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Northern peatlands are globally important carbon stores. With increasing fire frequency, the re-establishment of bryophytes becomes crucial for their carbon sequestration. Smoke-responsive germination is a common trait of seeds in fire-prone ecosystems but has not been demonstrated in bryophytes. To investigate the potential role of smoke in post-fire peatland recovery, we tested the germination of spores of 15 bryophyte species after treatment with smoke-water. The smoke responsiveness of spores with different laboratory storage times and burial depths/age (3–200 years) was subsequently tested. Smoke increased the germination percentage for 10 of the species and the germination speed for four of these. Smoke responsiveness increased along the fire frequency gradient from open expanse to forest margin, consistent with the theory that this selects for the maintenance of fire-adapted traits. Smoke enhanced the germinability of 1-year but not 4-year laboratory-stored spores, and considerably increased the germinability of spores naturally buried in peat for up to ~200 years. The effect of fire may be overlooked in non-fire-prone ecosystems, such as those in which wetland bryophytes dominate. Our study reveals a mechanism by which an increase in fire frequency may lead to shifts in species dominance, which may affect long-term carbon sequestration in peatlands.

Comision Nacional de Investigacion Cientifica y Tecnologica (CONICYT) CONICYT FONDECYT 11150301 11160152

Long amplicon metabarcoding has opened the door for phylogenetic analysis of the largely unknown communities of microeukaryotes in soil. Here, we amplified and sequenced the ITS and LSU regions of the rDNA operon (around 1500 bp) from grassland soils using PacBio SMRT sequencing. We tested how three different methods for generation of operational taxonomic units (OTUs) effected estimated richness and identified taxa, and how well large-scale ecological patterns associated with shifting environmental conditions were recovered in data from the three methods. The field site at Kungsängen Nature Reserve has drawn frequent visitors since Linnaeus's time, and its species rich vegetation includes the largest population of Fritillaria meleagris in Sweden. To test the effect of different OTU generation methods, we sampled soils across an abrupt moisture transition that divides the meadow community into a Carex acuta dominated plant community with low species richness in the wetter part, which is visually distinct from the mesic-dry part that has a species rich grass-dominated plant community including a high frequency of F. meleagris. We used the moisture and plant community transition as a framework to investigate how detected belowground microeukaryotic community composition was influenced by OTU generation methods. Soil communities in both moisture regimes were dominated by protists, a large fraction of which were taxonomically assigned to Ciliophora (Alveolata) while 30%–40% of all reads were assigned to kingdom Fungi. Ecological patterns were consistently recovered irrespective of OTU generation method used. However, different methods strongly affect richness estimates and the taxonomic and phylogenetic resolution of the characterized community with implications for how well members of the microeukaryotic communities can be recognized in the data.

Long amplicon metabarcoding has opened the door for phylogenetic analysis of the largely unknown communites of microeukaryotes in soil. Here, we amplified and sequenced the ITS and LSU regions of the rDNA operon (around 1500 bp) from grassland soils using PacBio SMRT sequencing and evaluated the performance of three different methods for generation of operational taxonomic units (OTUs). The field site at Kungsängen Nature Reserve has drawn frequent visitors since Linnaeus’s time, and its species rich vegetation includes the largest population of Fritillaria meleagris in Sweden. To test the effect of different OTU generation methods, we sampled soils across an abrupt moisture transition that divides the meadow community into a Carex acuta dominated plant community with low species richness in the wetter part, which is visually distinct from the mesic-dry part that has a species rich grass-dominated plant community including a high frequency of F. meleagris. We used the moisture and plant community transition as a framework to investigare how detected belowground microeukaryotic community composition was influenced by OTU generation methods. Soil communities in both moisture regimes were dominated by protists, a large fraction of which were taxonomically assigned to Ciliophora (Alveolata) while 30-40% of all reads were assigned to kingdom Fungi. Ecological patterns were consistently recovered irrespective of OTU generation method used. However, different methods strongly affect richness estimates and the taxonomic and phylogenetic resolution of the characterized community with implications for how well members of the microeukaryotic communities can be recognized in the data.

Species distribution models (SDM) predicting changes in species occurrences and abundance are increasingly being used as a tool in biogeography and conservation biology. However, we have little information on their predictive performance. Here we used archive-recorded predictor and field-observational verifier data associated with water level to evaluate the performance of response curves over 40 years for marsh plant species in Northeast China. A consensus approach (AUC: area-under-curve) was used as the test measure for internal evaluation and external evaluation (forecast and hindcast). Our results demonstrated that there is no significant differences between internal and external evaluation, and they both showed reasonable accuracy (AUC = 0.73, respectively). There was considerable variation across species and projection direction in model accuracy, and accuracy of model fitting in internal evaluation and restricting the environmental range of data in different time periods may impact the performance of model projection over time. The performance of generalized additive models (GAM) is similar with that of extended Huisman-Olff-Fresco models (eHOF). Cover model is a little better than presence/absence models in prediction over time. Our findings provide some guidelines for the use of SDM for predictions under environmental change.

There were mistakes in the data of three variables used in this study. The corrections listed below have minor effects on statistics and p-values, but do not change any conclusions. Correct data are stored in the Dryad repository ( https://doi.org/10.5061/dryad.4f8d2.2 ).

The response of photosynthetic CO2 exchange to tissue water content in five spp. of Sphagnum from contrasting microhabitats (S. fuscum (Schimp.) Klinggr., S. papillosum H. Lindb., S. magellanicum B ...

The effects of increased nitrogen influx on Sphagnum growth and on interspecific competition between Sphagnum species were studied in a 3-yr experiment in mires situated in two areas with different rates of airborne N deposition. Sphagnum growth was recorded after various supplementary N influxes (0, 1, 3, 5 and 10 g m −2 yr−1) in hummocks and lawn communities. Sphagnum biomass production decreased with increasing N influx in both areas. After the first season at the low-deposition site, Sphagnum showed an increased growth in length with the intermediate N treatment, but in the second and third seasons the control treatment had the highest growth in length. Capitulum dry mass increased with increasing N influx. Sphagnum N concentration and N/P quotient were higher at the high- than at the low-deposition site. The low quotient at the low-deposition site, together with the initial growth increase with intermediate N supplements, indicates that growth was N-limited at this site, but our lowest N supplement was sufficient to reduce growth. The N treatments had no effect on interspecific competition between the Sphagnum species. This indicates that the species have similar responses to N. The species studied all occur naturally on ombrotrophic, N-poor sites and show low tolerances to increased N influx. Reduced Sphagnum production may affect the carbon balance, changing the mires from C sinks to sources.

1. The relative importance of global versus local environmental factors for growth and thus carbon uptake of the bryophyte genus Sphagnum—the main peat‐former and ecosystem engineer in northern peatlands—remains unclear. 2. We measured length growth and net primary production (NPP) of two abundant Sphagnum species across 99 Holarctic peatlands. We tested the importance of previously proposed abiotic and biotic drivers for peatland carbon uptake (climate, N deposition, water table depth and vascular plant cover) on these two responses. Employing structural equation models (SEMs), we explored both indirect and direct effects of drivers on Sphagnum growth. 3. Variation in growth was large, but similar within and between peatlands. Length growth showed a stronger response to predictors than NPP. Moreover, the smaller and denser Sphagnum fuscum growing on hummocks had weaker responses to climatic variation than the larger and looser Sphagnum magellanicum growing in the wetter conditions. Growth decreased with increasing vascular plant cover within a site. Between sites, precipitation and temperature increased growth for S. magellanicum. The SEMs indicate that indirect effects are important. For example, vascular plant cover increased with a deeper water table, increased nitrogen deposition, precipitation and temperature. These factors also influenced Sphagnum growth indirectly by affecting moss shoot density. 4. Synthesis. Our results imply that in a warmer climate, S. magellanicum will increase length growth as long as precipitation is not reduced, while S. fuscum is more resistant to decreased precipitation, but also less able to take advantage of increased precipitation and temperature. Such species‐specific sensitivity to climate may affect competitive outcomes in a changing environment, and potentially the future carbon sink function of peatlands.

Background and AimsThe ecosystem engineers Sphagnum (peat mosses) are responsible for sequestering a large proportion of carbon in northern peatlands. Species may respond differently to hydrological changes, and water level changes may lead to vegetation shifts in peatlands, causing them to revert from sinks to sources of carbon. We aimed to compare species-specific responses to water level drawdown within Sphagnum, and investigate which traits affect water economy in this genus.MethodsIn a mesocosm experiment, we investigated how water level drawdown affected water content (WC) in the photosynthetically active apex of the moss and maximum quantum yield of photosystem II (i.e. Fv/Fm) of 13 Sphagnum species. Structural traits were measured, and eight anatomical traits were quantified from scanning electron microscopy micrographs.Key ResultsMixed-effects models indicated that at high water level, large leaves were the most influential predictor of high WC, and at low water level WC was higher in species growing drier in the field, with larger hyaline cell pore sizes and total pore areas associated with higher WC. Higher stem and peat bulk density increased WC, while capitulum mass per area and numerical shoot density did not. We observed a clear positive relationship between Fv/Fm and WC in wet-growing species.ConclusionsWhile we found that most hummock species had a relatively high water loss resistance, we propose that some species are able to maintain a high WC at drawdown by storing large amounts of water at a high water level. Our result showing that leaf traits are important warrants further research using advanced morphometric methods. As climate change may lead to more frequent droughts and thereby water level drawdowns in peatlands, a mechanistic understanding of species-specific traits and responses is crucial for predicting future changes in these systems.

The information of species' response (optimum or critical limits along environmental gradients) is a key to understanding ecological questions and to design management plans. A large number of plots (762) from 70 transects of 13 wetland sites in Northeast China were sampled along flooding gradient from marsh to wet meadow. Species response (abundance and occurrence) to flooding were modelled with Generalized Additive Models for 21 dominant plant species. We found that 20 of 21 species showed a significant response to flooding for the occurrence and abundance models, and four types of response were found: monotonically increasing, monotonically decreasing, skewed unimodal and symmetric unimodal. The species with monotonically increasing response have the deepest flooding optimum and widest niche width, followed by those with unimodal curve, and the monotonically decreasing ones have the smallest values. The optima and niche width (whether based on occurrence or abundance models) both significantly correlated with the frequency, but not with mean abundance. Abundance models outperformed occurrence models based on goodness of fit. The abundance models predicted a rather sharp shift from dominance of helophytes (Carex pseudo-curaica and C. lasiocarpa) to wet meadow species (Calamagrostis angustifolia and Carex appendiculata) if water levels drop from about 10cm above soil surface to below the surface. The defined optima and niche width based on the abundance models can be applied to better instruct restoration management. Given the time required to collect abundance data, an efficient strategy could be to monitor occurrence in many plots and abundance in a subset of these.

Many endangered plant species remain absent in rewetted, previously drained fens. We performed a 3-year introduction experiment with endangered fen species (9 Carex- and 6 bryophyte species) in 4 hydrologically restored fens to investigate which factors hamper establishment and survival. Carex species were introduced as adults and seedlings, mosses as gametophytes. Introductions were done on (initially) bare soil, which allowed us to exclude excessive competition for light during the first year. First year survival of the transplants was high in all fens (mean survival = 96%), indicating that there were no direct abiotic constraints on establishment. However, survival analysis revealed that a decrease in relative light intensity (RLI) at the soil surface during consecutive years (indicating an increase in biotic competition for light) drove high mortality rates in most species. As a result, overall final survival was lowest in the two most productive (low light) fens (mean survival = 38%), while most transplants persisted in the two less productive (high light) fens (mean survival = 79%). Taller and faster-growing Carex species were able to outgrow light limitation near the soil surface, and thus had a higher overall survivability than smaller and slower-growing species. Light limitation also drove the loss of 5 out of 6 bryophyte species. We conclude that both dispersal limitation and asymmetric competition for light may explain the lack and loss of small and endangered plant species in rewetted fens. A minimum empirical threshold of c. 30% relative light intensity near the soil surface is required for successful introduction.

Question We quantify tree dynamics over a century of free development in a small broadleaved forest dominated by Fraxinus excelsior and Ulmus glabra. What are the internal and external factors driving the changes, and how predictable are they? What were the time scale and effects of the spread of Dutch elm disease (DED)? Location Vardsatra, eastern central Sweden. Methods The survival, growth and recruitment of all trees (≥ 12 cm in girth) were monitored in 1912, 1967, 1988 and 2013 (more often for a part of the forest). Woody species in the field and shrub layers were surveyed in permanent plots in 1976 and 2012. We used transition matrix models to project changes in population sizes and species composition within the century and for 2050. Results The results indicate that the forest was in a successional development during the first period. The species composition had stabilised by 1967, except for an expansion of Acer platanoides and the drastic effect of DED that struck the forest around 2000. It took only a decade to kill virtually all large elms in the forest, leading to strong decrease in stem density and basal area. The evidence for effects of DED is still weak, but there has been an increase in saplings, notably of Fraxinus, Prunus padus, Ulmus, and of shoots of Corylus avellana. Several species that are abundant in the vicinity and as seeds fail to establish (Picea abies, Betula spp., Quercus robur, Populus tremula). Projections for 2050 based on the third period (1988-2013) are probably unrealistic since also Fraxinus may disappear because of the recent arrival of the ash dieback. Conclusions Slow dynamics in forests that could follow from climate change will locally probably be overruled by unforeseen catastrophes, such as invasions by forest pathogens. These initiate changes with long lag phases difficult to quantify. Still, a dense deciduous forest can resist invasion of colonist species and of regionally dominant conifers; the reason being unfavourable conditions for establishment rather than dispersal limitation This article is protected by copyright. All rights reserved.

1. Functional traits mechanistically capture plant responses to environmental gradients as well as plant effects on ecosystem functioning. Yet most trait-based theory stems from terrestrial systems ...

Mires (bogs and fens) are nutrient-limited peatland ecosystems, the vegetation of which is especially sensitive to nitrogen deposition and climate change. The role of mires in the global carbon cycle, and the delivery of different ecosystem services can be considerably altered by changes in the vegetation, which has a strong impact on peat-formation and hydrology. Mire ecosystems are commonly open with limited canopy cover but both nitrogen deposition and increased temperatures may increase the woody vegetation component. It has been predicted that such an increase in tree cover and the associated effects on light and water regimes would cause a positive feed-back loop with respect to the ground vegetation. None of these effects, however, have so far been confirmed in large-scale spatiotemporal studies. Here we analyzed data pertaining to mire vegetation from the Swedish National Forest Inventory collected from permanent sample plots over a period of 20 yr along a latitudinal gradient covering 14°. We hypothesized that the changes would be larger in the southern parts as a result of higher nitrogen deposition and warmer climate. Our results showed an increase in woody vegetation with increases in most ericaceous dwarf-shrubs and in the basal area of trees. These changes were, in contrast to our expectations, evenly distributed over most of the latitudinal gradient. While nitrogen deposition is elevated in the south, the increase in temperatures during recent decades has been larger in the north. Hence, we suggest that different processes in the north and south have produced similar vegetation changes along the latitudinal gradient. There was, however, a sharp increase in compositional change at high deposition, indicating a threshold effect in the response. Instead of a positive feed-back loop caused by the tree layer, an increase in canopy cover reduced the changes in composition of the ground vegetation, whereas a decrease in canopy cover lead to larger changes. Increased natural disturbances of the tree layer due to, for example, pathogens or climate is a predicted outcome of climate change. Hence, these results may have important implications for predictions of long-term effects of increased temperature on peatland vegetation.

Introduction.Sphagnum L. forms much of the ground cover in northern peatlands. Different species show affinities for bioclimatic regions in Europe (oceanic/continental; northern/southern) and species-specific tolerance of winter conditions can be a factor explaining their distribution. Methods. We focussed on low temperature in a series of experiments and tested (1) the innate ability of a selection of Sphagnum species to tolerate low temperature in relation to their micro-topographic (wetness) and geographical (climate) distribution; (2) the rate of cold tolerance acquisition; and (3) the ability of species to survive a range of low temperature once cold hardened. Key results. Our experiments showed that maximal PSII efficiency (Fv/Fm, chlorophyll fluorescence), growth rates and survival were all negatively affected by sub-zero temperatures. Environmental conditions associated with the onset of winter (colder nights and shorter days) triggered the acquisition of cold tolerance in Sphagnum. Conclusions. The results were not unequivocal, but species associated with colder climates were generally more tolerant of sub-zero conditions. Species associated with the wettest and driest ends of the wetness gradient were more consistent in their responses than those in between, with wetter-dwelling species being less sensitive to sub-zero temperature than species found in drier microhabitats. Overall, our results suggest that adaptation to winter conditions contribute to the current distribution patterns of Sphagnum species.

Rain-fed peatlands are dominated by peat mosses (Sphagnum sp.), which for their growth depend on elements from the atmosphere. As the isotopic composition of carbon (12,13C) and oxygen (16,18O) of these Sphagnum mosses are affected by environmental conditions, the dead Sphagnum tissue accumulated in peat constitutes a potential long-term archive that can be used for climate reconstruction. However, there is a lack of adequate understanding of how isotope values are influenced by environmental conditions, which restricts their current use as environmental and palaeoenvironmental indicators. Here we tested (i) to what extent C and O isotopic variation in living tissue of Sphagnum is species-specific and associated with local hydrological gradients, climatic gradients (evapotranspiration, temperature, precipitation), and elevation; (ii) if the C isotopic signature can be a proxy for net primary productivity (NPP) of Sphagnum; and (iii) to what extent Sphagnum tissue δ18O tracks the δ18O isotope signature of precipitation. In total, we analysed 337 samples from 93 sites across North America and Eurasia using two important peat-forming Sphagnum species (S. magellanicum, S. fuscum) common to the Holartic realm. There were differences in δ13C values between species. For S. magellanicum δ13C decreased with increasing height above the water table (HWT, R2 = 17 %) and was positively correlated to productivity (R2 = 7 %). Together these two variables explained 46 % of the between-site variation in δ13C values. For S. fuscum, productivity was the only significant predictor of δ13C (total R2 = 6 %). For δ18O values, ca. 90 % of the variation was found between sites. Globally-modelled annual δ18O values in precipitation explained 69% of the between-site variation in tissue δ18O. S. magellanicum showed lower δ18O enrichment than S. fuscum (−0.83 ‰ lower) . Elevation and climatic variables were weak predictors of tissue δ18O values after controlling for δ18O values of the precipitation. To summarise, our study provides evidence for (a) good predictability of tissue δ18O values from modelled annual δ18O values in precipitation, and (b) the possibility to relate tissue δ13C values to HWT and NPP, but this appears to be species-dependent. These results suggest that isotope composition can be used at a large scale for climatic reconstructions but that such models should be species-specific.

Considerable progress has been made in ecological and evolutionary genetics with studies demonstrating how genes underlying plant and microbial traits can influence adaptation and even 'extend' to influence community structure and ecosystem level processes. Progress in this area is limited to model systems with deep genetic and genomic resources that often have negligible ecological impact or interest. Thus, important linkages between genetic adaptations and their consequences at organismal and ecological scales are often lacking. Here we introduce the Sphagnome Project, which incorporates genomics into a long-running history of Sphagnum research that has documented unparalleled contributions to peatland ecology, carbon sequestration, biogeochemistry, microbiome research, niche construction, and ecosystem engineering. The Sphagnome Project encompasses a genus-level sequencing effort that represents a new type of model system driven not only by genetic tractability, but by ecologically relevant questions and hypotheses.

Background and aims Sphagnum mosses are ecosystem engineers that create and maintain boreal peatlands. With unique biochemistry, waterlogging and acidifying capacities, they build up meters-thick layers of peat, reducing competition and impeding decomposition. We quantify within-genus differences in biochemical composition to make inferences about decay rates, related to hummock–hollow and fen–bog gradients and to phylogeny. Methods We sampled litter from 15 Sphagnum species, abundant over the whole northern hemisphere. We used regression and Principal Components Analysis (PCA) to evaluate general relationships between litter quality parameters and decay rates measured under laboratory and field conditions. Results Both concentrations of the polysaccharide sphagnan and the soluble phenolics were positively correlated with intrinsic decay resistance, however, so were the previously understudied lignin-like phenolics. More resistant litter had more of all the important metabolites; consequently, PC1 scores were related to lab mass loss (R2 = 0.57). There was no such relationship with field mass loss, which is also affected by the environment. PCA also revealed that metabolites clearly group Sphagnum sections (subgenera). Conclusions We suggest that the commonly stated growth-decomposition trade-off is largely due to litter quality. We show a strong phylogenetic control on Sphagnum metabolites, but their effects on decay are affected by nutrient availability in the habitat.

Question Species interactions are one of the processes determining composition of plant communities. We used the community density series method to study competition in a multi-species community of bryophytes common in calcareous fens. The succession of mires is driven by Sphagnum species, which are supposedly superior to brown mosses in competition for resources and space, but little is known about the environmental conditions in which brown mosses can prevail when subject to neighbour interactions. How are interactions among peatland bryophytes affected by the environment? Location Field and garden experiments near Uppsala in mid-eastern Sweden. Methods To examine the effects of environment on competition and competitive hierarchies we assembled multi-species communities of ten bryophyte species from shoot fragments (brown mosses and Sphagnum species) at two densities and grew them on three types of peat (representing poor, intermediate and rich fens) under dry or wet conditions in a garden experiment and along pH and wetness gradients in the field. Results A multivariate analysis of the garden experiment showed that community composition was affected by peat type and wetness and their interactions. The brown mosses performed better in wet and rich fens, the Sphagnum species in drier and poorer fens. The Sphagnum species were overall the best competitors. Conclusions The experiments demonstrated contrasting responses of brown mosses and Sphagnum to properties of the microhabitat. Sphagnum species were generally less affected by competition than the brown mosses. Sphagnum species were competitive in habitats typically dominated by brown mosses and even responded positively to crowding. This can explain why Sphagnum can invade calcareous fens. In contrast, brown mosses performed poorly in habitats more typical of Sphagnum species.

The species pool theory helps us understand species distributions at different geographical scales. In theory, species pools consist of species passing the filters between different geographical scales. Filters of dispersal and environment act between the regional and local pools, while filters of biotic interactions act between the local and the community pools. We studied bryophytes and vascular plants restricted to rich (calcareous) fens that occur as patches in a forested landscape. We then examined their frequencies and abundances at the local and regional scales and related the results to traits important for dispersal and competition. Our results show that weft-forming bryophytes and vascular plants with far-creeping rhizomes have higher local frequency than predicted from their regional frequency. Dispersal traits did not explain any variation in the distributions. This indicates that environmental and biotic filters are more important than dispersal limitation at the regional scale, and that clonal expansion is the most important factor for high frequency and abundance at the local scale in these nutrient poor habitats.

Peat mosses (Sphagnum) are ecosystem engineers-species in boreal peatlands simultaneously create and inhabit narrow habitat preferences along two microhabitat gradients: an ionic gradient and a hydrological hummock-hollow gradient. In this article, we demonstrate the connections between microhabitat preference and phylogeny in Sphagnum. Using a dataset of 39 species of Sphagnum, with an 18-locus DNA alignment and an ecological dataset encompassing three large published studies, we tested for phylogenetic signal and within-genus changes in evolutionary rate of eight niche descriptors and two multivariate niche gradients. We find little to no evidence for phylogenetic signal in most component descriptors of the ionic gradient, but interspecific variation along the hummock-hollow gradient shows considerable phylogenetic signal. We find support for a change in the rate of niche evolution within the genus-the hummock-forming subgenus Acutifolia has evolved along the multivariate hummock-hollow gradient faster than the hollow-inhabiting subgenus Cuspidata. Because peat mosses themselves create some of the ecological gradients constituting their own habitats, the classic microtopography of Sphagnum-dominated peatlands is maintained by evolutionary constraints and the biological properties of related Sphagnum species. The patterns of phylogenetic signal observed here will instruct future study on the role of functional traits in peatland growth and reconstruction.

Growth and decomposition of Sphagnum controls turnover of a large global store of soil organic carbon. We investigated variation in morphological and physiological traits of Sphagnum shoots, and related this variation to canopy variables relevant to peatland carbon cycling. We sampled Sphagnum along a bog plateau-swamp forest gradient and measured a suite of shoot traits and canopy variables. Major axes of variation were identified using principal component analysis and correlated with canopy variables such as growth, biomass and decomposition. We also examined scaling of shoot traits with one another and with canopy variables. Two distinct tradeoffs in shoot traits emerged. From dry to wet habitats, individual metabolic rates and capitulum size increased while numerical density decreased, leading to faster growth and elongation on an individual basis. From treed to open habitats, photosynthetic efficiency decreased and photosynthetic biomass increased, driving faster growth on an area basis and slower litter mass loss. The tradeoffs identified have important implications for peatlands undergoing climate-related changes in water and light availability. Sphagnum trait comparisons, combined with scaling analyses, offer a promising approach to understanding and predicting the effects of environmental change on peatland carbon cycling.

Endozoochory is usually involved in seed dispersal mutualisms, whereas ectozoochory is non-rewarding, and therefore neutral (or even negative) for the animal vector. Synzoochory is an intermediate dispersal type between endo and ectozoochory in which propagules are deliberately transported (usually in the mouth) but with no ingestion or gut passage involved. We present empirical evidence of synzoochoric mutualism between the hummingbird Sephanoides sephaniodes and cryptogams (one fern and seven moss species). Two species (Lophosoria quadripinnata and Ancistrodes genuflexa) constituted the bulk of nest biomass, and another six moss species were present in lesser quantity. The hummingbird was selective when collecting nest material so that the nests contained a higher density of reproductive structures (that could be dispersed further) than natural patches of the cryptogam species. Even after one year, the nests maintained half of the original reproductive structures (sporangia, sporophytes) and biomass, constituting an important dispersal source. These results show a new type of mutualism in which mosses could be dispersed throughout longer distances (several km) by hummingbirds and to higher positions (particularly for ground-living species, promoting dispersal potential). The hummingbird benefits from collecting cryptogam material for nest building, and cryptogams benefit from the concentration and relocation of diaspore sources into more effective recruiting sites. Similar mutualistic relationships could be a general phenomenon, of importance in many ecosystems.

The stress-gradient hypothesis (SGH) predicts that the relative importance of competition decreases and facilitation increases with an increase in abiotic stress. In peatlands, Sphagnum faces the threat of drought and differentiates into hummock species (drought-tolerant) and hollow species. Whether interspecific interaction affects the influence of drought on bryophyte composition in peatlands is unknown. We established an experiment by simulating drought and building bryophyte communities with two hummock species (S. palustre and S. capillifolium) and one hollow species (S. fallax). In all three species, drought decreased biomass production, height increment and side-shoot production. Sphagnum stores water in the hyaline cells, and leaf hyaline cell percentage (HCP) in the two hummock species increased with drought while no effect was found in S. fallax, suggesting that adjusting HCP is not an effective response to drought for the hollow species. Morphological traits and carbon and nitrogen contents in hummock species responded more to drought than in the hollow species, indicating a rapid response in phenotypic plasticity is an important strategy to resist drought in the hummock species. The presence of neighboring Sphagnum species, rather than drought, decreased carbon content for all three species. All three bryophytes showed interaction between drought and neighbor in two or more plant traits. Our study, however, did not support SGH, and there were no changes from competition under wet to facilitation under dry treatments in any of the six species combinations. On the contrary, when S. fallax was the target species, a change from facilitation under wet to competition under dry treatments was observed. The results suggest that hummock species can facilitate hollow species in wet environments but they could suppress hollow species under drought conditions by competing for water resources. Both drought and strong competition are the probable reasons why hollow species rarely grow in hummocks.

Questions: Ecological restoration has traditionally been evaluated with analyses focused on species identities and abundances. These analyses provide no ecological explanation to why certain specie ...

Peat mosses (Sphagnum) largely govern carbon sequestration in Northern Hemisphere peatlands. We investigated functional traits related to growth and decomposition in Sphagnum species. We tested the importance of environment and phylogeny in driving species traits and investigated trade‐offs among them. We selected 15 globally important Sphagnum species, representing four sections (subgenera) and a range of peatland habitats. We measured rates of photosynthesis and decomposition in standard laboratory conditions as measures of innate growth and decay potential, and related this to realized growth, production, and decomposition in their natural habitats. In general, we found support for a trade‐off between measures of growth and decomposition. However, the relationships are not strong, with r ranging between 0.24 and 0.45 for different measures of growth versus decomposition. Using photosynthetic rate to predict decomposition in standard conditions yielded R 2 = 0.20. Habitat and section (phylogeny) affected the traits and the trade‐offs. In a wet year, species from sections Cuspidata and Sphagnum had the highest production, but in a dry year, differences among species, sections, and habitats evened out. Cuspidata species in general produced easily decomposable litter, but their decay in the field was hampered, probably due to near‐surface anoxia in their wet habitats. In a principal components analysis, PCA, photosynthetic capacity, production, and laboratory decomposition acted in the same direction. The species were imperfectly clustered according to vegetation type and phylogeny, so that some species clustered with others in the same section, whereas others clustered more clearly with others from similar vegetation types. Our study includes a wider range of species and habitats than previous trait analyses in Sphagnum and shows that while the previously described growth–decay trade‐off exists, it is far from perfect. We therefore suggest that our species‐specific trait measures offer opportunities for improvements of peatland ecosystem models. Innate qualities measured in laboratory conditions translate differently to field responses. Most dramatically, fast‐growing species could only realize their potential in a wet year. The same species decompose fast in laboratory, but their decomposition was more retarded in the field than that of other species. These relationships are crucial for understanding the long‐term dynamics of peatland communities.

Summary Epiphytes are an important component in many forest ecosystems. The proportion of threatened epiphyte species is high, and the impact of clearcuts on key demographic processes via edge-influence is still poorly understood. There are few studies on epiphyte growth, and even less is known about how reproduction is affected by proximity to forest edges. For retention trees, demographic studies are even scarcer. Based on the results from a 6-month transplant experiment and a 3-year study of natural colonies, we modelled growth and reproduction of epiphytic bryophytes used as indicators of old-growth forests and widespread epiphytes in relation to distance from the forest edge. We also modelled growth and reproduction on retention trees within the clearcut. Species responses were linked to variation in canopy openness. Unlike the widespread species, the old-growth-forest indicators grew exponentially with distance from the edge, and this response was more pronounced at the south-east than north-west forest edge. In one red-listed species, reproduction was thoroughly inhibited near the edge, whereas the reproductive rate of the widespread species tended to increase. However, the widespread species also showed reduced shoot lengths on the retention trees. Reduced growth and inhibited reproduction of sensitive epiphytes near edges decrease the number of dispersing diaspores and may, in combination with lower local connectivity and increased tree fall rates close to edges, increase the risk of metapopulation extinction. Synthesis and applications. Two general management implications for boreal forests are drawn. First, retention trees may not have the capacity to act as a ‘lifeboat’ for epiphytic bryophytes and support their populations during the regeneration phase. Second, the creation of buffer zones is a useful conservation strategy for bryophytes. The exact width of zones depends on the forest structure and should be orientated in relation to the requirements of the most sensitive species. For the rather dense experimental forest, a width of at least 30 m was required for the south-facing buffer, whereas for the north-facing buffer 10 m was sufficient.

Large peatland areas have been drained for forestry and agricultural purposes, resulting in the decline of characteristic biodiversity. Two measures commonly suggested for restoring drained fens is ditch blocking and tree removal to raise the groundwater table and increase light availability, respectively. In 2002, we initiated factorial restoration experiments, including ditch blocking and tree removal, in three former rich fens that had been drained for forestry purposes. Species cover of vascular plants and bryophytes were monitored during 8 years in permanent plots along transects perpendicular to the ditch for all four treatment combinations. Both methods had positive and independent effects on the cover of wetland vegetation. Specifically, Sphagnum species and wetland bryophytes showed a persistent positive response to both clear cutting and rewetting. Wetland vascular plants and grasses showed a persistent positive response to clear cutting. Sedges and species number responded positively to both clear cutting and ditch blocking, but the response was partly transient, and for species richness the response was limited when restoration methods were applied separately. Rich fen indicators of vascular plants and bryophytes did not respond to any of the restoration treatments. This indicates that species introduction in combination with further habitat restorations may be necessary to re-establish the original rich fen flora. Nevertheless, we conclude that the combination of ditch blocking and clear cutting are effective measures to partly restore wetland vegetation on previously drained and forested fens, while peat subsidence along the ditch may restrict the success further away from ditches.

Summary 1. Bogs are nutrient-poor peatland ecosystems that are sensitive to nitrogen (N) deposition. Production of peat mosses (i.e. the peat-forming genus Sphagnum) is known to decrease under elevated N deposition, but the causal mechanisms are poorly understood. 2. It is predicted that increased N deposition will cause changes in Sphagnum species composition, with fast-growing species benefiting from increased N availability in contrast to slow-growing species. Knowledge of species-specific responses to N availability can help us to understand interspecific competitive relationships. 3. We investigated the direct effects of N application on plant physiology in three Sphagnum species by exposing shoots to a range of N doses (corresponding to depositions of 0–5·6 g m−2 year−1), over 5 months, in a greenhouse experiment. The species investigated included one that grows high above the water-table (Sphagnum fuscum) and two that grow lower down (Sphagnum balticum and Sphagnum fallax). S. fuscum and S. balticum originate from ombrotrophic and S. fallax from minerotrophic environments. To estimate N responses, we measured the performance and light-capture kinetics of the photosynthetic apparatus (maximum photosynthetic rate and Fv/Fm), biomass production, shoot formation, and N and phosphorus (P) concentrations in the tissue. 4. Tissue nitrogen concentration generally increased with N application rate, and photosynthetic rate increased with N concentration, although S. balticum exhibited a unimodal response. With respect to production, a negative response to N application rate was found in S. fallax and S. fuscum (weak), while production in S. balticum was unrelated to application rate. S. fallax was the fastest-growing species, producing two to three times more biomass per shoot compared with the other species. 5. The mismatch between photosynthetic capacity and production could partly be explained by an increased N : P ratio following N application. Phosphorus limitation may not negatively affect photosynthetic capacity, but may hamper production. 6. The fast-growing species S. fallax is considered to benefit from increased N deposition, but we found a negative physiological response, suggesting stoichiometric constraints. Thus, we conclude that responses to N deposition cannot be predicted in a simple way from physiological traits related to growth rate without considering local environmental factors.

International audience; Peatlands in the northern hemisphere have accumulated more atmospheric carbon (C) during the Holocene than any other terrestrial ecosystem, making peatlands long-term C sinks of global importance. Projected increases in nitrogen (N) deposition and temperature make future accumulation rates uncertain. * Here, we assessed the impact of N deposition on peatland C sequestration potential by investigating the effects of experimental N addition on Sphagnum moss. We employed meta-regressions to the results of 107 field experiments, accounting for sampling dependence in the data. * We found that high N loading (comprising N application rate, experiment duration, background N deposition) depressed Sphagnum production relative to untreated controls. The interactive effects of presence of competitive vascular plants and high tissue N concentrations indicated intensified biotic interactions and altered nutrient stochiometry as mechanisms underlying the detrimental N effects. Importantly, a higher summer temperature (mean for July) and increased annual precipitation intensified the negative effects of N. The temperature effect was comparable to an experimental application of almost 4 g N m)2 yr)1 for each 1°C increase. * Our results indicate that current rates of N deposition in a warmer environment will strongly inhibit C sequestration by Sphagnum-dominated vegetation.

Stratigraphic records from peatlands suggest that the shift from a rich fen (calcareous fen) to an ombrotrophic bog can occur rapidly. This shift constitutes a switch from a species-rich ecosystem to a species-poor one with greater carbon storage. In this process, the invasion and expansion of acidifying bog species of Sphagnum (peat mosses) play a key role. To test under what conditions an acidifying bog species could invade a rich fen, we conducted three experiments, contrasting the bog species S. fucsum with the rich-fen species S. warnstorfii and S. teres. We first tested the effect of calcareous water by growing the three species at different constant height above the water table (HWT; 2, 7, and 14 cm) in a rich-fen pool and measured maximum photosynthetic rate and production and difference in length growth as an indicator of competition. In none of the species was the photosynthetic capacity negatively affected when placed at low HWT, but S. fuscum was a weaker competitor at low HWT. In our second experiment we transplanted the three species into microhabitats with different and naturally varying HWT in a rich fen. Here, S. fuscum nearly ceased to photosynthesize when transplanted to low HWT (brown moss carpet), while it performed similarly to the two rich-fen species at the intermediate level (S. warnstorfii hummock level). In contrast to S. fuscum, the rich-fen sphagna performed equally well in both habitats. The brown moss carpet was seasonally flooded, and in our third experiment we found that S. fuscum, but not S. teres, was severely damaged when submerged in rich-fen water. Our results suggest two thresholds in HWT affecting the ecosystem switch: one level that reduces the risk of submergence and a higher one that makes bog sphagna competitive against the rich-fen species.

The aim of this thesis was to increase knowledge about metapopulation and metacommunity processes in patchy, dynamic landscapes, using epiphytic bryophytes as a model system. Host trees and deciduous forest stands in the coniferous landscape are patchy, temporal and undergo changes in habitat quality during succession. Epiphytes must track this dynamic habitat network for their long-term survival. Community patterns at different spatial scales were explored and linked to regional metapopulation processes and local population dynamics. Spatial structuring in species richness both at a local and regional scale indicated stronger dispersal limitation but lower sensitivity to habitat quality in species with large asexual than in species with small sexual diaspores. In sexually dispersed species, a strong rescue effect was indicated by a bimodal frequency distribution of the species and by increasing local abundance with increasing patch connectivity. Present connectivity to other deciduous forest patches had positive effects on richness of asexually dispersed species, whereas richness of sexually dispersed species was instead related to the landscape connectivity 30 years ago. A study of local growth and reproduction suggested that this is caused by delayed sexual, but not asexual, reproduction. Habitat conditions affected the production of sporophytes, but not of asexual diaspores. No differences in either growth rates or competitive abilities among species with different dispersal and life-history strategies were found. In vitro experiments showed that establishment is higher from large asexual diaspores than from small sexual. Establishment of all diaspore types was limited by pH. There were indications of trade-offs between high germination and protonemal growth rates, desiccation tolerance and a rapid development of shoots from protonema. The results indicated that the epiphyte metacommunity is structured by two main trade-offs: dispersal distance (diaspore size) versus age at first reproduction, and dispersal distance versus sensitivity to habitat quality. Trade-offs in species traits may have evolved as a consequence of conflicting selection pressures imposed by habitat turnover, connectivity and irregular water supply rather than by species interactions. Syndromes of interrelated species traits imply that fairly small changes in habitat conditions can lead to distinct changes in metacommunity diversity: the results indicate that increasing distances among patches cause most harm to asexually dispersed species, whereas cuttings of forests of high age and quality as well as increasing patch dynamics are most harmful to sexually dispersed species.

We record the distribution of (137)Cs, K, Rb and Cs within individual Sphagnum plants (down to 20cm depth) as well as (137)Cs in vascular plants growing on a peatland in eastern central Sweden. In Calluna vulgaris(137)Cs was mainly located within the green parts, whereas Andromeda polifolia, Eriophorum vaginatum and Vaccinium oxycoccos showed higher (137)Cs activity in roots. Carex rostrata and Menyanthes trifoliata showed variable distribution of (137)Cs within the plants. The patterns of (137)Cs activity concentration distribution as well as K, Rb and Cs concentrations within individual Sphagnum plants were rather similar and were usually highest in the capitula and/or in the subapical segments and lowest in the lower dead segments, which suggests continuous relocation of those elements to the actively growing apical part. The (137)Cs and K showed relatively weak correlations, especially in capitula and living green segments (0-10cm) of the plant (r=0.50). The strongest correlations were revealed between (137)Cs and Rb (r=0.89), and between (137)Cs and stable Cs (r=0.84). This suggests similarities between (137)Cs and Rb in uptake and relocation within the Sphagnum, but that (137)Cs differs from K.

Among wetlands, rich fens are one of the most species-rich, and also one of the most threatened habitats due to drainage activities. We tested the effects of several restoration measures on vegetation composition and dynamics over 4 years in two rich fens in east central Sweden that had been drained about 50 years ago. The encroaching tree layer was removed, and the restoration measures examined were rewetting by ditch blocking, mowing, and peat disturbance (removing vegetation and mixing the surface peat). At both sites, rewetting alone (without mowing or peat disturbance) led to small changes in species composition. In plots with peat disturbance, several characteristic rich fen species established, probably from a persistent diaspore bank or from refugia nearby. The disturbance treatment also prevented a rapid reestablishment of dominant grasses and mosses that had spread since drainage. Late season mowing alone appeared not to be a very efficient treatment in the initial phase after restoration, but seemed to reinforce the positive effects of peat disturbance. The results indicate the necessity of combining several treatments to recreate an ecologically functional vegetation.

Competitive exclusion among species with high niche overlap has been questioned by plant ecologists over the last decades. In this study, we used a greenhouse experiment to test the role of competition for community dynamics and composition among bryophytes in boreal rich (calcareous) fens. We evaluate the rate and direction of competition between three of the most common bryophytes in these wetlands, Campylium stellatum, Scorpidium cossonii and Scorpidium scorpioides. A pair-wise experimental setup with equal amounts of initial plant material of the species was used and the experiment continued for two years. To assess the competitive outcome we measured the area expansion of one species into the part of the experimental containers covered by its competitor, and the height growth of each species. Differences were observed between tested species for total spatial expansion into the area of its competitor. C. stellatum and S. cossonii showed similar total spatial expansion, while S. scorpioides showed very low spatial expansion. The spatial expansion also depended on the identity of the other species in the combination. All three pair-wise interactions were strongly asymmetric, and in qualitative terms a competitive hierarchy could be established. However, this did not lead to competitive exclusions, even though the experimental containers were small and the two years of continuous growth in the greenhouse would correspond to several years of growth under field conditions. The results suggest that in the natural habitat, where interactions are further weakened by microtopographic niche separation and interrupted by fine-scale disturbances, competitive exclusions are probably rare, if they occur at all.

The aim of this thesis was to increase knowledge about metapopulation and metacommunity processes in patchy, dynamic landscapes, using epiphytic bryophytes as a model system. Host trees and deciduous forest stands in the coniferous landscape are patchy, temporal and undergo changes in habitat quality during succession. Epiphytes must track this dynamic habitat network for their long-term survival. Community patterns at different spatial scales were explored and linked to regional metapopulation processes and local population dynamics. Spatial structuring in species richness both at a local and regional scale indicated stronger dispersal limitation but lower sensitivity to habitat quality in species with large asexual than in species with small sexual diaspores. In sexually dispersed species, a strong rescue effect was indicated by a bimodal frequency distribution of the species and by increasing local abundance with increasing patch connectivity. Present connectivity to other deciduous forest patches had positive effects on richness of asexually dispersed species, whereas richness of sexually dispersed species was instead related to the landscape connectivity 30 years ago. A study of local growth and reproduction suggested that this is caused by delayed sexual, but not asexual, reproduction. Habitat conditions affected the production of sporophytes, but not of asexual diaspores. No differences in either growth rates or competitive abilities among species with different dispersal and life-history strategies were found. In vitro experiments showed that establishment is higher from large asexual diaspores than from small sexual. Establishment of all diaspore types was limited by pH. There were indications of trade-offs between high germination and protonemal growth rates, desiccation tolerance and a rapid development of shoots from protonema. The results indicated that the epiphyte metacommunity is structured by two main trade-offs: dispersal distance (diaspore size) versus age at first reproduction, and dispersal distance versus sensitivity to habitat quality. Trade-offs in species traits may have evolved as a consequence of conflicting selection pressures imposed by habitat turnover, connectivity and irregular water supply rather than by species interactions. Syndromes of interrelated species traits imply that fairly small changes in habitat conditions can lead to distinct changes in metacommunity diversity: the results indicate that increasing distances among patches cause most harm to asexually dispersed species, whereas cuttings of forests of high age and quality as well as increasing patch dynamics are most harmful to sexually dispersed species.

Peatlands cover only 3% of the Earth's land surface but boreal and subarctic peatlands store about 15–30% of the world's soil carbon (C) as peat. Despite their potential for large positive feedbacks to the climate system through sequestration and emission of greenhouse gases, peatlands are not explicitly included in global climate models and therefore in predictions of future climate change. In April 2007 a symposium was held in Wageningen, the Netherlands, to advance our understanding of peatland C cycling. This paper synthesizes the main findings of the symposium, focusing on (i) small-scale processes, (ii) C fluxes at the landscape scale, and (iii) peatlands in the context of climate change. The main drivers controlling C fluxes are largely scale dependent and most are related to some aspects of hydrology. Despite high spatial and annual variability in Net Ecosystem Exchange (NEE), the differences in cumulative annual NEE are more a function of broad scale geographic location and physical setting than internal factors, suggesting the existence of strong feedbacks. In contrast, trace gas emissions seem mainly controlled by local factors. Key uncertainties remain concerning the existence of perturbation thresholds, the relative strengths of the CO2 and CH4 feedback, the links among peatland surface climate, hydrology, ecosystem structure and function, and trace gas biogeochemistry as well as the similarity of process rates across peatland types and climatic zones. Progress on these research areas can only be realized by stronger co-operation between disciplines that address different spatial and temporal scales.

We present data on the accumulation of carbon and nitrogen into an open oceanic ombrotrophic bog, SW Sweden, with high levels of anthropogenic nitrogen deposition. The aim was to investigate if this peatland currently acts as a sink for atmospheric carbon. Peat cores were sampled from the top peat layer in five different vegetation types. Small pines were used to date the cores. The cores bulk density and carbon and nitrogen content were determined. Avegetation-classified satellite image was used to estimate the areal extent of the vegetation types and to scale up these results to bog level. The rate of current carbon input into the upper oxic acrotelm was 290gm � 2 yr � 1 , and there were no significant differences in accumulation rates among the vegetation types. This organic matter input to the acrotelm was almost completely decomposed before it was deposited for storage in the deeper peat layers (the catotelm) and only a small fraction (� 1%) or 0.012gm � 2 yr � 1 of the carbon would be left, assuming a residence time of 100 years in the acrotelm. Nitrogen accumulation rates differed between the vegetation classes, and the average input via primary production varied from 5.33 to 16.8gm � 2 yr � 1 . Current nitrogen input rates into the catotelm are much lower, 0‐0.059gm � 2 yr � 1 , with the highest accumulation rates in lawn-dominated communities. We suggest that one of the main causes of the low carbon input rates is the high level of nitrogen deposition, which enhances decomposition and changes the vegetation from peat-forming Sphagnumdominance to dominance by dwarf shrubs and graminoids.

Biodiversity in wetland habitats is strongly affected by drainage. In Sweden, rich fens (alkaline fens) have been particularly affected by drainage campaigns for forestry and conversion to agricult ...

We examined the influence of tree- and stand-level conditions on lichen diversity on 143 ash trees, varying in age from 11 to 140+ y, in 5 deciduous stands in southern Sweden. The number of lichen species per tree varied from 2 to 30 and was primarily explained by tree trunk diameter and to a lesser extent by tree age, crown cover, lichen cover, and stand identity. The positive relationship between species richness and lichen cover seems compatible with a random placement of species and suggests that similar factors affect both lichen growth and establishment. Species richness did not increase on trees above 65 y of age, while species composition changed with tree age. Together with the positive linear effect of trunk diameter, these results suggest a slight overall positive effect of area, but that species richness over time depends more on species turnover. In addition, we examined if lichens occurring on trees of different ages differed in life history traits, e.g., spore size, thallus height,...

There is a growing interest in the effects of deciduous trees on biodiversity, soil processes and long-term productivity in boreal, conifer-dominated forests. This study investigated whether individual birch trees allowed to grow to maturity in the coniferous forest can have a local effect on floristic richness and regeneration of tree saplings. The ground vegetation was compared in 2 m radius plots around the stem under the canopies of matched conifer-deciduous trees in a mature, conifer-dominated forest, and included in the analysis variables that could potentially mediate the tree effect (soil pH, cover of lichens, bryophytes, leaf and needle litter). The field layer vegetation was more species rich under birch (Betula pendula and B. pubescens) than under conifers (Picea abies and Pinus sylvestris), and several vascular plant species (including saplings of tree species) occurred more often under birch than under conifers. However, when the effect of the number of subordinate trees was taken into account the difference between birch and pine was not significant. The number of tree regenerations (saplings) was lowest under pines, but did not differ between spruce and birch. There were no effects of the canopy species on soil pH or on cover of lichens and bryophytes. The difference in diversity may be caused by the different effects of leaf and needle litter, and it is also likely that canopy structure has an influence via interception and throughfall and by affecting the light and microclimate.

Aim To test whether species richness of Sphagnum mosses on islands in a land uplift archipelago is related to island age, area or connectivity, and whether the frequency of different species can be predicted by their life history and autecology. Location The northern Stockholm archipelago in the Baltic Sea, east-central Sweden, with a current land uplift rate of 4.4 mm year−1. Methods We sampled 17 islands differing in area (0.55–55 ha), height (3.6–18 m, representing c. 800–4000 years of age) and distance from mainland (1.6–41 km). For each Sphagnum patch we measured area, height above sea level, horizontal distance from the shore and shading from vascular plants. Factors affecting island species richness, species frequency and habitats on the islands were tested by stepwise regressions. Species frequency was tested on nine life history and autecological variables, including estimated abundance and spore output on the mainland, habitat preference and distribution. Results We recorded 500 patches of 19 Sphagnum species, distributed in 83 rock pools on 14 islands. Island species richness correlated positively with island area and with degree of shelter by surrounding islands, while distance from the mainland, connectivity, height or age did not add to the model. Species frequency (number of colonized islands and rock pools) was mainly predicted by spore output on the mainland and by habitat preference (swamp forest species were more frequent than others), while spore size, for example, did not add to the model. Species differed in mean height above and horizontal distance from the shore, area of occupied rock pools and in the degree of shading of patches. The mean horizontal distance from the shore and the area of occupied rock pools correlated positively with the normal growth position above the water table among species. Spore capsules were found in only 2% of patches, mostly in the bisexual Sphagnum fimbriatum. Main conclusions The presence of Sphagnum in the Stockholm archipelago seems to be governed by regional spore production and habitat demands. Sphagnum does not appear to be dispersal limited at distances up to 40 km and time spans of centuries. Species with a high regional spore output have had a higher colonization rate, which, together with the rarity of spore capsules on the islands, indicate the mainland as a source for colonization rather than dispersal among islands. Swamp forest species seem more tolerant to the island conditions (summer droughts and some salt spray) than open mire species. The different distances from the sea occupied by the species indicate a slow, continuous succession and species replacement towards the island interior as islands are being uplifted and thus expand in area. This partly explains why larger islands harbour more species. Our results thus support some of the island biogeographical theories related to the species–area relationship.