Sphagnum limits : Physiology, morphology and climate

Sphagnum is the most important plant genus in terms of terrestrial carbon cycling. It and the habitats it creates store an equivalent of ~68% of the CO2 in the atmosphere. The genus has little dispersal limitation and the mire habitats are functionally similar at global scales. Sphagnum species are limited by water deficit at local and biogeographic scales, but this alone is not sufficient to explain local and global scale species patterns. As Sphagnum shoots are long-lived they may be limited by stochastic periods of cold temperature. Within Europe, species are associated with climate gradients along north-south (cold-warm) and oceanic-continental (wet-dry) clines. Within mires, species are sorted along a moisture (hummock-hollow) gradient. In this thesis I examined species responses to and recovery from freezing (I). I compared species with different water level niches in traits related to water management of individual shoots and colonies (II). Using distribution modelling of GBIF data, I estimated how different aspects of climate contributed to Sphagnum species distributions in Europe (III). Combining the approaches in papers II and III, I modelled the climatic distributions of the parapatric species S. cuspidatum and S. lindbergii and assessed how traits of water economy varied across the distribution boundary (IV). Species responses to winter stress were largely allied to both their hydrological niche and geographic range. Generally, hollow species managed better than hummock species, but species from intermediate positions were less consistent in their response. Species associated with boreal regions were generally less affected than those from temperate regions. Hardening against low temperature was triggered by shorter days and cold nights. Cold temperatures during late autumn may be more important for Sphagnum limits than the minimum temperature during winter. Water-related traits split the species into two groups; hollows species with large capitula and hummock species with small capitula. However, inter- and intra-specific trait variation and trait trends along the hydrological gradient were not necessarily the same at the shoot and canopy scale. Some trait correlations were common to all species. Canopy traits, which were emergent traits of colonies of shoots, had the strongest trait associations with the species position along the hummock-hollow gradient. At the continental scale the distribution of most Sphagnum species could be successfully modelled by a combination of annual degree days and water balance and the degree of seasonality in these two variables. Individual species distributions were shaped more by the seasonality in degree days than in water balance. Across the distributional border of S. cuspidatum and S. lindbergii divergence in the measured traits was mostly seen in the capitula indicating that limits to Sphagnum species are strongly linked to the functioning of the capitulum. Capitulum mass of both species was lower in sympatry than in allopatry, even though the measured values were similar. Canopy traits most strongly separated the species though did not change across the species boundaries. In summary, Sphagnum species in general are limited by the availability of water. Low temperature, particularly during late autumn are probably decisive for the biogeographic limits and for the distribution of species along the hydrological gradient