Interactive effects of nitrogen deposition and climate change on a globally rare forest geophyte.

Nitrogen (N) deposition and climate change are both known to threaten global biodiversity. However, we still have a limited understanding of how interactions between these global change drivers affect individuals and populations of specialist species, such as geophytes, within their natural habitat. We explored possible interactive effects of N, drought, and warming on population vitality (mean leaf length, leaf density, flowering probability) and morpho-physiological traits (e.g., leaf and bulb size, N allocation to leaves and bulbs) of the globally rare forest geophyte Gagea spathacea (Liliaceae) in deciduous forests of northern Germany by applying experimental N addition across a climate gradient over a 5-year period. Mean leaf growth and leaf density were not affected by N addition but were enhanced by warmer and drier conditions in the months before leaf emergence. N addition increased N allocation of individual plants towards their subterranean bulbs. Importantly, effects of N addition on morpho-physiological traits depended on warming and drought, with N-fertilized plants showing increased leaf length and decreased specific leaf and bulb N concentration after drier autumns and warmer winters. This indicates that N deposition may partially compensate for increased N demands during warming-induced growth, although this growth-promoting interaction effect is not (yet) reflected in population vitality. Our results highlight the importance of considering multiple global environmental change drivers and a whole plant perspective (above- and belowground traits) to predict long-term growth responses of (endangered) forest spring geophytes and to develop adapted long-term protection strategies.