Genotypic and environmental variation in specific leaf area in a widespread Alpine plant after transplantation to different altitudes.

Specific leaf area (SLA) is an important plant functional trait as it is an indicator of ecophysiological characteristics like relative growth rate, stress tolerance and leaf longevity. Substantial intraspecific variation in SLA is common and usually correlates with environmental conditions. For instance, SLA decreases with increasing altitude, which is understood as adjustment to temperature. It is generally assumed that intraspecific variation is mostly the result of environmentally induced phenotypic plasticity, but genetic effects may also be present, due to local adaptation or genetic drift. In this study, genotypic and environmental effects on SLA were experimentally separated for the widespread Alpine bell flower Campanula thyrsoides by transplanting plants to three common gardens at contrasting altitudes (600, 1,235 and 1,850 m a.s.l.). Seeds were sampled from 18 populations in four phylogeographic regions within the European Alps. A strong plastic response was observed: SLA decreased with increasing altitude of the common gardens (22.0% of variation). The phylogeographic regions were differentiated in SLA in the common gardens (10.1% of variation), indicating that SLA is at least partly genetically determined. Plants from the six easternmost populations experienced a submediterranean climate and showed decreased SLA values in the three common gardens compared to populations to the west, which may be explained as adaptation to drought. Within these submediterranean populations, SLA decreased with altitude of origin in two out of three common gardens. Concluding, SLA shows strong phenotypic plasticity as well as substantial genetic effects, the latter probably being the result of adaptation to local conditions rather than genetic drift.