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Facilitation of Quercus ilex recruitment by shrubs in Mediterranean open woodlands

Abstract Question: Insufficient tree regeneration threatens the long-term persistence of biodiverse Mediterranean open oak woodlands. Could shrubs, scarce due to decades of management (clearing and ploughing), facilitate holm oak recruitment at both acorn and seedling stages? Location: Open oak woodlands in Central Spain. Methods: Plots with four acorns were planted: (1) under the canopy of the spiny shrub Genista hirsuta, (2) in a small cage, protecting against ungulates, (3) in a shaded cage, protecting against ungulates and sun, and (4) in open grassland. Sets of these four treatments were spatially grouped according to a randomised block design, with 16 blocks near (< 10 m) and 16 away from (> 20 m) parent trees to test for distance-related survival. Plots were regularly checked for seed removal. After emergence one seedling per plot (97 in total) was selected and its survival monitored. Results: Three months after sowing, 199 of 512 acorns were removed, predominantly by rodents. Acorn removal occurred at each treatment but was highest under shrubs. Eight months after sowing, seedling survival was highest under shrubs (50%), followed by shaded cages (16%), open grassland (4%) and cages (0%). Main mortality cause was drought (90%), killing most seedlings between June and July. No seedlings died from ungulate browsing. Conclusion: Shrubs demonstrated clear net facilitative effects for Quercus ilex recruitment, despite higher seed removal. Shading appears the crucial factor facilitating seedling survival. We therefore propose that lack of shrubs contributes largely to tree recruitment failure in Mediterranean open woodlands; management should aim at conserving shrubs.