Short- to medium-term effects of crown and surface fires on soil respiration in a Canadian boreal forest

Fires are an important perturbation for the carbon (C) dynamics of boreal forests, especially when they are stand-replacing. In North American boreal forests, crown fires are predominant and, therefore, the most studied. However, surface fires can also lead to major tree mortality with substantial implications for the C balance. Here, we assess the short- (hours – days) to medium-term (1 – 3 years) effects of the different fire types (surface vs. crown) on the postfire soil C effluxes in jack pine and black spruce forest stands in the Northwest Territories, Canada. We found that while trees were instantly killed by the four crown fires studied, trees also died within one year after two of three surface fires studied. Associated with this tree mortality, soil autotrophic respiration decreased after both fire types, although at different timings. The soil heterotrophic respiration was either lower or unchanged when measured 1 – 3 years after either fire type, but was increased when measured immediately after a surface fire, possibly due to the interaction between ash generation and wetting performed to suppress the fire. Our results suggest that both fire types can thus substantially alter C fluxes in the short- to medium-term, both through changes in vegetation and the soil environment.

Mitigating post-fire regeneration failure in boreal landscapes with reforestation and variable retention harvesting: At what cost?

Successive disturbances such as fire can affect post-disturbance regeneration density, with documented adverse effects on subsequent stand productivity. We conducted a simulation study to assess the potential of reactive (reforestation) and proactive (variable retention harvesting) post-fire regeneration failure mitigation strategies in a 1.37-Mha fire-prone boreal landscape dominated by black spruce and jack pine. We quantified their respective capacity to maintain landscape productivity and post-fire resilience, as well as their associated financial returns under current and projected (RCP 8.5) fire regimes. While post-fire reforestation with jack pine revealed to be the most effective strategy to maintain potential production, associated costs quickly became prohibitive when applied over extensive areas. Proactive strategies such as an extensive use of variable retention harvesting, combined with replanting of fire-adapted jack pine only in easily accessible areas, appeared as a more promising approach. Despite this, our results suggest an inevitable erosion of forest productivity due to post-fire regeneration failure events, highlighting the importance to integrate fire a priori in strategic forest management planning as well as its effects on long-term regeneration dynamics.

Variation of Stem Radius in Response to Defoliation in Boreal Conifers

In the long term, defoliation strongly decreases tree growth and survival. Insect outbreaks are a typical cause of severe defoliation. Eastern spruce budworm (Choristoneura fumiferana Clem.) outbreaks are one of the most significant disturbances of Picea and Abies boreal forests. Nevertheless, in boreal conifers, a 2-year defoliation has been shown to quickly improve tree water status, protect the foliage and decrease growth loss. It suggests that defoliation effects are time-dependent and could switch from favorable in the short term to unfavorable when defoliation duration exceeds 5–10 years. A better understanding of the effect of defoliation on stem radius variation during the needle flushing time-window could help to elucidate the relationships between water use and tree growth during an outbreak in the medium term. This study aims to assess the effects of eastern spruce budworm (Choristoneura fumiferana Clem.) defoliation and bud phenology on stem radius variation in black spruce [Picea mariana (Mill.) B.S.P.] and balsam fir [Abies balsamea (L.) Mill.] in a natural stand in Quebec, Canada. We monitored host and insect phenology, new shoot defoliation, seasonal stem radius variation and daytime radius phases (contraction and expansion) from 2016 to 2019. We found that defoliation significantly increased stem growth at the beginning of needle flushing. Needles flushing influenced the amplitude and duration of daily stem expansion and contraction, except the amplitude of stem contraction. Over the whole growing season, defoliation increased the duration of stem contraction, which in turn decreased the duration of stem expansion. However, the change (increase/decrease) of the duration of contraction/expansion reflects a reduced ability of the potential recovery from defoliation. Black spruce showed significantly larger 24-h cycles of stem amplitude compared to balsam fir. However, both species showed similar physiological adjustments during mild stress, preventing water loss from stem storage zones to support the remaining needles’ transpiration. Finally, conifers react to defoliation during a 4-year period, modulating stem radius variation phases according to the severity of the defoliation.

Fine Root Growth of Black Spruce Trees and Understory Plants in a Permafrost Forest Along a North-Facing Slope in Interior Alaska

Permafrost forests play an important role in the global carbon budget due to the huge amounts of carbon stored below ground in these ecosystems. Although fine roots are considered to be a major pathway of belowground carbon flux, separate contributions of overstory trees and understory shrubs to fine root dynamics in these forests have not been specifically characterized in relation to permafrost conditions, such as active layer thickness. In this study, we investigated fine root growth and morphology of trees and understory shrubs using ingrowth cores with two types of moss substrates (feather- and Sphagnum mosses) in permafrost black spruce (Picea mariana) stands along a north-facing slope in Interior Alaska, where active layer thickness varied substantially. Aboveground biomass, litterfall production rate, and fine root mass were also examined. Results showed that aboveground biomass, fine root mass, and fine root growth of black spruce trees tended to decrease downslope, whereas those of understory Ericaceae shrubs increased. Belowground allocation (e.g., ratio of fine root growth/leaf litter production) increased downslope in both of black spruce and understory plants. These results suggested that, at a lower slope, belowground resource availability was lower than at upper slope, but higher light availability under open canopy seemed to benefit the growth of the understory shrubs. On the other hand, understory shrubs were more responsive to the moss substrates than black spruce, in which Sphagnum moss substrates increased fine root growth of the shrubs as compared with feather moss substrates, whereas the effect was unclear for black spruce. This is probably due to higher moisture contents in Sphagnum moss substrates, which benefited the growth of small diameter (high specific root length) fine roots of understory shrubs. Hence, the contribution of understory shrubs to fine root growth was greater at lower slope than at upper slope, or in Sphagnum than in feather-moss substrates in our study site. Taken together, our data show that fine roots of Ericaceae shrubs are a key component in belowground carbon flux at permafrost black spruce forests with shallow active layer and/or with Sphagnum dominated forest floor.

Increasing fire and the decline of fire adapted black spruce in the boreal forest

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.

Differences in C, N, δ13C and δ15N among plant functional types after a wildfire in a black spruce forest, interior Alaska

We measured differences in %C, %N,  13C and  15N of plant functional types 17 (PFTs) between burned and unburned ground surfaces soon after a wildfire on a north-18 facing slope in interior Alaska. The C and N were measured for 16 species and 19 Sphagnum litter.  13C differed among the PFTs and was low for trees and shrubs, 20 suggesting that woody stems slowed C dynamics or showed low water use efficiency. 21  15N concentrations suggested that the herbaceous plants depended less on the 22 mycorrhizal associations that became weak on the burned surfaces. The shrub leaves 23 showed the lowest  15N of PFTs and showed higher  15N on the burned surface, showing 24 that N transfer from the soils to the leaves in the shrubs was slowed by the wildfire. 25 Mosses showed the highest C/N ratio. Sphagnum litter decomposed faster on the burned 26 surface, and %N and  15N in the litter increased from the second to third year on both 27 burned and unburned surfaces, while %C changed little. In conclusion, the responses to 28 the wildfire differed among the PFTs as characterized by their C and N dynamics. 29 30 Key words: Burned and unburned ground surface, carbon (C) and nitrogen (N), Alaskan 31 taiga, plant functional type, stable isotope

Response of Northern Populations of Black Spruce and Jack Pine to Southward Seed Transfers: Implications for Climate Change

A variety of responses to climate change have been reported for northern tree populations, primarily from tree-ring and satellite-based studies. Here we employ provenance data to examine growth and survival responses of northern populations (defined here as those occurring north of 52° N) of black spruce (Picea mariana) and jack pine (Pinus banksiana) to southward seed transfers. This space for time substitution affords insights into potential climate change responses by these important northern tree species. Based on previous work, we anticipated relatively flat response curves that peak at much warmer temperatures than those found at seed source origin. These expectations were generally met for growth-related responses, with peak growth associated with seed transfers to environments with mean annual temperatures 2.2 and 3.6 °C warmer than seed source origin for black spruce and jack pine, respectively. These findings imply that northern tree populations harbor a significant amount of resilience to climate warming. However, survival responses told a different story, with both species exhibiting reduced survival rates when moved to warmer and drier environments. Together with the growth-based results, these findings suggest that the warmer and drier conditions expected across much of northern Canada under climate change may reduce survival, but surviving trees may grow at a faster rate up until a certain magnitude of climate warming has been reached. We note that all relationships had high levels of unexplained variation, underlining the many factors that may influence provenance study outcomes and the challenges in predicting tree responses to climate change. Despite certain limitations, we feel that the provenance data employed here provide valuable insights into potential climate change outcomes for northern tree populations.