Although changes in land-use, climate, and the spread of introduced tree species have increased the global importance of tree invasions into grasslands, our ability to predict any particular invasion is limited. To elucidate mechanisms driving tree invasions of grasslands, we studied in detail how seed dispersal and fine-scale environment control the expansion of an introduced Picea abies Karst. (Norway spruce) population into Western Carpathian grassland. We mapped invading trees and measured tree size, fecundity, seed rain, seedling density, plant community composition, and light and soil environment within a 200 · 60 m belt across the invasion front. Maximum likelihood estimates of dispersal kernels suggested peak seed deposition directly underneath tree crowns where germination was poor, but mean dispersal distances were sufficiently large to generate over- lapping seed shadows from multiple trees that saturated the invasion front with seeds further away from seed-dispersing trees. Partial Mantel tests indicated that germinant density was affected con- siderably less by seed rain than by moss cover (r = 0.54), overstory tree influence (r = 0.32), soil moisture (r = 0.21), grass cover (r = 0.15), and diffuse radiation (r = 0.13). However, these vari- ables were not independent but formed complex multivariate gradients within the invasion front. Moss cover and soil moisture were negatively correlated with overstory tree influence and the resulting gradient clearly affected germinant density (partial Mantel r = 0.45). In contrast, positively correlated light and grass cover defined a gradient related weakly to germinant density (partial Mantel r = 0.05) as it integrated opposing effects of these variables on germinants. Seedlings had similar environmental associations, but except for the lasting positive effects of moss these tended to weaken with seedling size. Although a few seedlings may establish and survive in the more adverse environment of the outer edges of the invasion front, a significant population expansion may require a gradual build-up of the critical density of invading trees to reduce grass cover and facilitate germination on moist mossy seedbeds within uncolonized areas. Thus, Picea abies appears more likely to spread within temperate grasslands by gradual expansion of its population frontier rather than by advanced groups.
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