Echoes of the past: Agricultural legacies shape the successional dynamics of protected semi-natural dry grasslands.

European semi-natural dry grasslands are among the most endangered terrestrial ecosystems, being recognised as habitats of community interest by the EU Habitats Directive. The occurrence and preservation of these habitats depend on a combination of anthropogenic and natural factors, although little is known regarding the role of past land-use changes. Here, we investigated the role of time since cultivation abandonment as a major driver of grassland successional dynamics in the Mediterranean agro-pastoral system of Alta Murgia, southern Italy. By integrating cartographic information on the past agricultural land-use with the main abiotic constraints (patch area, slope and aspect), we used generalised additive mixed models to test for the probability of occurrence of current grassland habitat types along time since cultivation abandonment (10 to 200 years). Our results disclosed the successional sequence of grassland plant communities since crop abandonment in the study area, highlighting that the distribution of semi-natural grassland communities largely depends on land use history besides current environmental patterns. Among the habitat types protected under the EU Habitats Directive, we highlighted that xero-thermic communities may represent an intermediate step of grassland succession after cultivation abandonment, while more mesic perennial communities indicate a late successional stage. These successional dynamics are further modulated by mesoclimatic conditions associated with slope and aspect, especially in case of long-standing pastures that were not historically affected by agricultural transformations. Our findings can contribute to a deeper understanding of dynamics relevant to spontaneous vegetation recovery in open environments, which is a prerequisite for setting up effective grassland conservation and restoration actions. Furthermore, our results underline the value of integrating historical maps and current information for the assessment of habitat conservation status.