Effects of increased temperature, drought and nitrogen supply on two upland perennials of contrasting functional type: Calluna vulgaris and Pteridium aquilinum

Mature heather (Calluna vulgaris) and bracken (Pteridium aquilinum) turfs, transplanted from the field, were subjected to factorially combined experimental treatments for three consecutive years. Summer drought had the greatest effect, decreasing photosynthesis, growth and reproductive output in both species, and opening the bracken canopy. The timing of the drought relative to plant development was critical to which species was worst affected; bracken was worst affected by an early drought, heather by a later drought. Both species showed physiological damage during drought but, as predicted on the basis of their functional types, heather showed greater acclimation of water-use efficiency to drought stress. Contrary to expectations based on functional types, heather responded more rapidly than bracken to increased nitrogen supply (50 kg N ha(-1) yr(-1)). Added nitrogen caused both species to start above-ground growth earlier in the spring. For bracken this stimulation was short-lived; added nitrogen might be preferentially allocated to the rhizome and the longer-term consequences of this are unknown. For heather, nitrogen promoted growth and flowering throughout the season. There was no positive effect on the photosynthetic physiology of either species; changes in resource partitioning, and thus photosynthate production at the canopy level, are the most likely mechanism for the increase in heather shoot growth. Warmer temperatures increased heather shoot growth from early spring onwards but did not advance bracken crozier emergence, although frond height and the proportion of fertile fronds were subsequently increased. No significant effects of warming on the photosynthetic physiology of either species were found. Predictions of responses of heather and bracken to environmental change are complicated by the strong interactive effects of unpredictable climatic events such as drought and extreme winter temperatures. When drought was imposed, damage to heather was much greater in plants receiving increased nitrogen supply. Stimulation of growth by nitrogen resulted in a water demand that was unsustainable in drought conditions, leading to wilting, reduced shoot growth and some acclimation of water-use efficiency. Additionally, a very cold winter spell proved most damaging to heather that had been droughted in the previous summer. For bracken, winter damage occurred in plants that had been warmed, with significantly fewer fronds emerging in the next spring and thus canopy photosynthetic potential being reduced. We predict that positively managed heather has the potential to limit the bracken problem in conditions of environmental change, provided that high levels of nitrogen deposition do not coincide with increased drought frequency.