Catalogus

A continuum of assemblages existed along the salinity gradient from freshwater river to the North Sea, with shifts in the ranges of organisms apparent in response to changes in freshwater flow. This pattern closely fits an ecocline model. However, the estuary differs from previously defined ecoclines in having two overlapping gradients in the major stressor: from river to mid-estuary for freshwater species and from sea to mid-estuary for marine species. We propose, therefore, that the estuary represents a two-ecocline model, with fauna inhabiting the mid-estuary being either freshwater or marine species at the edge of their range, rather than ‘ true estuarine organisms ’. This allows a redefinition of the Remane diagram, with estuarine species removed, and supports previous arguments that brackish-water species do not exist. Such two-ecocline models may also exist in other marine systems, such as rocky shores.