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Cascading effects of changes in land use on the invasion of the walnut Juglans regia in forest ecosystems

Plant invasions are affected by many factors that must be favourable in order for invasions to occur. Factors can be grouped into three major categories: propagule pressure, biotic factors and abiotic characteristics; all may be moderated by human activity. However, studies examining all factors simultaneously are rare, and most are limited to a single factor. This hampers our understanding of the mechanisms driving invasions. In recent decades, an alien walnut (Juglans regia) has become invasive in Central Europe due to an increase in the populations of native dispersers, rooks (Corvus frugilegus) and political changes that have resulted in the abandonment of farmland. Here, we test whether increased propagule pressure resulting from the presence of seed‐bearing walnuts in abandoned fields interacts with native forest seed‐dispersing animals (biotic factors), landscape structure and management, and passive secondary dispersal by gravity (abiotic factors) to facilitate the further invasion of walnuts into forests. Seed‐dispersing animals were observed visiting walnuts growing in abandoned fields and in human settlements. Walnuts and seed dispersers were surveyed in 102 forest patches. Forest characteristics were examined, and field experiments examining passive dispersal were conducted. Walnut seeds were carried into forests by native birds, jays (Garrulus glandarius). Jays harvested seeds from both wild walnuts growing in abandoned fields and walnuts planted in human settlements. The density of walnuts in forest patches was correlated with jay density. Forest occupancy by walnuts and walnut densities were correlated with the cover of abandoned fields and human settlements containing seed‐bearing walnuts. Secondary seed dispersal also affected forest colonization. Walnut seeds hidden by native rooks in sloping, arable fields may be transported passively to forest edges. Synthesis. Our results show that this invasion is a multifaceted process in which human‐related alterations to propagule pressure and biotic and abiotic factors have led to the spread of alien walnuts from human settlements and abandoned fields into forest ecosystems. Thus, politically related land use changes can create an invasion debt that causes unexpected linkages among the invasive plant, native dispersers, land management and topography that together can cause cascading changes in ecosystems.