The benthoneuston of the Black Sea: Composition and environmental factors influencing its nocturnal dynamic

Plankton fauna nocturnally migrating from the sea-floor or near-bottom layer to the uppermost surface layer (benthoneuston) links benthic, benthopelagic, pelagic, and neustonic realms. We conducted five intervals of sampling (every 1-2 h during five nights) synchronously in the neustal (surface to 10-cm depth layer) and in the water column below to examine which taxa concentrate in the neustal, and under which circumstances they do so. We tested the following environmental factors: sea-floor biotope type, temperature, time after sunset, time to midnight, moon phase, and moon altitude. Of the 77 taxa recorded, about half (41) were found in the neustal. Among these, less than half (16) of the taxa showed a quantified attraction (by L-index) to the neustal and may be called the true benthoneuston. In contrast to the benthopelagic zone, where the contribution of the characteristic benthopelagic fauna exceeded 50%, the neustal was not dominated by a specific benthoneuston fauna. Nocturnal dynamics of the benthoneuston was mainly controlled by the proximity of the twilight time, then by the sea-floor biotope type and time to midnight. Neustonic taxa were more affected by moon illumination (moon phase and moon altitude) than those in the water column below. The benthoneuston in the studied area was represented by either juveniles or reproducing adults. This component of plankton communities is thus temporary and seasonal, at least in the temperate Black Sea. In the "high" summer-autumn season, contribution of benthoneuston to the coastal plankton communities is significant, whilst in winter this contribution may be negligible. The next step in the understanding of the role of benthoneuston should be associated with tropical areas where seasonal changes in reproduction are less distinctive and this group may represent an important permanent component of coastal communities.