The influence of cattle activity on tree regeneration in wood-pastures

Wood-pastures are semi-natural, highly biodiverse systems maintained by traditional extensive agriculture. They are used for grazing and timber. The shifting mosaics of grassland, shrub thickets and woodland patches in these systems are driven by large herbivores. Current changes in agricultural practices are leading to either intensification or abandonment and they result in a segregation of grassland and forest. This presents a complex conservation problem for these endangered ecosystems. Large herbivores have been identified as an important factor preventing the establishment of trees and the regeneration of woodland. However, successful management is severely hindered by the lack of ecological knowledge about the effects of large herbivores on woody species. This research focussed on the quantification of the impact of cattle activity on the early stages of tree establishment for four tree species (Abies alba, Picea abies, Acer pseudoplatanus and Fagus sylvatica) common in the wood-pastures of the Swiss Jura Mountains. Cattle can have both direct and indirect effects on tree establishment, and the interaction between cattle activity and both competition and facilitation by surrounding vegetation was studied. Experimental field studies (involving seed sowing and sapling planting) were carried out. Both cattle activity and competition from neighbours reduced first-year tree seedling recruitment in wood-pastures. Seedling establishment decreased with increasing cattle grazing intensity. The effects of gaps in the vegetation on seedling emergence were complex, as a result of biotic (i.e. competition, facilitation and predation from rodents, invertebrates and large herbivores) and abiotic (e.g. moisture, temperature and light) interactions, leading to different results in different years. Although overall the chances for tree seeds to make it to seedlings were low, seedling establishment was relatively high for Picea, intermediate for Abies and Acer and low for Fagus. The probability of a sapling of being browsed increased with sapling size and decreased with the height of surrounding vegetation and in the presence of nurse shrubs. It was similar for the four species and under low and high grazing intensities. Saplings had species- and size-specific responses in survival and growth to cattle browsing. Under both low and high grazing intensity, small coniferous saplings (36 %) had a lower survival rate than small deciduous saplings (53 %), which we hypothesize is due to the larger biomass losses stemming from a more vulnerable plant architecture. Under low grazing intensities, large Picea (80 %) and Fagus (67 %) saplings had a higher survival than large Acer (50 %) and Abies (33 %) saplings. This was likely due to differences in sapling tolerances to loss of biomass. Simulated browsing damage resulted in relatively smaller growth losses when small saplings were growing slowly in the presence of neighbours. Shade increased this effect. At least in t