Vertical distribution of benthic community responses to fish predators, and effects on algae and suspended material
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The vertical positioning of benthic invertebrates should be a trade-off between the risky, but productive, sediment surface and the safer, but physiologically harsher, conditions deeper down in the sediment. This is because the foraging efficiency of benthic fish decreases with sediment depth, whereas the sediment surface is generally better oxygenated and has a higher resource quality than lower layers. We studied how two benthic fish predators, bream (Abramis brama) and ruffe (Gymnocephalus cernuus), affected the community composition and vertical distribution of benthos, and their indirect effects on algae and suspended material, in field enclosures. Whereas bream had significant effects on the density, composition and distribution of the benthos, ruffe had no such effects. The total benthos biomass in bream treatments was an-order of magnitude lower in the upper sediment layer (0–1 cm) and three times lower in the middle layer (1–3 cm) than in the controls, whereas there were no significant effects in the deepest layer (3–10 cm). Bivalves persisted in the deepest layer although their density was reduced in shallow sediment, whereas gastropods faced the risk of local extinction in the presence of bream. As indirect effects, small-bodied cladocerans, phytoplankton, periphyton and both organic and inorganic suspended material were higher in the bream treatments. We␣conclude that the impact of bream diminished substantially with increasing sediment depth, enabling invertebrates to survive in the sediment and to persist in the presence of bream. However, there were␣no␣indications of any group adjusting their vertical position behaviourally as a response to predation threat.

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