To what extent can we predict variation of bryophyte and tracheophyte community composition at fine spatial scale along an elevation gradient?

Mounting evidence points to the need for high-resolution climatic data in biodiversity analyses under global change. As we move to finer resolution, other factors than climate, including other abiotic variables and biotic interactions play, however, an increasing role, raising the question of our ability to predict community composition at fine scales. Focusing on two lineages of land plants, bryophytes and tracheophytes, we determine the relative contribution of climatic, non-climatic environmental drivers, spatial effects, community architecture and composition of one lineage to predict community composition of the other lineage, and how our ability to predict community composition varies along an elevation gradient. The relationship between community composition of one lineage and 68 environmental variables at 2-25 m spatial resolution, architecture and composition of the other lineage, and spatial factors, was investigated by hierarchical and variance partitioning across 413 2x2m plots in the Swiss Alps. Climatic data, although significant, contributed less to the model than any other variable considered. Community composition of one lineage, reflecting both direct interactions and unmeasured (hidden) abiotic factors, was the best predictor of community composition of the other lineage. Total explained variance substantially varied with elevation, underlining the fact that the strength of the species composition-environment relationship varies depending on environmental conditions. Total variance explained increased towards high elevation up to 50 %, with an increasing importance of spatial effects and vegetation architecture, pointing to increasing positive interactions and aggregated species distribution patterns in alpine environments. In tracheophytes, an increase of the contribution of non-climatic environmental factors was also observed at high elevation, in line with the hypothesis of a stronger environmental control under harsher conditions. Further improvements of our ability to predict changes in plant community composition may involve the implementation of historical variables and higher-resolution climatic data to better describe the microhabitat conditions actually experienced by organisms.

Morphological, genetic and ecological divergence in near-cryptic bryophyte species widespread in the Holarctic: the Dicranum acutifolium complex (Dicranales) revisited in the Alps.

There is mounting evidence that reproductively isolated, but morphologically weakly differentiated species (so-called cryptic species) represent a substantial part of biological diversity, especially in bryophytes. We assessed the evolutionary history and ecological differentiation of a species pair, Dicranum brevifolium and D. septentrionale, which have overlapping ranges in the Holarctic. Despite their morphological similarity, we found similar genetic differentiation as between morphologically well-differentiated Dicranum species. Moreover, we detected gene tree discordance between plastid and nuclear markers, but neither of the two datasets resolved the two as sister species. The signal in trnL-trnF better reflects the morphological and ecological affinities and indicates a close relationship while ITS sequence data resolved the two taxa as phylogenetically distantly related. The discordance is probably unrelated to the ecological differentiation of D. septentrionale to colonise subneutral to alkaline substrates (vs. acidic in D. brevifolium), because this ability is rare in the genus and shared with D. acutifolium. This taxon is the closest relative of D. septentrionale according to the trnL-trnF data and does not share the discordance in ITS. We furthermore demonstrate that beside D. acutifolium, both D. septentrionale and D. brevifolium occur in the Alps but D. brevifolium is most likely rarer. Based on morphological analyses including factor analysis for mixed data of 45 traits we suggest treating the latter two as near-cryptic species and we recommend verifying morphological determinations molecularly.

Thermophilisation of communities differs between land plant lineages, land use types and elevation.

Bryophytes provide key ecosystem services at the global scale such as carbon storage and primary production in resource limited habitats, but compared to vascular plants knowledge on how these organisms face recent climate warming is fragmentary. This is particularly critical because bryophytes differ fundamentally from vascular plants in their ecophysiological and biological characteristics, so that community alterations most likely have different dynamics. In a comparative approach, we analysed thermophilisation of bryophyte and vascular plant communities in 1146 permanent plots distributed along an elevational gradient of nearly 3.000 m in Switzerland (Central Europe) that were visited in 5-years intervals between 2001 and 2021. We estimated thermophilisation from changes in unweighted mean temperature indicator values of species, compared it to expected thermophilisation rates given the shift of isotherms and addressed differences between the two lineages, major land use types (managed grasslands, forests, unmanaged open areas), life strategy types (long- and short-lived species) and in elevation. Thermophilisation of bryophyte communities was on average 2.1 times higher than of vascular plant communities and at high elevations it approximated the expected rate given the shift of isotherms. Thermophilisation of both, bryophyte and vascular plant communities was not driven by a loss of cryophilic species but by an increase in thermophilic and mesophilic species, indicating an in-filling process. Furthermore, our data show that thermophilisation is higher in managed grasslands than in forests. We suggest that the higher responsiveness of bryophytes compared to vascular plants depends on their poikilohydry and dispersal capacity and that lower thermophilisation of forests communities is related to the buffering effect of microclimatic conditions in the interior of forests. Our study emphasises the heterogeneity of climate warming effects on plants because response dynamics differ between taxonomic groups as well as between land use types and along elevational gradients.

Epiphytes in wooded pastures: Isolation matters for lichen but not for bryophyte species richness.

Sylvo-pastoral systems are species-rich man-made landscapes that are currently often severely threatened by abandonment or management intensification. At low tree densities, single trees in these systems represent habitat islands for epiphytic cryptogams. Here, we focused on sycamore maple (Acer pseudoplatanus) wooded pastures in the northern European Alps. We assessed per tree species richness of bryophytes and lichens on 90 sycamore maple trees distributed across six study sites. We analysed the effects of a range of explanatory variables (tree characteristics, environmental variables and isolation measures) on the richness of epiphytic bryophytes and lichens and various functional subgroups (based on diaspore size, habitat preference and red list status). Furthermore, we estimated the effect of these variables on the occurrence of two specific bryophyte species (Tayloria rudolphiana, Orthotrichum rogeri) and one lichen species (Lobaria pulmonaria) of major conservation concern. Bryophytes and lichens, as well as their subgroups, were differently and sometimes contrastingly affected by the variables considered: tree diameter at breast height had no significant effect on bryophytes but negatively affected many lichen groups; tree phenological age positively affected red-listed lichens but not red-listed bryophytes; increasing isolation from neighbouring trees negatively affected lichens but not bryophytes. However, the high-priority bryophyte species T. rudolphiana was also negatively affected by increased isolation at small spatial scales. Orthotrichum rogeri was more frequent on young trees and L. pulmonaria was more frequent on trees with thin stems and large crowns. The results indicate that local dispersal is important for lichens, whereas long distance dispersal seems to be more important for colonisation by bryophytes. Furthermore, our study highlights that different conservation measures need to be taken depending on the taxonomic and functional species group or the individual species that is addressed. In practice, for the conservation of a high overall richness in sylvo-pastoral systems, it is crucial to sustain not only old and large trees but rather a wide range of tree sizes and ages.