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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 similar 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.

Plants in flammable ecosystems have different response strategies to fire, such as increasing germination after exposure to smoke and break of dormancy through heat shock. Peatlands are ecosystems that are more likely to be disturbed by fire with increasing temperatures, but it is not clear how fire affects spore germination of Sphagnum, the dominant plants in peatlands. Here, we hypothesize that Sphagnum spores respond positively to single and combined treatments of moderate heat and smoke (by increased germinability), while spore germinability decreases in response to high temperature. We exposed the Sphagnum spores of four selected species (S. angustifolium, S. fuscum, S. magellanicum and S. squarrosum) collected from peatlands in the Changbai Mountains to heat (40, 60 and 100 degrees C), on its own and combined with smoke-water treatments. Our results showed that a heat of 100 degrees C inhibited the spore germination or even killed spores of all species, while spore germination of three (Sphagnum angustifolium, S. fuscum and S. squarrosum) of the four species was promoted by 40 and 60 degrees C heat compared to the control (20 degrees C). Hollow species (S. angustifolium and S. squarrosum) showed a stronger positive responsive to heat than hummock species (S. fuscum and S. magellanicum). Sphagnum fuscum spores responded positively to the combined heat and smoke treatment while the other species did not. For the first time, we demonstrate the positive effects of heat on its own and in combination with smoke on spore germination in wetland mosses, which may be important for the establishment and persistence of peatmoss populations after fire.

The relative importance of global versus local environmental factors for growth and thus carbon uptake of the bryophyte genusSphagnum-the main peat-former and ecosystem engineer in northern peatlands-remains unclear. We measured length growth and net primary production (NPP) of two abundantSphagnumspecies 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 onSphagnumgrowth. 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 denserSphagnum fuscumgrowing on hummocks had weaker responses to climatic variation than the larger and looserSphagnum magellanicumgrowing in the wetter conditions. Growth decreased with increasing vascular plant cover within a site. Between sites, precipitation and temperature increased growth forS. 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 influencedSphagnumgrowth indirectly by affecting moss shoot density. Synthesis. Our results imply that in a warmer climate,S. magellanicumwill increase length growth as long as precipitation is not reduced, whileS. fuscumis 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.

Spore production of bryophytes is common in subalpine ecosystems, but how spore banks of bryophytes vary with elevations is unknown. We used peat cores from twoSphagnumdominated peatlands, a low- (780 m a.s.l.) and a high-elevation (1480 m a.s.l.) site to test the hypotheses that spore concentration, accumulation rate, germinability and longevity are lower at a high-elevation peatland than those at lower elevation due to the assumed climatic limitations at high elevation. Along the cores,Sphagnumspores were extracted, burial time of spores was dated and spore germinability was tested layer by layer.Sphagnumspore concentration, accumulation rate and germinability were greater at the high-elevation peatland than at the low-elevation site. Both peatlands had long-term persistentSphagnumspore banks with a predicted longevity (the burial time when 1% spores are viable) ofc. 390 and 140 years for the low- and high-elevation sites, respectively. Our study suggests thatSphagnum, the ecological engineer of peatlands, by regulating spore production and longevity can maintain relatively constant spore banks which may be of great importance in population regeneration and persistence in a changing world.

Question: What environmental variables and plant–plant interactions affect mire bryophyte distribution and does the surrounding landscape with human disturbance play a role in the mire bryophyte distribution? Location: Jinchuan mire, Northeast China. Methods: We studied the spatial distribution of bryophytes in 100 1 × 1 m quadrats in the mire. Spatial variables were simulated by analysis of the distance-based Moran’s eigenvector maps (dbMEM). Variation partitioning analysis was used to reveal the relative contribution of spatial and environmental variables to bryophytes. The relationship between environmental variables and bryophytes was tested by redundancy analysis (RDA). We used co-occurrence and niche overlap models to detect interactions among bryophytes. We also studied the influence of the surrounding landscape on the distribution of bryophytes in relation to water chemistry. Results: The eight bryophytes occupying part of the mire had both a general distribution trend and a local spatial structure. Over 40% of the total variation in cover among bryophytes could be explained by spatial and environmental variables. In this fraction, the environmental variables explained 29.7% of the variation, of which only 4.5% was not spatially structured. RDA showed the contribution of dwarf shrub cover (SC), Na, and P to the bryophyte distribution was relatively large. Concentration of Na and SC decreased gradually from north to south, and contributed most to the variation in species composition along the first axis. The concentrations of P decreased from east to west, and contributed along the second axis. All the bryophyte species were spatially isolated but with large niche overlaps, indicating that the bryophyte community was structured by interspecific competition. Conclusion: Sodium mainly originating from the volcanic hill and P from the paddy fields were the main environmental factors affecting the bryophyte distribution. Concentrations of Na and P showed spatial

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, 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 S. 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 structural equation models indicated 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., 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, 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 S. 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 structural equation models indicated 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.