Plant and vegetation functional responses to cumulative high nitrogen deposition in rear-edge heathlands

Elevated atmospheric nitrogen (N) deposition is a major driver of change, altering the structure/functioning of nutrient-poor Calluna vulgaris-heathlands over Europe. These effects amply proven for north-western/central heathlands may, however, vary across the ecosystem's distribution, especially at the range limits, as heathlands are highly vulnerable to land-use changes combined with present climate change. This is an often overlooked and greatly understudied aspect of the ecology of heathlands facing global change. We investigated the effects of five N-fertilisation treatments simulating a range of N deposition rates (0, 10, 20, and 50 kg N ha^-1 yr^-1 for 1 year; and 56 kg N ha^-1 yr^-1 for 9 years) on the Calluna-plants, the plant functional groups, species composition and richness of two life-cycle stages (building/young- and mature-phase) of Calluna-heathlands at their rear-edge limit. Our findings revealed a dose-related response of the shoot length and number of flowers of young and mature Calluna-plants to the addition of N, adhering to the findings from other heathland locations. However, cumulative high-N loading reduced the annual growth and flowering of young plants, showing early signs of N saturation. The different plant functional groups showed contrasting responses to the cumulative addition of N: annual/perennial forbs and annual graminoids increased with quite low values; perennial graminoids were rather abundant in young heathlands but only slightly augmented in mature ones; while bryophytes and lichens strongly declined at the two heathland life-cycle stages. Meanwhile there were no significant N-driven changes in plant species composition and richness. Our results demonstrated that Calluna-heathlands at their low-latitude distribution limit are moderately resistant to cumulative high-N loading. As north-western/central European heathlands under high-N inputs broadly experienced the loss of plant diversity and pronounced changes in plant species dominance, rear-edge locations may be of critical importance to unravel the mechanisms of heathland resilience to future global change.