Spatial variability of ultraviolet-absorbing compounds in an aquatic liverwort and their usefulness as biomarkers of current and past UV radiation: A case study in the Atlantic-Mediterranean transition

The spatial variability of ultraviolet-absorbing compounds (UVACs) in the freshwater liverwort Jungermannia exsertifolia subsp. cordifolia was studied in mid-latitudes (the Atlantic-Mediterranean transition) across a wide lati-altitudinal gradient, with the aim of testing the usefulness of UVACs as biomarkers of current ambient levels of UV radiation. We analysed 17 samples from streams located in the main mountain ranges of the Iberian Peninsula, differentiating methanol-soluble (SUVACs, mainly located in the vacuoles) and methanol-insoluble (IUVACs, bound to cell walls) compounds, since they represent different manners to cope with UV radiation. In both fractions, the bulk level of UVACs and the concentrations of several individual compounds were measured. In addition, we measured Fv/Fm, DNA damage and sclerophylly index (SI) as possible additional UV biomarkers. UVACs showed a high variability, probably due not only to the gradients of macroenvironmental factors (UV radiation, PAR, and water temperature), but also to microenvironmental factors inherent to the dynamic nature of mountain streams. Two soluble coumarins were positively correlated with UV levels and could be used for ambient UV biomonitoring in the spatial scale. In contrast to the variability in UVACs, the relatively homogeneous values of Fv/Fm and the lack of any DNA damage made these variables useless for ambient UV biomonitoring, but suggested a strong acclimation capacity of this liverwort to changing environmental conditions (in particular, to UV levels). Finally, UVACs of fresh samples of the liverwort were compared to those of herbarium samples collected in the same lati-altitudinal gradient. SUVACs were significantly higher in fresh samples, whereas IUVACs generally showed the contrary. Thus, IUVACs were more stable than SUVACs and hence more adequate for retrospective UV biomonitoring. In conclusion, UVAC compartmentation should be taken into account for bryophyte-based UV biomonitoring in future studies.