The term 'bioturbation' is frequently used to describe how living organisms affect the substratum in which they live. A closer look at the aquatic science literature reveals, however, an inconsistent usage of the term with increasing perplexity in recent years. Faunal disturbance has often been referred to as particle reworking, while water movement (if considered) is re ferred to as bioirrigation in many cases. For consistency, we therefore propose that, for contemporary aquatic scientific disciplines, faunal bioturbation in aquatic environments includes all transport processes carried out by animals that directly or indirectly affect sediment matrices. These processes include both particle reworking and burrow ventilation. With this definition, bioturbation acts as an 'umbrella' term that covers all transport processes and their physical effects on the substratum. Particle reworking occurs through burrow construction and maintenance, as well as ingestion and defecation, and causes biomixing of the substratum. Organic matter and microorganisms are thus displaced vertically and laterally within the sediment matrix. Particle reworking animals can be categorized as biodiffusors, upward conveyors, downward conveyors and regenerators depending on their behaviour, life style and feeding type. Burrow ventilation occurs when animals flush their open-or blind-ended burrows with overlying water for respiratory and feeding purposes, and it causes advective or diffusive bioirrigation ex change of solutes between the sediment pore water and the overlying water body. Many bioturbating species perform reworking and ventilation simultaneously. We also propose that the effects of bioturbation on other organisms and associated processes (e.g. microbial driven biogeochemical transformations) are considered within the conceptual framework of ecosystem engineering.
The effects of organic enrichment on sediment biogeochemistry was studied in diffusion controlled sediment mesocosms, where labile organic matter (OM) (fish feed) pulses were added once a week to the sediment surface. Two types of sediments, differing mainly in content of reactive Fe, were used. The aim of this experiment was two-fold, (1) to evaluate the importance of Fe-driven sulfide buffering for sulfide accumulation in surface enriched sediments, and (2) to estimate the diagenetic capacity for degradation of labile OM near the sediment surface. The simulated OM loading rate of 375 mmol C m -2 day -1 led to a 5-6 times increase in CO 2 -production and a 4-5 times increase in O 2 -uptake. Sulfate reduction estimated by radiotracer experiments and CO 2 -release was 105-131 mmol m -2 day -1 , but accumulation of porewater sulfide was low in both sediment types. Instead 99% of sulfide was oxidized with O 2 at the sediment water interface in the low Fe treatment, whereas 46% of produced sulfide precipitated as Fe-S compound in the high Fe treatment resulting in significantly lower O 2 -uptake. Furthermore, the accumulation of up to 30% of added OM by the end of the experiment indicated a saturation of the heterotrophic microbial communities in the upper enriched surface layer. These results suggest a maximum diagenetic capacity for OM degradation in the range of *25 lmol C cm -3 day -1 or 260 mmol m -2 day -1 for the present sediment types.
We aim at identifying how ecosystem functioning in shallow estuaries is affected by rapid shifts in benthic fauna communities. We use the shallow estuary, Odense Fjord, Denmark, as a case study to test our hypotheses that (1) shifts in benthic fauna composition and species functional traits affect biogeochemical cycling with cascading effects on the ecological functioning, which may (2) modulate primary productivity in the overlying water column with feedbacks to the benthic system. Odense Fjord is suitable because it experienced dramatic shifts in benthic fauna community structure from 1998 to 2008. We focused on infaunal species composition and functional traits with emphasis on three dominating burrow-dwelling polychaetes: the native Nereis (Hediste) diversicolor and Arenicola marina, and the invasive Marenzelleria viridis. The impact of functional traits in the form of particle reworking and ventilation on biogeochemical cycles, i.e., sediment metabolism and nutrient dynamics, was determined from literature data. Historical records of summer nutrient levels in the water column of the inner Odense Fjord show elevated concentrations of NH + 4 and NO − 3 (DIN) during the years 2004-2006, exactly when the N. diversicolor population declined and A. marina and M. viridis populations expanded dramatically. In support of our first hypothesis, we show that excess NH + 4 delivery from the benthic system during the A. marina and M. viridis expansion period enriched the overlying water in DIN and stimulated phytoplankton concentration. The altered benthic-pelagic coupling and stimulated pelagic production may, in support of our second hypothesis, have feedback to the benthic system by changing the deposition of organic material. We therefore advice to identify the exact functional traits of the species involved in a community shift before investigating its impact on ecosystem functioning. We also recommend studying benthic community shifts in shallow environments to obtain sufficient knowledge about the drivers and controls before exploring deep-water environments.
Eelgrass coverage in Odense Fjord (Denmark) has declined by 90% since 1983, due to eutrophication and its associated pressures, and the state of low eelgrass coverage has remained stable despite 10 to 15 yr of reduced nutrient loading and improved water quality. We hypothesize that the survival of eelgrass seedlings, and thus recolonization through reproductive dispersal, is negatively affected by physical disturbances. The 3 most likely physical mechanisms involved are uprooting or burial through drifting macroalgae, Arenicola marina sediment reworking and current-driven sediment resuspension. Our hypothesis was tested by field observations during the summer of 2009, when the mortality of seedlings was followed through time. The density of seedlings decreased dramatically by 80% during the first month of observations, and no seedlings survived past August, corresponding to an average seedling mortality of 1.5% d -1. This was > 3 times higher than the mortality for seedlings protected from physical disturbance by enclosures (0.4% d -1 ), indicating that physical disturbance contributed to high seedling mortality. A significant correlation (p = 0.02) between macroalgal drift and seedling mortality suggested that ~40% of seedlings were lost due to the physical disturbance of drifting algae. In contrast, no correlations were found between A. marina reworking or resuspension and seedling mortality, despite a mobility of up to 400 cm 3 sediment m -2 d -1 by these mechanisms. Given the observed intensity of macroalgal drift, we speculate that this mechanism severely hampers eelgrass reestablishment in certain parts of Odense Fjord.KEY WORDS: Eelgrass · Macroalgal drift · Bedload transport · Fucus sp. · Sediment · Reworking · Resuspension · Arenicola marina Resale or republication not permitted without written consent of the publisherMar Ecol Prog Ser 418: [119][120][121][122][123][124][125][126][127][128][129][130] 2010 elongation (Olesen & Sand-Jensen 1994, Townsend & Fonseca 1998, Boese et al. 2009). Recolonization of larger areas, therefore, depends primarily on seed dispersal and the subsequent growth of seedlings (Olesen & Sand-Jensen 1994, Plus et al. 2003, Greve et al. 2005, Jarvis & Moore 2010. Z. marina has a high capacity for sexual reproduction, whereby seeds are released during late summer and fall, resulting in densities of from 10 2 to 10 4 seeds m -2 inside eelgrass beds and up to 30-50 seeds m -2 at a distance of 15 m from established beds (Harrison 1993, Olesen & Sand-Jensen 1994, Orth et al. 1994, Olesen 1999, Probert & Brenchley 1999, Boese & Robbins 2008. However, only 5 to 15% of seeds germinate to produce seedlings the following spring (Harrison 1993, Orth et al. 2003, resulting in low seedling density (0.2 to 26 m -2 ) outside established beds (Olesen & Sand-Jensen 1994, Greve et al. 2005, Boese & Robbins 2008. Hence, net expansion of eelgrass coverage depends primarily on the survival of a low number of seedlings to form new patches and eventually to create a continuous seagra...
Global expansion of salmon aquaculture is contingent upon finding new, large-capacity farming locations that are, increasingly, situated in dispersive environments with atypical ecological characteristics. The capacity of such sites to assimilate organic waste and the type and spatial extent of effects remain poorly understood. This study couples intensive spatial and temporal measurements of waste outputs with measurements of benthic ecological diversity and sediment biogeochemistry at a shallow dispersive site on the central west coast of Norway. Despite minimal visual changes to the seabed, pronounced biological effects were evident out to 600−1000 m away from the farm. Greatly enhanced faunal and microbial activity was tightly coupled with strong sediment respiratory responses in the form of oxygen uptake and ammonia efflux. The benthic response was highly dynamic, with rapid and prolific colonization by opportunistic fauna, followed by substantive recovery during the subsequent 7 mo fallowing period. The fate and pathways of farm waste through the environment was compartmentalized by converting measured parameters into equivalent carbon fluxes. During early production, approximately 30−40% of the waste was accounted for by the measured benthic processes, attributed to faunal respiratory activity and the physical properties including sediment type. Later in the production cycle, a sediment burial event was observed down-current from the farm. The remaining 60−70% of organic waste was assumed to be either assimilated in the water column, consumed by large, unquantified benthic fauna, or exported and dispersed. Shallow, dispersive sites therefore appear relatively resilient to acute near-field enrichment, but are also more likely to result in accumulative, far-field effects.
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