Although not widely used, ecosystem engineers represent a promising and sustainable tool in nature-based ecosystem management and restoration. In grassland ecosystems, a few invertebrates that engineer soils have been identified as key species regulating soil nutrients and plant communities' diversity and dynamics. Here, we assessed the role of the harvester ant Messor barbarus, an ecological engineer, in a Mediterranean dry grassland under restoration by characterising its nest environment, particularly the soil and vegetation. We found profound differences in soil physical and chemical variables and plant community structure between nests and antfree patches in the restored grassland. Messor barbarus has improved soil fertility, driven the seed bank towards the reference grassland and significantly increased plant biomass, species richness and micro-local-heterogeneity. As biological filters, M. barbarus has driven plant communities towards a new trajectory in the restored site. Ant patches are characterised by mesotrophic species, whereas ant-free patches are dominated by species characteristic of compacted soils. They have accelerated the ecological recovery of Mediterranean dry grassland plants by directly and indirectly facilitating their re-establishment. These results illustrate the potential key role of ants as ecological engineers for the conservation and restoration of Mediterranean grasslands.

Within a local assemblage, ecosystem engineers can have major impacts on population dynamics, community composition and ecosystem functions by transforming or creating new habitats. They act as an ecological filter altering community composition through a set of environmental variables. The impact of ants on their environment has been widely studied, but their multi-component effects (both trophic and non-trophic) have been rarely addressed. We investigated the roles of Messor barbarus , one of the commonest harvester ant species in south-western European Mediterranean grasslands. We analysed soil physico-chemical parameters, above-ground vegetation (e.g. species richness, plant community, micro-local heterogeneity, plant biomass) and above- and below-ground fauna (macrofauna, Collembola, Acari and nematodes). A clear and strong local impact of M. barbarus on soil, vegetation and fauna compartments emerges. The environmental filter is altered by modifications to soil physico-chemical properties, and the biotic filter by changes to plant communities and altered above- and below-ground fauna abundance, occurrence and community structure. The engineering activity of M. barbarus affects not only these separate ecosystem components but also the trophic and non-trophic relationships between them. By altering ecological filters at a local scale, M. barbarus creates habitat heterogeneity that may in turn increase ecological niches in these highly diverse ecosystems.