Current conceptual models predict that an increase in stress shifts interactions between plants from competitive to facilitative; hence, facilitation is expected to gain in ecological importance with increasing stress. Little is known about how facilitative interactions between plants change with increasing biotic stress, such as that incurred by consumer pressure or herbivory (i.e. disturbance sensu Grime). In grazed ecosystems, the presence of unpalatable plants is reported to protect tree saplings against cattle grazing and enhance tree establishment. In accordance with current conceptual facilitation-stress models, we hypothesised a positive relationship between facilitation and grazing pressure. We tested this hypothesis in a Weld experiment in which tree saplings of four diVerent species (deciduous Fagus sylvatica, Acer pseudoplatanus and coniferous Abies alba, Picea abies) were planted either inside or outside of the canopy of the spiny nurse shrub Rosa rubiginosa in enclosures diVering in grazing pressure (low and high) and in exclosures. During one grazing season we followed the survival of the diVerent tree saplings and the level of browsing on these; we also estimated browsing damage to the nurse shrubs. Shrub damage was highest at the higher grazing pressure. Correspondingly, browsing increased and survival decreased in saplings located inside the canopy of the shrubs at the high grazing pressure compared to the low grazing pressure. Saplings of both deciduous species showed a higher survival than the evergreens, while sapling browsing did not diVer between species. The relative facilitation of sapling browsing and sapling survival -i.e. the diVerence between saplings inside and outside the shrub canopy -decreased at high grazing pressure as the facilitative species became less protective. Interestingly, these Wndings do not agree with current conceptual facilitationstress models predicting increasing facilitation with abiotic stress. We used our results to design a conceptual model of facilitation along a biotic environmental gradient. Empirical studies are needed to test the applicability of this model. In conclusion, we suggest that current conceptual facilitation models should at least consider the possibility of decreasing facilitation at high levels of stress.

Summary1. According to the insurance hypothesis, more diverse plant communities are more likely to be resistant to drought. Whilst many experiments have been carried out to determine the effects of plant diversity on plant community insurance, the results are still contradictory. 2. Here, we conducted a drought experiment where we tested whether the presence of subordinate species increases plant community insurance. In Swiss Jura grassland, we combined a removal experiment of subordinate species with a summer drought event using rainout shelters. 3. Plant community composition was determined after the drought and based on biomass measurements; we estimated resistance, recovery and resilience of the plant community for each combination of treatments. Moreover, to assess drought impacts on water-use efficiency (WUE), we analysed carbon isotope ratios (d 13 C values) in plant leaves of two dominants and two subordinates collected at the end of the drought period. 4. We showed that subordinate species are more resistant to drought and increased community resistance by enhancing their above-ground biomass production during the imposed drought. These patterns were associated with decreased competitiveness of dominant species whose biomass decreased during drought. Significant increase in d 13 C values in plant tissue under drought indicated a better WUE for the measured species. Interestingly, the WUE was significantly higher in plots where subordinates were removed. Recovery and resilience were not affected by the summer drought, but the absence of subordinates reduced overall above-ground biomass in both watered and drought plots. 5. Synthesis. We demonstrated that, independent of plant diversity, the presence of drought-resistant subordinate species increases plant community insurance against drought and, hence, is important for the functioning of grassland ecosystems.

Summary1. Light-demanding trees and thorny shrubs in temperate plant communities may reflect adaptations to now-extinct large grazers, such as aurochs and tarpans, rendering these adaptations ecological anachronisms. 2. We explored the ecological functions of plant traits of Quercus robur and Prunus spinosa in areas grazed by cattle and horses, the domesticated descendants of aurochs and tarpans. Specifically, we tested the hypothesis that grazing induces a shifting mosaic of grassland, shrub thickets and woodlands through the key process of associational resistance: the protection of palatable young trees by thorny shrubs.3. An exclosure experiment with transplanted Q. robur seedlings revealed that Q. robur grew best in grassland exclosures and on the edge of thorny shrub thickets, which may be viewed as an optimal balance between sufficient protection from large herbivores and sufficient light availability. 4. A cross-site comparison of four floodplain woodlands in north-western Europe showed that Q. robur can regenerate in the presence of large herbivores through spatial association with P. spinosa . However, we found that expansion of P. spinosa shrubs and Q. robur coincided with periods of low rabbit abundance and not with livestock density. From this, it appears that the process of associational resistance does not work with rabbits. 5. Synthesis and applications . With extensive grazing by large (domesticated) grazers in temperate floodplains, a shifting mosaic of grassland, shrubs and trees may develop that has high conservation value. Palatable, light-demanding Q. robur seedlings can successfully regenerate in spiny P. spinosa shrubs through associational resistance. This process does not offer protection from abundant small herbivores, such as rabbits, that can inhibit the recruitment of shrubs and trees in this mosaic vegetation. In floodplain meadows frequent flooding may be an efficient way to reduce rabbit populations, with dry conditions in summer and wet in winter. When floodplain meadows are combined with adjacent higher grounds, large herbivores can escape the floods through migration.