Regionally Varying Habitat Relationships in Lichens: The Concept and Evidence with an Emphasis on North-Temperate Ecosystems.

Habitat ecology of lichens (lichen-forming fungi) involves diverse adaptations to stressful environments where lichens use specific habitat conditions. Field observations confirm that such habitat 'preferences' can vary significantly across species' distribution ranges, sometimes revealing abrupt changes over short distances. We critically review and generalize such empirical evidence as broad ecological patterns, link these with the likely physiological mechanisms and evolutionary processes involved, and outline the implications for lichen conservation. Non-replicated correlative studies remain only suggestive because the data are frequently compromised by sampling bias and pervasive random errors; further noise is related to unrecognized cryptic species. Replicated evidence exists for three macroecological patterns: (a) regional limiting factors excluding a species from a part of its microhabitat range in suboptimal areas; (b) microhabitat shifts to buffer regionally adverse macroclimates; (c) substrate suitability changed by the chemical environment, notably air pollution. All these appear to be primarily buffering physiological challenges of the adverse conditions at the macrohabitat scale or, in favorable environments, coping with competition or predation. The roles of plasticity, adaptation, dispersal, and population-level stochasticity remain to be studied. Although lichens can inhabit various novel microhabitats, there is no evidence for a related adaptive change. A precautionary approach to lichen conservation is to maintain long-term structural heterogeneity in lichen habitats, and consider lichen ecotypes as potential evolutionarily significant units and a bet-hedging strategy for addressing the climate change-related challenges to biodiversity.