The study area was the Friuli Venezia-Giulia region (FVG, centroid coordinates = 46° 24′ 00″ N, 13° 04′ 50″ E, Fig. 1a). It covers an area of about 7900 km2 and is bounded by the Julian and Carnic Alps to the north, the Adriatic Sea to the south and the Karst of Trieste to the east. FVG exhibits a high geological and morphological complexity77, showing a prevalence of calcareous soils (i.e. limestone and dolomites) in the mountain region, alluvial calcareous deposits in the lowlands and the occurrence of both rock and sandy coasts77. FVG exhibits a continental climate in the inner valleys and temperate conditions in the outer area, with a mean annual air temperature of about 12 °C. Due to its ecological complexity, FVG has been recognised for its considerable plant diversity since early botanical surveys78. The area encompasses several protected areas including well-conserved wetlands, grasslands and ancient woods. Landscape variability determines a large complexity of land uses, mainly dedicated to the intensive agriculture in the plain. In these cultivated areas both annual (maize, barley, soybean) and perennial crops (vineyards, orchards, poplar) are frequent79. FVG was selected as study area because of the occurrence of clear altitudinal and climatic gradients, ranging from the coast to the Alps and the strong impact of human land uses. Moreover, an intensive and updated floristic survey was conducted during the last 40 years resulting in a solid database of presence and location of native or exotic species (see below).

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