Properties of ecotones: Evidence from five ecotones objectively determined from a coastal vegetation gradient

Several properties have been suggested to be characteristic of ecotones, but their prevalence has rarely been tested. We sampled five ecotones to seek evidence on seven generalizations that are commonly made about ecotones: vegetational sharpness, physiognomic change, occurrence of a spatial community mosaic, many exotic species, ecotonal species, spatial mass effect, and species richness higher or lower than either side of the ecotone. The ecotones were in a sequence from scattered mangroves, through salt marsh, rush-marsh, scrub, woodland, to pasture. We developed a method to objectively define, by rapid vegetational change, the position and depth of an ecotone, identifying five ecotones. Their positions were consistent across three sampling schemes and two spatial grain sizes. One ecotone is a switch ecotone, produced by positive feedback between community and environment. Another is anthropogenic, due to clearing for agriculture. Two others are probably environmental in cause, and one may be largely a relict environmental ecotone. Sharp changes in species composition occurred. Three ecotones were associated with a change in plant physiognomy. In two, the ecotone was located just outside a woodland canopy, in the zone influenced by the canopy. Community mosaicity was evident at only one ecotone. There were few strictly ecotonal species; many species occurred more frequently within ecotones than in adjacent vegetation, but there were never significantly more ecotonal species than expected at random. There was little evidence for the spatial mass effect reducing ecotonal sharpness, or leading to higher species richness within ecotones. Species richness was higher than in the adjacent habitat in only one ecotone. It seems that supposedly characteristic ecotone features depend on the particular ecological situation, and the ecology of the species present, rather than being intrinsic properties of ecotones.