While climatic research about treeline has a long history, the climatic conditions corresponding to the upper limit of closed alpine grasslands remain poorly understood. Here, we propose a climatic definition for this limit, the ‘grassline’, in analogy to the treeline, which is based on the growing season length and the soil temperature. Eighty-seven mountain summits across ten European mountain ranges, covering three biomes (boreal, temperate, Mediterranean), were inventoried as part of the GLORIA project. Vascular plant cover was estimated visually in 326 plots of 1 × 1 m. Soil temperatures were measured in situ for 2–7 years, from which the length of the growing season and mean temperature were derived. The climatic conditions corresponding to 40% plant cover were defined as the thresholds for alpine grassland. Closed vegetation was present in locations with a mean growing season soil temperature warmer than 4.9 °C, or a minimal growing season length of 85 days, with the growing season defined as encompassing days with daily mean ≥ 1 °C. Hence, the upper limit of closed grasslands was associated with a mean soil temperature close to that previously observed at the treeline, and in accordance with physiological thresholds to growth in vascular plants. In contrast to trees, whose canopy temperature is coupled with air temperature, small-stature alpine plants benefit from the soil warmed by solar radiation and consequently, they can grow at higher elevations. Since substrate stability is necessary for grasslands to occur at their climatic limit, the grassline rarely appears as a distinct linear feature.
Variation in plant sex ratios is often attributable to sex-specific mortality in heterogeneous environments that differentially limit male and female plant reproduction. Yet sexual dimorphism and plastic responses to environmental heterogeneity are common and may co-vary with variation in sex ratios. Here, we show that the sex ratio and the degree of sexual dimorphism for a number of plant traits varied along climatic and elevation gradients in three wind-pollinated dioecious species, Rumex lunaria, Urtica dioica and Salix helvetica. Some of the observed sex-specific responses to climatic variation are consistent with greater sensitivity of females to water scarcity, but most responses rather point to the greater sensitivity of males to ecological stress, consistent with larger male reproductive effort, as has been commonly reported for wind-pollinated plants.In contrast, we found no evidence for variation in either sex ratios or sexual dimorphism expected under sexual selection. Interestingly, sex ratios and sexual dimorphism varied both along distinct and the same ecological axes of variation, suggesting that the evolution of sexual dimorphism in the measured traits was not sufficient to prevent sex-specific mortality.
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