The resurrection flowering plant Ramonda nathaliae on serpentine soil – coping with extreme mineral element stress

Ramonda nathaliae (Gesneriaceae) is a rare desiccation tolerant flowering plant species of the Northern Hemisphere. This, mainly calcicole, preglacial relict species is endemic in the Balkan Peninsula, where it has survived in its refugial habitats of gorges and mountain slopes. At present, only two localities within its narrow range are known where it thrives in hostile serpentine habitats, and the adverse serpentine environment is bound to present further challenge to the adaptive capacity of R. nathaliae. In general, the occurrence of a resurrection flowering plant on serpentine soil is exceptional and the soil-plant relation of R. nathaliae in those circumstances is described here for the first time. The aim of this study was (i) to analyze mineral elements composition in soil from both serpentine and limestone habitats of the species and to compare the way peculiarities of the substrates are reflected in roots and leaves of plants from the respective soils; (ii) to evaluate the effect of heavy metal overload on the habit of serpentine R. nathaliae individuals. Serpentine soil, characterized by high levels of phytotoxic heavy metals (Ni, Cr, Co, Mn), hosts plants showing elevated metal contents in their organs. Ramonda plants from serpentine populations are able to maintain balance of Ca to Mg favourable to Ca (2.0 in roots, 2.7 in leaves) despite a strongly unfavourable Ca/Mg ratio in the soil (0.09). The greatest difference in concentrations was observed for the Ni content in plant tissues: serpentine plants had 57 and 20 times more Ni in their roots and leaves than the plants from limestone. Aluminium was present in similar concentrations in both soils, and was highly accumulated in plant tissues of the plants from both substrates. Metal-specific metabolic activity is demonstrated in bioaccumulation indices of several essential minerals (Ca, Mg, Cu, Zn). A significantly higher metal content found in roots in relation to leaves might indicate the plant's ability to immobilize the metals within the root tissues. Mycorrhizal fungi colonize plant roots from both substrates and apparently are important in improving the supply of nutrients, but they could also take part in toxic metal immobilization. The price of adaptation to the hostile environment is evident in the habit of R. nathaliae plants growing on serpentine: reduced size of rosettes and leaves, as well as chlorotic and necrotic leaf tips and margins.