Several explanations have been given for the decline in species richness with altitude. However, separating the influences of altitude, area, and isolation is difficult because of the conical shape of mountains. We used species lists of aquatic plants from >300 lakes in a small geographical area to investigate the influence of altitude on species richness. Altitude and/or surface area were better predictors of species richness than any measure of water chemistry. The surface area and depth of individual lakes were not related to altitude, neither was the degree of isolation from other waterbodies. Although species range size increased with altitude, range sizes of all but the rarer species (in the data set) encompassed the lowest altitudes, indicating species loss rather than turnover and no influence of the Rapoport rescue effect. Nevertheless we found a decline in species richness with altitude, additive to the effect of area. Species were ascribed to attribute groups according to an a priori classification based on morphological and life-history traits. The number of attribute groups and number of species within each group increased with area, suggesting both increased diversity and coexistence within niches. With altitude, the number of attribute groups declined, but the number of species per group increased, consistent with a loss of richness and reduced competition. The species remaining at high altitudes were characterised by stress tolerant traits, associated with sites of low productivity.
Our results suggest an absolute effect of altitude on species richness, irrespective of other influences and consistent with a decline in productivity.