Hierarchical organization of a Sardinian sand dune plant community

Coastal sand dunes have attracted the attention of plant ecologists for over a century, but they have largely relied on correlations to explain dune plant community organization. We experimentally examined longstanding hypotheses that sand binding, interspecific interactions, abiotic factors and seedling recruitment are drivers of sand dune plant community structure in Sardinia, Italy. Removing foundation species from the fore, middle and back dune habitats over 3 years led to erosion and habitat loss on the fore dune and limited plant recovery that increased with dune elevation. Reciprocal species removals in all zones suggested that interspecific competition is common, but that dominance is transient, particularly due to sand burial disturbance in the middle dune. A fully factorial 2-year physical factor manipulation of water, nutrient availability and substrate stability revealed no significant proximate response to these abiotic factors in any dune zone. In the fore and middle dune, plant seeds are trapped under adult plants during seed germination, and seedling survivorship and growth generally increase with dune height in spite of increased herbivory in the back dune. Sand and seed erosion lead to limited seed recruitment on the fore dune while high summer temperatures and allelopathy lead to competitive dominance of woody plants in the back dune. Our results suggest that Sardinian sand dune plant communities are hierarchically organized, structured by sand binding foundation species on the fore dune, sand burial in the middle dune and increasingly successful seedling recruitment, growth and competitive dominance in the back dune.