We studied the effects of two fertilization treatments (N and NPKCa) on wood nitrogen (N) isotope composition (delta(15)N), water-use efficiency (WUE) estimated by carbon isotope composition (delta(13)C) analyses, and ring width of trees in 80-year-old beech (Fagus sylvatica L.) stands in the forest of Fougères, western France. Four replicates were fertilized in two successive years (1973 and 1974), 20 years before core sampling. Unfertilized control trees displayed a decreasing delta(15)N trend with time. The N and NPKCa treatments both increased delta(15)N compared with the control treatment. Wood extraction by organic solvents enhanced the delta(15)N signal. Thus, N addition to the beech ecosystem, even in moderate amounts, could be traced back in tree-ring delta(15)N, suggesting that wood N isotope analysis is a promising tool for studying the long-term effects of N deposition on forests. Although WUE decreased for about 6 years after N fertilization, WUE in NPKCa-treated trees did not differ significantly from that in control trees. Results were similar whether based on cellulose or total wood delta(13)C analysis, suggesting that extraction of cellulose is not necessary when studying fertilization impacts on WUE. The NPKCa treatment had a large impact on radial growth, causing a significant long-lasting increase of 29% compared with the control treatment. Nitrogen alone did not change radial growth significantly.

Large wild ungulates are a major biotic factor shaping plant communities. They influence species abundance and occurrence directly by herbivory and plant dispersal, or indirectly by modifying plant-plant interactions and through soil disturbance. In forest ecosystems, researchers' attention has been mainly focused on deer overabundance. Far less is known about the effects on understory plant dynamics and diversity of wild ungulates where their abundance is maintained at lower levels to mitigate impacts on tree regeneration. We used vegetation data collected over 10 years on 82 pairs of exclosure (excluding ungulates) and control plots located in a nation-wide forest monitoring network (Renecofor). We report the effects of ungulate exclusion on (i) plant species richness and ecological characteristics, (ii) and cover percentage of herbaceous and shrub layers. We also analyzed the response of these variables along gradients of ungulate abundance, based on hunting statistics, for wild boar (Sus scrofa), red deer (Cervus elaphus) and roe deer (Capreolus capreolus). Outside the exclosures, forest ungulates maintained higher species richness in the herbaceous layer (+15%), while the shrub layer was 17% less rich, and the plant communities became more light-demanding. Inside the exclosures, shrub cover increased, often to the benefit of bramble (Rubus fruticosus agg.). Ungulates tend to favour ruderal, hemerobic, epizoochorous and non-forest species. Among plots, the magnitude of vegetation changes was proportional to deer abundance. We conclude that ungulates, through the control of the shrub layer, indirectly increase herbaceous plant species richness by increasing light reaching the ground. However, this increase is detrimental to the peculiarity of forest plant communities and contributes to a landscape-level biotic homogenization. Even at population density levels considered to be harmless for overall plant species richness, ungulates remain a conservation issue for plant community composition.

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