Wildfire in Swedish boreal forests - Impacts of fire severity and salvage logging on tree seedling regeneration and the belowground system

Wildfires are projected to occur more frequently and become more severe as the climate warms. The increasing release of carbon to the atmosphere from more severe fires may contribute to climate warming, and thus initiate a positive feedback loop. Following wildfire, salvage logging is a common management practice, but the impact on soil and vegetation may vary depending on fire severity. Hence, it is important to understand how wildfire severity and salvage logging affect soil abiotic and biotic properties and tree seedling regeneration. Also, in the future, forest recovery from fire and salvage logging may be affected by climate warming. In this thesis, I have conducted laboratory and greenhouse experiments to investigate the abiotic and biotic effects of fire severity and post-fire management on biomass production of Betula pendula, Picea abies and Pinus sylvestris seedlings regenerated from seeds and associated soil processes. Further, I have performed a mesocosm field experiment to investigate the impacts of fire severity, salvage logging and experimental warming on P. abies and P. sylvestris nursery seedlings in situ, and undertaken field measurements of soil N transformation rates and C fluxes in stands of varying fire severity and management. This thesis shows that the effect of fire severity on the soil system is stronger than that of salvage logging and that the impact of salvage logging on forest regeneration varies depending on fire severity; these effects can further be directly or indirectly altered by experimental climate warming. This thesis also shows that the effect of soil abiotic and biotic properties following fire on tree seedling regeneration are species specific. Altogether, it contributes with new fundamental knowledge about the interactive effects of multiple drivers of seedling regeneration and soil processes in the post-fire boreal forest landscape. The conclusions of this thesis can be applied, in combination with other research, in decision making about management of boreal forest following wildfire

Effects of Soil Abiotic and Biotic Factors on Tree Seedling Regeneration Following a Boreal Forest Wildfire

Wildfire disturbance is important for tree regeneration in boreal ecosystems. A considerable amount of literature has been published on how wildfires affect boreal forest regeneration. However, we lack understanding about how soil-mediated effects of fire disturbance on seedlings occur via soil abiotic properties versus soil biota. We collected soil from stands with three different severities of burning (high, low and unburned) and conducted two greenhouse experiments to explore how seedlings of tree species (Betula pendula, Pinus sylvestris and Picea abies) performed in live soils and in sterilized soil inoculated by live soil from each of the three burning severities. Seedlings grown in live soil grew best in unburned soil. When sterilized soils were reinoculated with live soil, seedlings of P. abies and P. sylvestris grew better in soil from low burn severity stands than soil from either high severity or unburned stands, demonstrating that fire disturbance may favor post-fire regeneration of conifers in part due to the presence of soil biota that persists when fire severity is low or recovers quickly post-fire. Betula pendula did not respond to soil biota and was instead driven by changes in abiotic soil properties following fire. Our study provides strong evidence that high fire severity creates soil conditions that are adverse for seedling regeneration, but that low burn severity promotes soil biota that stimulates growth and potential regeneration of conifers. It also shows that species-specific responses to abiotic and biotic soil characteristics are altered by variation in fire severity. This has important implications for tree regeneration because it points to the role of plant-soil-microbial feedbacks in promoting successful establishment, and potentially successional trajectories and species dominance in boreal forests in the future as fire regimes become increasingly severe through climate change

Mid-term Effects of Wildfire and Salvage Logging on Gross and Net Soil Nitrogen Transformation Rates in a Swedish Boreal Forest

Wildfires are natural and important disturbances of boreal forest ecosystems, and they are expected to increase in parts of the boreal zone through climate warming. There is a broad understanding of the immediate effects of fire on soil nitrogen (N) transformation rates, but less is known about these effects several years after fire. In July 2014, a large wildfire in the boreal forest zone of Central Sweden took place. Four years after the wildfire, we measured processes linked to the soil N cycle using the 15N pool dilution method (for gross N mineralization, consumption and nitrification) and the buried bags method (for net N mineralization), in soils from stands of different fire severity that had or had not been subjected to salvage logging. Gross N mineralization and consumption rates per unit carbon (C) increased by 81 % and 85 % respectively, in response to high fire severity, and nitrification rates per unit C basis decreased by 69 % in response to high fire severity, while net N mineralization was unresponsive. There was no difference in the effect of salvage logging across stands of differing fire severity on N transformation rates, although concentrations of resin adsorbed nitrate (NO3–) were overall 50 % lower in logged compared to unlogged stands. We also found that irrespective of burn severity, N immobilization rates exceeded N nitrification rates, and immobilization was therefore the dominant pathway of gross N consumption. Gross N consumption rates were higher in burned than unburned stands, despite there being a higher active microbial biomass in unburned soil, which suggests an even higher immobilization of N over time as the microbial biomass recovers following fire. Our study shows that soil N transformation rates were more affected by changes in fire severity than by salvage logging, and that four years after the fire many aspects of the N cycle did not differ between burned and unburned stands, suggesting substantial resilience of the N cycle to fire and salvage logging. However, we note that long term impact and many additional ecosystem properties or processes should be evaluated before concluding that salvage logging has no ecosystem impact. Furthermore, shortened fire regimes following climate warming accompanied with shorter intervals between salvage logging practices, could still impact the capability for the N cycle to recover after an intense fire. While wildfire in the boreal region results in a shift from nutrient conserving to nutrient demanding plant species, our results suggest this shift is dependent on a relatively short-lived pulse of higher N cycling processes that would have likely dissipated within a few years after the fire

The biological controls of soil carbon accumulation following wildfire and harvest in boreal forests: A review

Boreal forests are frequently subjected to disturbances, including wildfire and clear-cutting. While these disturbances can cause soil carbon (C) losses, the long-term accumulation dynamics of soil C stocks during subsequent stand development is controlled by biological processes related to the balance of net primary production (NPP) and outputs via heterotrophic respiration and leaching, many of which remain poorly understood. We review the biological processes suggested to influence soil C accumulation in boreal forests. Our review indicates that median C accumulation rates following wildfire and clear-cutting are similar (0.15 and 0.20 Mg ha(-1) year(-1), respectively), however, variation between studies is extremely high. Further, while many individual studies show linear increases in soil C stocks through time after disturbance, there are indications that C stock recovery is fastest early to mid-succession (e.g. 15-80 years) and then slows as forests mature (e.g. >100 years). We indicate that the rapid build-up of soil C in younger stands appears not only driven by higher plant production, but also by a high rate of mycorrhizal hyphal production, and mycorrhizal suppression of saprotrophs. As stands mature, the balance between reductions in plant and mycorrhizal production, increasing plant litter recalcitrance, and ectomycorrhizal decomposers and saprotrophs have been highlighted as key controls on soil C accumulation rates. While some of these controls appear well understood (e.g. temporal patterns in NPP, changes in aboveground litter quality), many others remain research frontiers. Notably, very little data exists describing and comparing successional patterns of root production, mycorrhizal functional traits, mycorrhizal-saprotroph interactions, or C outputs via heterotrophic respiration and dissolved organic C following different disturbances. We argue that these less frequently described controls require attention, as they will be key not only for understanding ecosystem C balances, but also for representing these dynamics more accurately in soil organic C and Earth system models

Fire severity as a key determinant of aboveground and belowground biological community recovery in managed even-aged boreal forests

Changes in fire regime of boreal forests in response to climate warming are expected to impact postfire recovery. However, quantitative data on how managed forests sustain and recover from recent fire disturbance are limited.Two years after a large wildfire in managed even-aged boreal forests in Sweden, we investigated how recovery of aboveground and belowground communities, that is, understory vegetation and soil microbial and faunal communities, responded to variation in the severity of soil (i.e., consumption of soil organic matter) and canopy fires (i.e., tree mortality).While fire overall enhanced diversity of understory vegetation through colonization of fire adapted plant species, it reduced the abundance and diversity of soil biota. We observed contrasting effects of tree- and soil-related fire severity on survival and recovery of understory vegetation and soil biological communities. Severe fires that killed overstory Pinus sylvestris promoted a successional stage dominated by the mosses Ceratodon purpureus and Polytrichum juniperinum, but reduced regeneration of tree seedlings and disfavored the ericaceous dwarf-shrub Vaccinium vitis-idaea and the grass Deschampsia flexuosa. Moreover, high tree mortality from fire reduced fungal biomass and changed fungal community composition, in particular that of ectomycorrhizal fungi, and reduced the fungivorous soil Oribatida. In contrast, soil-related fire severity had little impact on vegetation composition, fungal communities, and soil animals. Bacterial communities responded to both tree- and soil-related fire severity.Synthesis: Our results 2 years postfire suggest that a change in fire regime from a historically low-severity ground fire regime, with fires that mainly burns into the soil organic layer, to a stand-replacing fire regime with a high degree of tree mortality, as may be expected with climate change, is likely to impact the short-term recovery of stand structure and above- and belowground species composition of even-aged P. sylvestris boreal forests

Boreal forest soil carbon fluxes one year after a wildfire: Effects of burn severity and management

The extreme 2018 hot drought that affected central and northern Europe led to the worst wildfire season in Sweden in over a century. The Ljusdal fire complex, the largest area burnt that year (8995 ha), offered a rare opportunity to quantify the combined impacts of wildfire and post-fire management on Scandinavian boreal forests. We present chamber measurements of soil CO2 and CH4 fluxes, soil microclimate and nutrient content from five Pinus sylvestris sites for the first growing season after the fire. We analysed the effects of three factors on forest soils: burn severity, salvage-logging and stand age. None of these caused significant differences in soil CH4 uptake. Soil respiration, however, declined significantly after a high-severity fire (complete tree mortality) but not after a low-severity fire (no tree mortality), despite substantial losses of the organic layer. Tree root respiration is thus key in determining post-fire soil CO2 emissions and may benefit, along with heterotrophic respiration, from the nutrient pulse after a low-severity fire. Salvage-logging after a high-severity fire had no significant effects on soil carbon fluxes, microclimate or nutrient content compared with leaving the dead trees standing, although differences are expected to emerge in the long term. In contrast, the impact of stand age was substantial: a young burnt stand experienced more extreme microclimate, lower soil nutrient supply and significantly lower soil respiration than a mature burnt stand, due to a thinner organic layer and the decade-long effects of a previous clear-cut and soil scarification. Disturbance history and burn severity are, therefore, important factors for predicting changes in the boreal forest carbon sink after wildfires. The presented short-term effects and ongoing monitoring will provide essential information for sustainable management strategies in response to the increasing risk of wildfire

Two Novel DNAs that enhance symptoms and overcome CMD2 resistance to cassava mosaic disease

Cassava mosaic begomoviruses (CMBs) cause cassava mosaic disease (CMD) across Africa and the Indian subcontinent. Like all members of the geminivirus family, CMBs have small, circular single-stranded DNA genomes. We report here the discovery of two novel DNA sequences, designated SEGS-1 and SEGS-2 (for sequences enhancing geminivirus symptoms), that enhance symptoms and break resistance to CMD. The SEGS are characterized by GC-rich regions and the absence of long open reading frames. Both SEGS enhanced CMD symptoms in cassava (Manihot esculenta Crantz) when coinoculated with African cassava mosaic virus (ACMV), East African cassava mosaic Cameroon virus (EACMCV), or East African cassava mosaic virus-Uganda (EACMV-UG). SEGS-1 also overcame resistance of a cassava landrace carrying the CMD2 resistance locus when coinoculated with EACMV-UG. Episomal forms of both SEGS were detected in CMB-infected cassava but not in healthy cassava. SEGS-2 episomes were also found in virions and whiteflies. SEGS-1 has no homology to geminiviruses or their associated satellites, but the cassava genome contains a sequence that is 99% identical to full-length SEGS-1. The cassava genome also includes three sequences with 84 to 89% identity to SEGS-2 that together encompass all of SEGS-2 except for a 52-bp region, which includes the episomal junction and a 26-bp sequence related to alphasatellite replication origins. These results suggest that SEGS-1 is derived from the cassava genome and facilitates CMB infection as an integrated copy and/or an episome, while SEGS-2 was originally from the cassava genome but now is encapsidated into virions and transmitted as an episome by whiteflies.Cassava mosaic begomoviruses (CMBs) cause cassava mosaic disease (CMD) across Africa and the Indian subcontinent. Like all members of the geminivirus family, CMBs have small, circular single-stranded DNA genomes. We report here the discovery of two novel DNA sequences, designated SEGS-1 and SEGS-2 (for sequences enhancing geminivirus symptoms), that enhance symptoms and break resistance to CMD. The SEGS are characterized by GC-rich regions and the absence of long open reading frames. Both SEGS enhanced CMD symptoms in cassava (Manihot esculenta Crantz) when coinoculated with African cassava mosaic virus (ACMV), East African cassava mosaic Cameroon virus (EACMCV), or East African cassava mosaic virus-Uganda (EACMV-UG). SEGS-1 also overcame resistance of a cassava landrace carrying the CMD2 resistance locus when coinoculated with EACMV-UG. Episomal forms of both SEGS were detected in CMB-infected cassava but not in healthy cassava. SEGS-2 episomes were also found in virions and whiteflies. SEGS-1 has no homology to geminiviruses or their associated satellites, but the cassava genome contains a sequence that is 99% identical to full-length SEGS-1. The cassava genome also includes three sequences with 84 to 89% identity to SEGS-2 that together encompass all of SEGS-2 except for a 52-bp region, which includes the episomal junction and a 26-bp sequence related to alphasatellite replication origins. These results suggest that SEGS-1 is derived from the cassava genome and facilitates CMB infection as an integrated copy and/or an episome, while SEGS-2 was originally from the cassava genome but now is encapsidated into virions and transmitted as an episome by whiteflies

Fire severity as a key determinant of aboveground and belowground biological community recovery in managed even‐aged boreal forests

Abstract Changes in fire regime of boreal forests in response to climate warming are expected to impact postfire recovery. However, quantitative data on how managed forests sustain and recover from recent fire disturbance are limited. Two years after a large wildfire in managed even‐aged boreal forests in Sweden, we investigated how recovery of aboveground and belowground communities, that is, understory vegetation and soil microbial and faunal communities, responded to variation in the severity of soil (i.e., consumption of soil organic matter) and canopy fires (i.e., tree mortality). While fire overall enhanced diversity of understory vegetation through colonization of fire adapted plant species, it reduced the abundance and diversity of soil biota. We observed contrasting effects of tree‐ and soil‐related fire severity on survival and recovery of understory vegetation and soil biological communities. Severe fires that killed overstory Pinus sylvestris promoted a successional stage dominated by the mosses Ceratodon purpureus and Polytrichum juniperinum, but reduced regeneration of tree seedlings and disfavored the ericaceous dwarf‐shrub Vaccinium vitis‐idaea and the grass Deschampsia flexuosa. Moreover, high tree mortality from fire reduced fungal biomass and changed fungal community composition, in particular that of ectomycorrhizal fungi, and reduced the fungivorous soil Oribatida. In contrast, soil‐related fire severity had little impact on vegetation composition, fungal communities, and soil animals. Bacterial communities responded to both tree‐ and soil‐related fire severity. Synthesis: Our results 2 years postfire suggest that a change in fire regime from a historically low‐severity ground fire regime, with fires that mainly burns into the soil organic layer, to a stand‐replacing fire regime with a high degree of tree mortality, as may be expected with climate change, is likely to impact the short‐term recovery of stand structure and above‐ and belowground species composition of even‐aged P. sylvestris boreal forests

The biological controls of soil carbon accumulation following wildfire and harvest in boreal forests : a review

Boreal forests are frequently subjected to disturbances, including wildfire and clear-cutting. While these disturbances can cause soil carbon (C) losses, the long-term accumulation dynamics of soil C stocks during subsequent stand development is controlled by biological processes related to the balance of net primary production (NPP) and outputs via heterotrophic respiration and leaching, many of which remain poorly understood. We review the biological processes suggested to influence soil C accumulation in boreal forests. Our review indicates that median C accumulation rates following wildfire and clear-cutting are similar (0.15 and 0.20 Mg ha−1 year−1, respectively), however, variation between studies is extremely high. Further, while many individual studies show linear increases in soil C stocks through time after disturbance, there are indications that C stock recovery is fastest early to mid-succession (e.g. 15–80 years) and then slows as forests mature (e.g. >100 years). We indicate that the rapid build-up of soil C in younger stands appears not only driven by higher plant production, but also by a high rate of mycorrhizal hyphal production, and mycorrhizal suppression of saprotrophs. As stands mature, the balance between reductions in plant and mycorrhizal production, increasing plant litter recalcitrance, and ectomycorrhizal decomposers and saprotrophs have been highlighted as key controls on soil C accumulation rates. While some of these controls appear well understood (e.g. temporal patterns in NPP, changes in aboveground litter quality), many others remain research frontiers. Notably, very little data exists describing and comparing successional patterns of root production, mycorrhizal functional traits, mycorrhizal-saprotroph interactions, or C outputs via heterotrophic respiration and dissolved organic C following different disturbances. We argue that these less frequently described controls require attention, as they will be key not only for understanding ecosystem C balances, but also for representing these dynamics more accurately in soil organic C and Earth system models