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.

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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.

The contrasting effects of the invasive Marenzelleria viridis and the native Nereis diversicolor on benthic metabolism, partitioning of reaction pathways and distribution of inorganic porewater (C and N) solutes in homogenized sandy sediment were investigated experimentally over a period of 1 mo. The 2 species were studied separately and in combination to observe possible effects and interactions. Benthic O 2 uptake and total CO 2 (TCO 2 ) release were affected similarly by M. viridis, N. diversicolor and the two in combination, with roughly a doubling after 1 to 2 wk compared to defaunated sediment. Sulfate reduction after 1 mo, on the other hand, was more than twice as high in sediment inhabited by M. viridis alone than in any other treatment, even when combined with N. diversicolor. Denitrification estimated from benthic TCO 2 release, porewater reaction stoichiometry and nutrient fluxes was largely unaffected by the presence of fauna. Accordingly, the partitioning of reaction pathways after 1 mo revealed that M. viridis stimulated sulfate reduction at the expense of aerobic respiration. Most of the oxygen uptake in M. viridis sediment was apparently due to enhanced oxidation resulting from an upward drifting front of sulfide as indicated by low redox and the appearance of Beggiatoa sp. near the surface. Porewater solute profiles showed that M. viridis was capable of stronger and deeper irrigation than N. diversicolor despite ~10 times higher burrow ventilation by the latter species. This effect was caused by percolation of return water in the deep (> 20 cm) I-or J-shaped burrows of M. viridis compared to the flushing of the more shallow (6 to 8 cm) U-shaped burrows of N. diversicolor. A replacement of the native N. diversicolor with the invasive M. viridis as the dominant burrow-dwelling polychaete in shallow coastal sediments will probably affect the biogeochemical functioning and ecological stability of the ecosystem. Among other things, organisms tolerant to sulfide are likely to be favored at the expense of more sensitive species.

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