Terricolous lichens are relevant associates of biological soil crusts in arid and semiarid environments. Dunes are ecosystems of high conservation interest, because of their unique, vulnerable and threatened features. The function of lichens is affected by the changing seasons and different microhabitat conditions. At the same time, inland dunes are less investigated areas from the terricolous lichens point of view. We explored the effect of seasonal variation and different micro-environmental conditions (aspect) on the metabolic activity of five terricolous lichen species, representing various growth forms, in temperate semiarid grasslands. Populations of Cladonia foliacea, C. furcata, C. pyxidata group, Diploschistes muscorum and Thalloidima physaroides were investigated. Thalli sampled from the south-west and north-east facing microhabitats were studied by chlorophyll fluorescence analysis for 2 years. The present study aims to understand how changing climate (during the year) and aspect affect photosynthetic activity and photoprotection. Microclimatic data were also continuously recorded to reveal the background of the difference between microhabitat types. As a result, the air temperature, photosynthetically active radiation, soil temperature and vapour pressure deficit were significantly higher on south-west than on north-east facing microsites, where relative humidity and water content of soil proved to be considerably higher. Higher photosynthetic activity, as well as a higher level of photoprotection, was detected in lichens from north-east-oriented microsites compared with south-west populations. In addition, the difference between sun and shade populations varied seasonally. Since a species-specific response to both aspect and season was detected, we propose to investigate more than one species of different growth forms, to reveal the response of lichens to the changing environment in space and time.

In lichens, each symbiotic partner cooperates for the survival of the symbiotic association. The protection of the susceptible photosynthetic apparatus is essential for both participants. The mycobiont and photobiont contribute to the protection against the damaging effect of excess light by various mechanisms. The present study investigated the effect of seasonality and microhabitat exposure on photoprotection and photoacclimation in the photo- and the mycobiont of six lichen species with different thallus morphology in inland dune system in the Kiskunság region (Hungary) with shaded, more humid and exposed, drier dune sides. High-Performance Liquid Chromatography, spectrophotometry, chlorophyll a fluorescence kinetic technique were used, and micrometeorological data were collected. The four years data series revealed that the north-east-facing side was characterized by higher relative humidity and lower light intensities compared to the south-west-facing drier and more exposed sides. The south-west facing side was exposed to direct illumination 3–4 hours longer in winter and 1–2 hours shorter in summer than the north-east facing side of the dune, influencing the metabolism of sun and shade populations of various species. Because rapid desiccation caused short active periods of lichens during bright and drier seasons and on exposed microhabitats, the rapid, non-regulated non-photochemical quenching mechanisms in the photobiont had a significant role in protecting the photosynthetic system in the hydrated state. In dehydrated conditions, thalli were mainly defended by the solar screening metabolites produced by the mycobiont and curling during desiccation (also caused by the mycobiont). Furthermore, the efficacy of light use (higher chlorophyll and carotenoid concentration) increased because of short hydrated periods. Still, a lower level of received irradiation was appropriate for photosynthesis in dry seasons and on sun exposed habitats. In humid seasons and microhabitats, more extended active periods lead to increased photosynthesis and production of solar radiation protectant fungal metabolites, allowing a lower level of photoprotection in the form of regulated non-photochemical quenching by the photobiont. Interspecific differences were more pronounced than the intraspecific ones among seasons and microhabitat types.

The terricolous species Cladonia foliacea (Cladoniaceae, lichenised Ascomycota) widely distributed in open, dry lowland steppe and rocky mountain grassland vegetation in Europe was chosen as a potential test organism for ecological experiments, since their thalli are producing cortical solar radiation-protective and UV screening pigment dibenzofuran usnic acid and medullary secondary substance depsidone fumarprotocetraric acid. Significant seasonal differences were found in the amounts of lichen secondary metabolites analysed by HPTLC and HPLC-PDA between summer and winter collected thalli in sandy grassland area in Hungary. The concentrations of usnic acid varied between 7.34 and 15.52 mg/g in summer collected samples and 13.90 and 21.61 mg/g in winter collected ones. A comparable amount (11.61±0.29 mg/g) was measured in pulverised samples. The concentrations of fumarprotocetraric acid varied between 0.60 and 3.01 mg/g in summer collected samples and 2.26 and 5.81 mg/g in winter collected thalli. A comparable amount (2.45±0.21 mg/g) was found in pulverised samples. The range of concentration values is comparable with data known from lichens. A higher amount of usnic acid is produced in winter probably to ensure sufficient protection also for summer. The fumarprotocetraric acid content of the medulla might contribute to the solar irradiation reflecting role of the pale lower surface lobes turning upwards in dry condition.

Terricolous lichens are abundant in semi-arid areas, where they are exposed to high irradiation. Photoprotection is essential for the algae as the photobiont provides the primer carbon source for both symbionts. The UV-protectant lichen metabolites and different quenching procedures of the alga ensure adequate photoprotection. Since the long-term effect of diminishing UV-protectant lichen metabolites is unknown, a major part of lichen secondary metabolites was removed from Cladonia foliacea thalli by acetone rinsing, and the lichens were then maintained under field conditions to investigate the effect on both symbionts for 3 years. Our aim was to determine if the decreased level of UV-protectant metabolites caused an elevated photoprotection in the algae and to reveal the dynamics of production of the metabolites. Photosynthetic activity and light protection were checked by chlorophyll a fluorescence kinetics measurements every 6 months. The concentrations of fumarprotocetraric and usnic acids were monitored by chromatographic methods. Our results proved that seasonality had a more pronounced effect than that of acetone treatment on the function of lichens over a long-term scale. Even after 3 years, the acetone-treated thalli contained half as much usnic acid as the control thalli, and the level of photoprotection remained unchanged in the algae. However, the amount of available humidity was a more critical limiting environmental factor than the amount of incoming irradiation affecting usnic acid production. The lichenicolous fungus Didymocyrtis cladoniicola became relatively more abundant in the acetone-treated samples than in the control samples, indicating a slight change caused by the treatment.

Distribution data originating from earlier herbarium collections and recent biodiversity records form the basis of distribution analyses in lichen species with different ecological requirements, where the records allowed comparisons or showed clear trends. As the occurrences of lichens are strongly correlated to background environmental conditions (e.g., air pollution, global warming), confirmed by Wirth’s ecological indicator values, the analysis of distribution types has a great value for bioindication and the establishment of current and future climatic and pollution situations. Five distribution types were introduced—presented by characteristic examples (13)—according to lichen distribution maps prepared in different periods of time (representing changing environmental conditions): (1) species of decreasing occurrences by time (e.g., Lobaria pulmonaria, Menegazzia terebrata, suboceanic, acidic pollution sensitive species), (2) species with no or few former records but with increasing occurrences in recent decades (e.g., Flavoparmelia soredians, Hyperphyscia adglutinata, Solenopsora candicans, sub-Mediterranean species), (3) species with increasing and then (from c. 2000) decreasing occurrences (e.g., Scoliciosporum chlorococcum, Straminella conizaeoides, acidofrequent species), (4) species with widely increasing occurrences in recent decades (e.g., Physcia aipolioides, Piccolia ochrophora, Xanthoria parietina, nitrofrequent species), and (5) species with rapidly increasing occurrences (e.g., Absconditella lignicola, Coenogonium pineti, Evernia divaricata, rapidly spreading species). The proposed distribution types of lichen species may be applied to wider regions (the European or the global level).