Keeping things in order: multivariate direct gradient analysis of a strongly fluctuating benthic community

Two spatial surveys of the macrobenthos of an estuarine intertidal area, the Oosterschelde in the southwestern part of The Netherlands, were conducted in different years. In the period between the surveys changes in the hydrodynamics of the estuary took place as a result of the construction of a storm-surge barrier. The works reduced the tidal volume in the basin and as a consequence the altitude of the tidal flats decreased by approximately 0.1 to 0.2 m. Considerable differences in species densities were found between the two surveys, but these large differences could not be explained by changes in the `spatialʹ environmental variables (such as altitude) that were taken into account in a direct gradient analysis. Unknown and `only-to-time-relatedʹ (e.g. weather-related) factors had considerably influenced species abundance. This implies that the impact of the works on the macrobenthos could not be evaluated on the basis of observed species densities in the before- and after-the-works surveys. Generally, it implies that even if the levels of future `spatialʹ environmental variables are known, direct gradient analysis based on a single survey cannot be used for prediction of future densities in absolute terms in such strongly fluctuating communities. Yet, the gradient analyses for the two years revealed rather similar estimates of the major environmental gradients. The same was true for the parameters that described the relative change in species densities in response to a change in the environmental gradient levels. Hence the most appropriate way of evaluating the changes that have occurred in the benthic community as a result of the works was in terms of the observed changes in the `spatialʹ environmental gradient levels. The levels of the first environmental gradient, which was strongly related to the sediment structure, did not change between years. Levels of the second gradient, which could be interpreted as a tidal zone gradient, decreased. This suggests that the works particularly affected taxa that are strongly related to this second gradient, such as Corophium living in the upper tidal areas and Nephthys living in the lower tidal zone.