Abstract Key message Black spruce ( Picea mariana (Mill.) B.S.P.) has historically self-replaced following wildfire, but recent evidence suggests that this is changing. One factor could be negative impacts of intensifying fire activity on black spruce seed rain. We investigated this by measuring black spruce seed rain and seedling establishment. Our results suggest that increases in fire activity could reduce seed rain meaning reductions in black spruce establishment. Context Black spruce is an important conifer in boreal North America that develops a semi-serotinous, aerial seedbank and releases a pulse of seeds after fire. Variation in postfire seed rain has important consequences for black spruce regeneration and stand composition. Aims We explore the possible effects of changes in fire regime on the abundance and viability of black spruce seeds following a very large wildfire season in the Northwest Territories, Canada (NWT). Methods We measured postfire seed rain over 2 years at 25 black spruce-dominated sites and evaluated drivers of stand characteristics and environmental conditions on total black spruce seed rain and viability. Results We found a positive relationship between black spruce basal area and total seed rain. However, at high basal areas, this increasing rate of seed rain was not maintained. Viable seed rain was greater in stands that were older, closer to unburned edges, and where canopy combustion was less severe. Finally, we demonstrated positive relationships between seed rain and seedling establishment, confirming our measures of seed rain were key drivers of postfire forest regeneration. Conclusion These results indicate that projected increases in fire activity will reduce levels of black spruce recruitment following fire.

Resilience of plant communities to disturbance is supported by multiple mechanisms, including ecological legacies affecting propagule availability, species' environmental tolerances, and biotic interactions. Understanding the relative importance of these mechanisms for plant community resilience supports predictions of where and how resilience will be altered with disturbance. We tested mechanisms underlying resilience of forests dominated by black spruce (Picea mariana) to fire disturbance across a heterogeneous forest landscape in the Northwest Territories, Canada. We combined surveys of naturally regenerating seedlings at 219 burned plots with experimental manipulations of ecological legacies via seed addition of four tree species and vertebrate exclosures to limit granivory and herbivory at 30 plots varying in moisture and fire severity. Black spruce recovery was greatest where it dominated pre-fire, at wet sites with deep residual soil organic layers, and fire conditions of low soil or canopy combustion and longer return intervals. Experimental addition of seed indicated all species were seed-limited, emphasizing the importance of propagule legacies. Black spruce and birch (Betula papyrifera) recruitment were enhanced with vertebrate exclusion. Our combination of observational and experimental studies demonstrates black spruce is vulnerable to effects of increased fire activity that erode ecological legacies. Moreover, black spruce relies on wet areas with deep soil organic layers where other species are less competitive. However, other species can colonize these areas if enough seed is available or soil moisture is altered by climate change. Testing mechanisms underlying species' resilience to disturbance aids predictions of where vegetation will transform with effects of climate change.The online version contains supplementary material available at 10.1007/s10021-022-00772-7.

Intensifying wildfire activity and climate change can drive rapid forest compositional shifts. In boreal North America, black spruce shapes forest flammability and depends on fire for regeneration. This relationship has helped black spruce maintain its dominance through much of the Holocene. However, with climate change and more frequent and severe fires, shifts away from black spruce dominance to broadleaf or pine species are emerging, with implications for ecosystem functions including carbon sequestration, water and energy fluxes, and wildlife habitat. Here, we predict that such reductions in black spruce after fire may already be widespread given current trends in climate and fire. To test this, we synthesize data from 1,538 field sites across boreal North America to evaluate compositional changes in tree species following 58 recent fires (1989 to 2014). While black spruce was resilient following most fires (62%), loss of resilience was common, and spruce regeneration failed completely in 18% of 1,140 black spruce sites. In contrast, postfire regeneration never failed in forests dominated by jack pine, which also possesses an aerial seed bank, or broad-leaved trees. More complete combustion of the soil organic layer, which often occurs in better-drained landscape positions and in dryer duff, promoted compositional changes throughout boreal North America. Forests in western North America, however, were more vulnerable to change due to greater long-term climate moisture deficits. While we find considerable remaining resilience in black spruce forests, predicted increases in climate moisture deficits and fire activity will erode this resilience, pushing the system toward a tipping point that has not been crossed in several thousand years.

Fungi play key roles in carbon (C) dynamics of ecosystems: saprotrophs decompose organic material and return C in the nutrient cycle, and mycorrhizal species support plants that accumulate C through photosynthesis. The identities and functions of extremophile fungi present after fire can influence C dynamics, particularly because plant-fungal relationships are often species-specific. However, little is known about the function and distribution of fungi that survive fires. We aim to assess the distribution of heat-resistant soil fungi across burned stands of boreal forest in the Northwest Territories, Canada, and understand their functions in relation to decomposition and tree seedling growth. We cultured and identified fungi from heat-treated soils and linked sequences from known taxa with high throughput sequencing fungal data (Illumina MiSeq, ITS1) from soils collected in 47 plots. We assessed functions under controlled conditions by inoculating litter and seedlings with heat-resistant fungi to assess decomposition and effects on seedling growth, respectively, for black spruce ( Picea mariana ), birch ( Betula papyrifera ), and jack pine ( Pinus banksiana ). We also measured litter decomposition rates and seedling densities in the field without inoculation. We isolated seven taxa of heat-resistant fungi and found their relative abundances were not associated with environmental or fire characteristics. Under controlled conditions, Fayodia gracilipes and Penicillium arenicola decomposed birch, but no taxa decomposed black spruce litter significantly more than the control treatment. Seedlings showed reduced biomass and/or mortality when inoculated with at least one of the fungal taxa. Penicillium turbatum reduced growth and/or caused mortality of all three species of seedlings. In the field, birch litter decomposed faster in stands with greater pre-fire proportion of black spruce, while black spruce litter decomposed faster in stands experiencing longer fire-free intervals. Densities of seedlings that had germinated since fire were positively associated with ectomycorrhizal richness while there were fewer conifer seedlings with greater heat-resistant fungal abundance. Overall, our study suggests that extremophile fungi present after fires have multiple functions and may have unexpected negative effects on forest functioning and regeneration. In particular, heat-resistant fungi after fires may promote shifts away from conifer dominance that are observed in these boreal forests.