Zonation of vegetation on lacustrine coastal dunes: effects of burial by sand

More than 50 years ago it was proposed that zonation of major plant species on coastal dunes was determined by salt spray. Here, we argue against this hypothesis because (i) salt concentrations rarely exceed toxic levels; (ii) high precipitation in temperate latitudes washes the salt off the plants; (iii) major salt spray events occur in late autumn and winter when plants are dormant; and (iv) zonation also occurs on lacustrine dunes. Instead, we show evidence that zonation may be caused by burial because plant distribution was correlated with sand deposition and species were eliminated when burial exceeded their limits of tolerance, thus creating zones of different plant species. We conclude that in temperate regions (i) burial by sand may be among the most important factors in zonation, while salt spray may play a secondary role; and (ii) single environmental factors cannot be the determinants of a phenomenon as complex as species zonation.