Landscape metrics as functional traits in plants: perspectives from a glacier foreland.

Spatial patterns of vegetation arise from an interplay of functional traits, environmental characteristics and chance. The retreat of glaciers offers exposed substrates which are colonised by plants forming distinct patchy patterns. The aim of this study was to unravel whether patch-level landscape metrics of plants can be treated as functional traits. We sampled 46 plots, each 1 m × 1 m, distributed along a restricted range of terrain age and topsoil texture on the foreland of the Nardis glacier, located in the South-Eastern Alps, Italy. Nine quantitative functional traits were selected for 16 of the plant species present, and seven landscape metrics were measured to describe the spatial arrangement of the plant species' patches on the study plots, at a resolution of 1 cm × 1 cm. We studied the relationships among plant communities, landscape metrics, terrain age and topsoil texture. RLQ-analysis was used to examine trait-spatial configuration relationships. To assess the effect of terrain age and topsoil texture variation on trait performance, we applied a partial-RLQ analysis approach. Finally, we used the fourth-corner statistic to quantify and test relationships between traits, landscape metrics and RLQ axes. Floristically-defined relevé clusters differed significantly with regard to several landscape metrics. Diversity in patch types and size increased and patch size decreased with increasing canopy height, leaf size and weight. Moreover, more compact patch shapes were correlated with an increased capacity for the conservation of nutrients in leaves. Neither plant species composition nor any of the landscape metrics were found to differ amongst the three classes of terrain age or topsoil texture. We conclude that patch-level landscape metrics of plants can be treated as species-specific functional traits. We recommend that existing databases of functional traits should incorporate these type of data.