Acacia karroo invasion of grassland: environmental and biotic effects influencing seedling emergence and establishment

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Abstract
Acacia karroo Hayne is the most important woody invader of grassland in South Africa, and can greatly reduce the productivity of grassland. A field experiment was conducted to test the hypotheses that emergence, growth and the 1st year's survival of Acacia karroo would be enhanced by (1) defoliation of the grass sward, (2) increased irradiance, (3) increased moisture availability and (4) its germination within cattle dung pats. The study was conducted on one site above and one below the natural altitudinal treeline of this species in grassland of the eastern Cape, South Africa. Not one seedling emerged from dung pats. Neither location nor the other treatments affected the density of emerging seedlings, although only 40.4 seedlings m−2 emerged of the 200 seeds m−2 planted. Shading dramatically increased the density of surviving seedlings. In the open, only 3 and 1.5 seedlings m−2 remained respectively at the end of the growing season or the beginning of the next, compared to 23.3 and 19.5 seedlings m−2 under shading for these respective times. This was attributed to the effect of shade on moisture availability in a season which received only 54% of average rainfall. Seedling survival until the end of the growing season was enhanced (30%) by shade at both sites, but also by supplemental water (24%) and defoliation of the sward (7%) at the site above the treeline. Across sites and treatments, seedling survival was related to moisture availability, with no or poor survival for < 500 mm rainfall, indicating this species can only establish in certain rainfall years. Seedling survival over winter was not influenced by treatment, but was greater for larger seedlings. Treatments affected seedling size, in particular seedlings growing under shade and within a dense grass canopy were etiolated. A. karroo seedlings are capable of establishing and surviving within a dense grass sward for at least a year, tolerant of low irradiance and of interference, which, because most seeds do not persist for much longer than a year, suggests this species forms predominantly a seedling bank. This has implications for the invasion of grassland by woody species.