by real benthic species.     

Food supply impacts sediment reworking by <Emphasis Type="Italic">Nereis diversicolor</Emphasis>

This study assessed the effects of food supply on sediment reworking by Nereis diversicolor. We hypothesized that food supply would enhance sediment reworking and that the frequency of food supply would affect the intensity of bioturbation. Mesocosm experiments consisted of four treatments: (1) without worms (control cores), (2) with worms and no food supply, (3) with worms and daily food supply, (4) with worms and weekly food supply. Fluorescent particles, used as tracers, were spread over the sediment surface. Sediment reworking was quantified after 28 days based on the tracer distribution profiles. Results showed that sediment reworking by N. diversicolor was exclusively due to non-local transport processes. Food supply greatly increased non-local transport coefficients (more than 3 times) in comparison with those measured in the absence of a food supply. However, the intensity of sediment bioturbation by these worms was unaffected by the frequency of food supply. This study showed that environmental conditions affecting the quantity of food supply at the water-sediment interface could strongly influence bioturbation process. Handling editor: P. Viaroli     

Growth and uptake kinetics of phosphate by benthic microalga Nitzschia sp. isolated from Hiroshima Bay,Japan

In the present study, we experimentally investigated the phosphate uptake kinetics of benthic microalga Nitzschia sp. isolated from Hiroshima Bay, Japan. The maximum uptake rate (ρ_max) obtained by short‐term experiments was 6.84 pmol cell^?1 h^?1 for phosphate. The half‐saturation constant for uptake (K_S) was 61.2 µmol cell^?1 h^?1. Both the ρ_max and Ks of this species were extremely high, suggesting that Nitzschia sp. is adapted to benthic environments, where nutrient concentrations are much higher than in the water column. The specific maximum growth rate (µ'_max) and minimum cell quota (Q₀) for the P‐limited condition, obtained by a semi‐continuous growth experiment, were 0.48 day^?1 and 0.045 pmol cell^?1, respectively. It is concluded that Nitzschia sp. could be a ‘storage strategist’ species, meaning it adapts so as to minimize the influence of fluctuations in phosphate conditions resulting from the change in redox conditions of sediment due to bioturbation.     

生物扰动对沉积物中污染物环境行为的影响研究进展

生物扰动由于显著改变沉积物结构和性质,进而影响沉积物中污染物的环境行为。综述生物扰动对沉积物中氮、磷、重金属和疏水性有机污染物环境行为的影响。生物扰动促进这些污染物从沉积物向水体释放。生物扰动还对不同的污染物产生其它不同的影响。对于氮,生物扰动还影响其硝化与反硝化作用;对于磷,生物扰动不仅改变其化学形态,还提高有机磷降解。对于重金属,生物扰动还能改变其在沉积物中的分布及化学形态。对于疏水性有机污染物,生物扰动主要增强生物富集和代谢,以及提高生物降解。     

Rare but large bivalves alter benthic respiration and nutrient recycling in riverine sediments

Bioturbation studies have generally analyzed small and abundant organisms while the contribution to the benthic metabolism by rare, large macrofauna has received little attention. We hypothesize that large, sporadic bivalves may represent a hot spot for benthic processes due to a combination of direct and indirect effects as their metabolic and bioturbation activities. Intact riverine sediments with and without individuals of the bivalve Sinanodonta woodiana were collected in a reach with transparent water, where the occurrence of the mollusk was clearly visible. The bivalve metabolism and its effects on sedimentary fluxes of dissolved gas and nutrients were measured via laboratory incubations of intact cores under controlled conditions. S. woodiana contributed significantly to O₂ and TCO₂ benthic fluxes through its respiration and to \({\text{NH}}_{4}^{ + }\), SRP and SiO₂ regeneration via its excretion. The bivalve significantly stimulated also microbial denitrification and determined a large efflux of CH₄, likely due a combination of bioturbation and biodeposition activities or to anaerobic metabolism within the mollusk gut. This study demonstrates that a few, large individuals of this bivalve produce significant effects on aerobic and anaerobic benthic metabolism and nutrient mobilization. Random sediment sampling in turbid waters seldom catches these important effects due to low densities of large fauna.     

Invertebrate bioturbation can reduce the clogging of sediment: an experimental study using infiltration sediment columns

1. Invertebrate bioturbation can strongly affect water‐sediment exchanges in aquatic ecosystems. The objective of this study was to quantify the influence of invertebrates on the physical characteristics of an infiltration system clogged with fine sediment. 2. Two taxa (chironomids and tubificids) with different bioturbation activities were studied in experimental slow infiltration columns filled with sand and gravel and clogged with a 2 cm layer of fine sediment at the surface. We measured the effects of each taxon separately and combined on hydraulic head, water mobility and sediment reworking. 3. The results showed that invertebrates could reduce sediment clogging and this effect was linked to the functional mode of bioturbation of each group. Tubificid worms dug networks of galleries in the fine sediment, creating pathways for water flow, which reduced the clogging of sediment. In contrast, the U‐shaped tubes of chironomids were restricted to the superficial layer of fine sediments and did not modify the hydraulic conductivity of experimental columns. The combination of invertebrates did not show any interactive effects between tubificids and chironomids. The occurrence of 80 tubificids in the combination was enough to maintain the same hydraulic conductivity that 160 worms did in monospecific treatment. 4. The invertebrates like tubificid worms can have a great benefit on functioning of clogged interfaces by maintaining high hydraulic conductivity, which contributes to increased water‐sediment exchanges and stimulates biogeochemical and microbial processes occurring in river sediments.     

Bioturbation: a fresh look at Darwin's last idea

Bioturbation refers to the biological reworking of soils and sediments, and its importance for soil processes and geomorphology was first realised by Charles Darwin, who devoted his last scientific book to the subject. Here, we review some new insights into the evolutionary and ecological role of bioturbation that would have probably amazed Darwin. In modern ecological theory, bioturbation is now recognised as an archetypal example of 'ecosystem engineering', modifying geochemical gradients, redistributing food resources, viruses, bacteria, resting stages and eggs. From an evolutionary perspective, recent investigations provide evidence that bioturbation had a key role in the evolution of metazoan life at the end of the Precambrian Era.     

A trait-based approach to assess niche overlap and functional distinctiveness between non-indigenous and native species

Our understanding of the community assembly processes acting on non-indigenous species (NIS), as well as the relationship with native species is limited, especially in marine ecosystems. To overcome this knowledge gap we here develop a trait-based approach based on the functional distinctiveness metric to assess niche overlap between NIS and native species, using high-resolution data on benthic invertebrate communities in the Baltic Sea. Our results show that NIS retain a certain degree of similarity with native species, but display one or a few singular unique traits (e.g., bioturbation ability). Furthermore, we demonstrate that community assembly processes, including both environmental filtering and limiting similarity affect NIS establishment, but that their effects may be highly context dependent, as illustrated by pronounced spatial patterns in distinctiveness. Finally, our trait-based approach provides a generic framework applicable to other areas and organisms, to better understand and address biological invasions.     

The importance of Planolites in the Cambrian substrate revolution

Early Cambrian subtidal shelf substrates were characterized by low water content and steep chemical gradients, conditions likely facilitated by the presence of microbial mats as reflected by an abundance of microbially-mediated sedimentary structures in Lower Cambrian strata. Such substrate conditions would have been unfavourable for burrowing by benthic metazoans. A combination of environmental restrictions and a lack of adaptations to vertical burrowing likely prevented most benthic metazoans from burrowing infaunally in Early Cambrian subtidal shelf substrates. The eventual acquisition of burrowing adaptations by benthic metazoans later in the Cambrian promoted an increase in the depth and intensity of bioturbation and initiated a transition toward well-hydrated substrates in which extensive infaunal activity was possible.Siliciclastic units of the Lower Cambrian succession in the White–Inyo Mountains, eastern California, contain abundant horizontal bioturbation on bedding planes, as documented by bedding plane bioturbation indices, but little vertical bioturbation, as shown by ichnofabric indices and x-radiography. Planolites, a simple horizontal trace fossil, represents the dominant type of bioturbation in these units. Planolites is found in a range of diameters, indicating that more than one species of tracemaker likely produced this type of trace. Although these Planolites do not have a vertical component, their abundance on bedding planes indicates that the activities of Planolites tracemakers had a significant impact on subtidal shelf substrates, represented by Lower Cambrian units in the White–Inyo Mountains, early in the Cambrian substrate revolution.     

Consequences of Increasing Hypoxic Disturbance on Benthic Communities and Ecosystem Functioning

Disturbance-mediated species loss has prompted research considering how ecosystem functions are changed when biota is impaired. However, there is still limited empirical evidence from natural environments evaluating the direct and indirect (i.e. via biota) effects of disturbance on ecosystem functioning. Oxygen deficiency is a widespread threat to coastal and estuarine communities. While the negative impacts of hypoxia on benthic communities are well known, few studies have assessed in situ how benthic communities subjected to different degrees of hypoxic stress alter their contribution to ecosystem functioning. We studied changes in sediment ecosystem function (i.e. oxygen and nutrient fluxes across the sediment water-interface) by artificially inducing hypoxia of different durations (0, 3, 7 and 48 days) in a subtidal sandy habitat. Benthic chamber incubations were used for measuring responses in sediment oxygen and nutrient fluxes. Changes in benthic species richness, structure and traits were quantified, while stress-induced behavioral changes were documented by observing bivalve reburial rates. The initial change in faunal behavior was followed by non-linear degradation in benthic parameters (abundance, biomass, bioturbation potential), gradually impairing the structural and functional composition of the benthic community. In terms of ecosystem function, the increasing duration of hypoxia altered sediment oxygen consumption and enhanced sediment effluxes of NH₄ ⁺ and dissolved Si. Although effluxes of PO₄ ³⁻ were not altered significantly, changes were observed in sediment PO₄ ³⁻ sorption capability. The duration of hypoxia (i.e. number of days of stress) explained a minor part of the changes in ecosystem function. Instead, the benthic community and disturbance-driven changes within the benthos explained a larger proportion of the variability in sediment oxygen- and nutrient fluxes. Our results emphasize that the level of stress to the benthic habitat matters, and that the link between biodiversity and ecosystem function is likely to be affected by a range of factors in complex, natural environments.     

Quantification of sediment reworking by the Asiatic clam Corbicula fluminea Müller, 1774

Active organisms modify the substratum in which they dwell. This process, called “bioturbation”, affects the way that biogeochemical fluxes are mediated at the substratum–water interface. In the frame of this work, the bioturbation potential of the Asiatic clam Corbicula fluminea was characterized and quantified. We measured the displacement of fluorescent particles by C. fluminea burying in a size-based experimental design in order to explore the effects of body-size on sediment reworking. Our results stress that C. fluminea belongs to the functional group of biodiffusors, and that C. fluminea can be considered as an intermediate sediment reworker. We suggest that bioturbation was mainly induced by the pedal-feeding activity of the clams. Results also showed that, though large clams induced displacement of particles deeper into the sediment, small clams showed the highest net sediment reworking activity. This result was in contrast to the initial hypothesis of biovolume as the main driver for particle displacement by bioturbating organisms. Life-history traits and specific features of pedal-feeding could explain the observed pattern.