Leaf size and leaf area index in Fagus sylvatica forests:: Competing effects of precipitation, temperature, and nitrogen availability

Plants across diverse biomes tend to produce smaller leaves and a reduced total leaf area when exposed to drought. For mature trees of a single species, however, the leaf area-water supply relationship is not well understood. We tested the paradigm of leaf area reduction upon drought by a transect study with 14 mature Fagus sylvatica forests along a steep precipitation gradient (970-520 mm y(-1)) by applying two independent methods of leaf size determination. Contrary to expectation, average leaf size in dry stands (520-550 mm y(-1)) was about 40% larger and SLA was higher than in moist stands (910-970 mm y(-1)). As a result of increased leaf sizes, leaf area index significantly increased from the high- to the low-precipitation stands. Multiple regression analyses suggested that average leaf size was primarily controlled by temperature, whereas the influence of soil moisture and soil C/N ratio was low. Summer rainfall of the preceding year was the most significant predictor of total leaf number. We assume that leaf expansion of beech was independent of water supply, because it takes place in May with ample soil water reserves along the entire transect. In contrast, bud formation, which determines total leaf number, occurs in mid-summer, when droughts are severest. We conclude that leaf expansion and stand leaf area of beech along this precipitation gradient are not a simple function of water availability, but are controlled by several abiotic factors including spring temperature and possibly also nitrogen supply, which both tend to increase toward drier sites, thus overlaying any negative effect of water shortage on leaf development.