Ecotone or ecocline: Ecological boundaries in estuaries

Two main ecological boundaries, ecotone and ecocline, have been defined in landscape ecology. At this scale, the estuary represents a boundary between rivers and the sea, but there has been no attempt to fit empirical data for estuaries to these boundary models. An extensive data set from the Thames estuary was analysed using multivariate techniques and species-range analysis, in order to investigate whether the ecocline or the ecotone model was most relevant to this estuary. Data for periods of high and low freshwater flow allowed the impact of large-scale fluctuations implicit in both models to be determined. 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. (C) 2002 Elsevier Science Ltd. All rights reserved.