Contributions of insects and droughts to growth decline of trembling aspen mixed boreal forest of western Canada

Insects, diseases, fire and drought and other disturbances associated with global climate change contribute to forest decline and mortality in many parts of the world. Forest decline and mortality related to drought or insect outbreaks have been observed in North American aspen forests. However, little research has been done to partition and estimate their relative contributions to growth declines. In this study, we combined tree‐ring width and basal area increment series from 40 trembling aspen (Populus tremuloides Michx.) sites along a latitudinal gradient (from 52° to 58°N) in western Canada and attempted to investigate the effect of drought and insect outbreaks on growth decline, and simultaneously partition and quantify their relative contributions. Results indicated that the influence of drought on forest decline was stronger than insect outbreaks, although both had significant effects. Furthermore, the influence of drought and insect outbreaks showed spatiotemporal variability. In addition, our data suggest that insect outbreaks could be triggered by warmer early spring temperature instead of drought, implicating that potentially increased insect outbreaks are expected with continued warming springs, which may further exacerbate growth decline and death in North America aspen mixed forests.     

Net aboveground biomass declines of four major forest types with forest ageing and climate change in western Canada's boreal forests

Biomass change of the world's forests is critical to the global carbon cycle. Despite storing nearly half of global forest carbon, the boreal biome of diverse forest types and ages is a poorly understood component of the carbon cycle. Using data from 871 permanent plots in the western boreal forest of Canada, we examined net annual aboveground biomass change (ΔAGB) of four major forest types between 1958 and 2011. We found that ΔAGB was higher for deciduous broadleaf (DEC) (1.44 Mg ha^?1 year^?1, 95% Bayesian confidence interval (CI), 1.22–1.68) and early‐successional coniferous forests (ESC) (1.42, CI, 1.30–1.56) than mixed forests (MIX) (0.80, CI, 0.50–1.11) and late‐successional coniferous (LSC) forests (0.62, CI, 0.39–0.88). ΔAGB declined with forest age as well as calendar year. After accounting for the effects of forest age, ΔAGB declined by 0.035, 0.021, 0.032 and 0.069 Mg ha^?1 year^?1 per calendar year in DEC, ESC, MIX and LSC forests, respectively. The ΔAGB declines resulted from increased tree mortality and reduced growth in all forest types except DEC, in which a large biomass loss from mortality was accompanied with a small increase in growth. With every degree of annual temperature increase, ΔAGB decreased by 1.00, 0.20, 0.55 and 1.07 Mg ha^?1 year^?1 in DEC, ESC, MIX and LSC forests, respectively. With every cm decrease of annual climatic moisture availability, ΔAGB decreased 0.030, 0.045 and 0.17 Mg ha^?1 year^?1 in ESC, MIX and LSC forests, but changed little in DEC forests. Our results suggest that persistent warming and decreasing water availability have profound negative effects on forest biomass in the boreal forests of western Canada. Furthermore, our results indicate that forest responses to climate change are strongly dependent on forest composition with late‐successional coniferous forests being most vulnerable to climate changes in terms of aboveground biomass.     

Species-specific flowering phenology responses to experimental warming and drought alter herbaceous plant species overlap in a temperate–boreal forest community

Background and AimsWarmer temperatures and altered precipitation patterns are expected to continue to occur as the climate changes. How these changes will impact the flowering phenology of herbaceous perennials in northern forests is poorly understood but could have consequences for forest functioning and species interactions. Here, we examine the flowering phenology responses of five herbaceous perennials to experimental warming and reduced summer rainfall over 3 years.MethodsThis study is part of the B4WarmED experiment located at two sites in northern Minnesota, USA. Three levels of warming (ambient, +1.6 °C and +3.1 °C) were crossed with two rainfall manipulations (ambient and 27 % reduced growing season rainfall).Key ResultsWe observed species-specific responses to the experimental treatments. Warming alone advanced flowering for four species. Most notably, the two autumn blooming species showed the strongest advance of flowering to warming. Reduced rainfall alone advanced flowering for one autumn blooming species and delayed flowering for the other, with no significant impact on the three early blooming species. Only one species, Solidago spp., showed an interactive response to warming and rainfall manipulation by advancing in +1.6 °C warming (regardless of rainfall manipulation) but not advancing in the warmest, driest treatment. Species-specific responses led to changes in temporal overlap between species. Most notably, the two autumn blooming species diverged significantly in their flowering timing. In ambient conditions, these two species flowered within the same week. In the warmest, driest treatment, flowering occurred over a month apart.ConclusionsHerbaceous species may differ in how they respond to future climate conditions. Changes to phenology may lead to fewer resources for insects or a mismatch between plants and pollinators.     

Spatial and temporal variations in boreal forest fire frequency in northern Alberta

Abstract. Spatial and temporal variations in fire frequency in the boreal forest of Wood Buffalo National Park (WBNP) were assessed using forest stand age, fire scar and historical data. I test the hypotheses that (1) fire frequency is higher in jack pine forests and aspen forests than in black spruce forests and white spruce forests, (2) these variations in fire frequency can be related to the mean waterbreak distance (MWD) around a site and (3) fire frequency has changed over the past 300 years. The fire cycles (the time required to burn an area equal in size to the entire study area) in jack pine forests (39 years) and in aspen forests (39 years) were significantly shorter than those in black spruce forests (78 years) and in white spruce forests (96 years). The length of the fire cycle varies inversely with the MWD around a site, and the MWD was significantly higher in jack pine and aspen forests than in black or white spruce forests. It is suggested that covariations between soil type and the MWD influence, respectively, variations in forest dominant and fire frequency. A change in fire frequency at 1860 was apparent in the fire history for all of WBNP, the black spruce dominated stands, and the near and medium MWD classes. The fire cycle estimates for these classes were all significantly shorter during the period 1750 to 1859 (fire cycles = 25–49 years) than they were in the period 1860 to 1989 (fire cycles = 59–89 years). The possible roles of changes in climate and aboriginal burning practices in causing the temporal change in fire frequency are discussed.     

Spatially explicit fire-climate history of the boreal forest-tundra (Eastern Canada) over the last 2000 years

Across the boreal forest, fire is the main disturbance factor and driver of ecosystem changes. In this study, we reconstructed a long-term, spatially explicit fire history of a forest-tundra region in northeastern Canada. We hypothesized that current occupation of similar topographic and edaphic sites by tundra and forest was the consequence of cumulative regression with time of forest cover due to compounding fire and climate disturbances. All fires were mapped and dated per 100 year intervals over the last 2,000 years using several fire dating techniques. Past fire occurrences and post-fire regeneration at the northern forest limit indicate 70% reduction of forest cover since 1800 yr BP and nearly complete cessation of forest regeneration since 900 yr BP. Regression of forest cover was particularly important between 1500s-1700s and possibly since 900 yr BP. Although fire frequency was very low over the last 100 years, each fire event was followed by drastic removal of spruce cover. Contrary to widespread belief of northward boreal forest expansion due to recent warming, lack of post-fire recovery during the last centuries, in comparison with active tree regeneration more than 1,000 years ago, indicates that the current climate does not favour such expansion.     

Disturbance legacies and climate jointly drive tree growth and mortality in an intensively studied boreal forest

Most North American forests are at some stage of post‐disturbance regrowth, subject to a changing climate, and exhibit growth and mortality patterns that may not be closely coupled to annual environmental conditions. Distinguishing the possibly interacting effects of these processes is necessary to put short‐term studies in a longer term context, and particularly important for the carbon‐dense, fire‐prone boreal forest. The goals of this study were to combine dendrochronological sampling, inventory records, and machine‐learning algorithms to understand how tree growth and death have changed at one highly studied site (Northern Old Black Spruce, NOBS) in the central Canadian boreal forest. Over the 1999–2012 inventory period, mean tree diameter increased even as stand density and basal area declined significantly. Tree mortality averaged 1.4 ± 0.6% yr^?1, with most mortality occurring in medium‐sized trees; new recruitment was minimal. There have been at least two, and probably three, significant influxes of new trees since stand initiation, but none in recent decades. A combined tree ring chronology constructed from sampling in 2001, 2004, and 2012 showed several periods of extreme growth depression, with increased mortality lagging depressed growth by ~5 years. Higher minimum and maximum air temperatures exerted a negative influence on tree growth, while precipitation and climate moisture index had a positive effect; both current‐ and previous‐year data exerted significant effects. Models based on these variables explained 23–44% of the ring‐width variability. We suggest that past climate extremes led to significant mortality still visible in the current forest structure, with decadal dynamics superimposed on slower patterns of fire and succession. These results have significant implications for our understanding of previous work at NOBS, the carbon sequestration capability of old‐growth stands in a disturbance‐prone landscape, and the sustainable management of regional forests in a changing climate.     

Effects of forest fire and drought on acidity of a base-poor boreal forest stream: similarities between climatic warming and acidic precipitation

In a boreal forest catchment in the Experimental Lakes Area in northwestern Ontario, wildfire caused an increase in the concentrations of strong acid anions and base cations of the stream. In the naturally base-poor Northwest (NW) Subbasin, a 1980 wildfire caused exports of strong acid anions to increase more than export of base cations, causing a 2.5 fold increase in the acidity of the stream. Mean annual stream pH declined from 5.15 prior to fire to 4.76 two years after fire. Acid-neutralizing capacity (ANC), calculated as the difference between total base cations and strong acid anions, decreased to 20% of pre-fire values. Sulfate and chloride were the strong acid anions responsible for the decline in ANC, increasing four-fold. While nitrate increased eleven-fold, concentrations were too low to significantly affect ANC. There was a significant correlation between weekly sulfate concentration and base cation concentration (r ² = 0.83) in the two years after fire. Recovery of ANC was caused by the more rapid decline in concentration of sulfate than by changes in base cations. Drought produced a similar but weaker response than fire, with increased sulfate concentrations and decreased stream pH. Climatic warming that increases drought and fire frequency would have effects that mimic the impacts of acidic precipitation (i.e. higher sulfate concentrations and acidic stream waters). Areas which have higher concentrations of stored S from past acid precipitation or have large areas of peatlands in the watershed may have aggravated losses of S and H⁺ after drought and fire.     

Impact of fire on active layer and permafrost microbial communities and metagenomes in an upland Alaskan boreal forest

Permafrost soils are large reservoirs of potentially labile carbon (C). Understanding the dynamics of C release from these soils requires us to account for the impact of wildfires, which are increasing in frequency as the climate changes. Boreal wildfires contribute to global emission of greenhouse gases (GHG—CO₂, CH₄ and N₂O) and indirectly result in the thawing of near-surface permafrost. In this study, we aimed to define the impact of fire on soil microbial communities and metabolic potential for GHG fluxes in samples collected up to 1 m depth from an upland black spruce forest near Nome Creek, Alaska. We measured geochemistry, GHG fluxes, potential soil enzyme activities and microbial community structure via 16SrRNA gene and metagenome sequencing. We found that soil moisture, C content and the potential for respiration were reduced by fire, as were microbial community diversity and metabolic potential. There were shifts in dominance of several microbial community members, including a higher abundance of candidate phylum AD3 after fire. The metagenome data showed that fire had a pervasive impact on genes involved in carbohydrate metabolism, methanogenesis and the nitrogen cycle. Although fire resulted in an immediate release of CO₂ from surface soils, our results suggest that the potential for emission of GHG was ultimately reduced at all soil depths over the longer term. Because of the size of the permafrost C reservoir, these results are crucial for understanding whether fire produces a positive or negative feedback loop contributing to the global C cycle.     

Drivers of postfire soil organic carbon accumulation in the boreal forest

The accumulation of soil carbon (C) is regulated by a complex interplay between abiotic and biotic factors. Our study aimed to identify the main drivers of soil C accumulation in the boreal forest of eastern North America. Ecosystem C pools were measured in 72 sites of fire origin that burned 2–314 years ago over a vast region with a range of ? mean annual temperature of 3°C and one of ? 500 mm total precipitation. We used a set of multivariate a priori causal hypotheses to test the influence of time since fire (TSF), climate, soil physico‐chemistry and bryophyte dominance on forest soil organic C accumulation. Integrating the direct and indirect effects among abiotic and biotic variables explained as much as 50% of the full model variability. The main direct drivers of soil C stocks were: TSF >bryophyte dominance of the FH layer and metal oxide content >pH of the mineral soil. Only climate parameters related to water availability contributed significantly to explaining soil C stock variation. Importantly, climate was found to affect FH layer and mineral soil C stocks indirectly through its effects on bryophyte dominance and organo‐metal complexation, respectively. Soil texture had no influence on soil C stocks. Soil C stocks increased both in the FH layer and mineral soil with TSF and this effect was linked to a decrease in pH with TSF in mineral soil. TSF thus appears to be an important factor of soil development and of C sequestration in mineral soil through its influence on soil chemistry. Overall, this work highlights that integrating the complex interplay between the main drivers of soil C stocks into mechanistic models of C dynamics could improve our ability to assess C stocks and better anticipate the response of the boreal forest to global change.     

Mechanisms influencing changes in lake area in Alaskan boreal forest

During the past ～50 years, the number and area of lakes have declined in several regions in boreal forests. However, there has been substantial finer‐scale heterogeneity; some lakes decreased in area, some showed no trend, and others increased. The objective of this study was to identify the primary mechanisms underlying heterogeneous trends in closed‐basin lake area. Eight lake characteristics (δ¹⁸O, electrical conductivity, surface : volume index, bank slope, floating mat width, peat depth, thaw depth at shoreline, and thaw depth at the forest boundary) were compared for 15 lake pairs in Alaskan boreal forest where one lake had decreased in area since ～1950, and the other had not. Mean differences in characteristics between paired lakes were used to identify the most likely of nine mechanistic scenarios that combined three potential mechanisms for decreasing lake area (talik drainage, surface water evaporation, and terrestrialization) with three potential mechanisms for nondecreasing lake area (subpermafrost groundwater recharge through an open talik, stable permafrost, and thermokarst). A priori expectations of the direction of mean differences between decreasing and nondecreasing paired lakes were generated for each scenario. Decreasing lakes had significantly greater electrical conductivity, greater surface : volume indices, shallower bank slopes, wider floating mats, greater peat depths, and shallower thaw depths at the forest boundary. These results indicated that the most likely scenario was terrestrialization as the mechanism for lake area reduction combined with thermokarst as the mechanism for nondecreasing lake area. Terrestrialization and thermokarst may have been enhanced by recent warming which has both accelerated permafrost thawing and lengthened the growing season, thereby increasing plant growth, floating mat encroachment, transpiration rates, and the accumulation of organic matter in lake basins. The transition to peatlands associated with terrestrialization may provide a transient increase in carbon storage enhancing the role of northern ecosystems as major stores of global carbon.     

Spatial patterns and drivers of fire occurrence and its future trend under climate change in a boreal forest of Northeast China

Understanding the spatial patterns of fire occurrence and its response to climate change is vital to fire risk mitigation and vegetation management. Focusing on boreal forests in Northeast China, we used spatial point pattern analysis to model fire occurrence reported from 1965 to 2009. Our objectives were to quantitate the relative importance of biotic, abiotic, and human influences on patterns of fire occurrence and to map the spatial distribution of fire occurrence density (number of fires occurring over a given area and time period) under current and future climate conditions. Our results showed human‐caused fires were strongly related to human activities (e.g. landscape accessibility), including proximity to settlements and roads. In contrast, fuel moisture and vegetation type were the most important controlling factors on the spatial pattern of lightning fires. Both current and future projected spatial distributions of the overall (human‐ + lightning‐caused) fire occurrence density were strongly clustered along linear components of human infrastructure. Our results demonstrated that the predicted change in overall fire occurrence density is positively related to the degree of temperature and precipitation change, although the spatial pattern of change is expected to vary spatially according to proximity to human ignition sources, and in a manner inconsistent with predicted climate change. Compared to the current overall fire occurrence density (median value: 0.36 fires per 1000 km² per year), the overall fire occurrence density is projected to increase by 30% under the CGCM3 B1 scenario and by 230% under HadCM3 A2 scenario in 2081–2100, respectively. Our results suggest that climate change effects may not outweigh the effects of human influence on overall fire occurrence over the next century in this cultural landscape. Accurate forecasts of future fire‐climate relationships should account for anthropogenic influences on fire ignition density, such as roads and proximity to settlements.     

Regional atmospheric cooling and wetting effect of permafrost thaw‐induced boreal forest loss

In the sporadic permafrost zone of North America, thaw‐induced boreal forest loss is leading to permafrost‐free wetland expansion. These land cover changes alter landscape‐scale surface properties with potentially large, however, still unknown impacts on regional climates. In this study, we combine nested eddy covariance flux tower measurements with satellite remote sensing to characterize the impacts of boreal forest loss on albedo, eco‐physiological and aerodynamic surface properties, and turbulent energy fluxes of a lowland boreal forest region in the Northwest Territories, Canada. Planetary boundary layer modelling is used to estimate the potential forest loss impact on regional air temperature and atmospheric moisture. We show that thaw‐induced conversion of forests to wetlands increases albedo: and bulk surface conductance for water vapour and decreases aerodynamic surface temperature. At the same time, heat transfer efficiency is reduced. These shifts in land surface properties increase latent at the expense of sensible heat fluxes, thus, drastically reducing Bowen ratios. Due to the lower albedo of forests and their masking effect of highly reflective snow, available energy is lower in wetlands, especially in late winter. Modelling results demonstrate that a conversion of a present‐day boreal forest–wetland to a hypothetical homogeneous wetland landscape could induce a near‐surface cooling effect on regional air temperatures of up to 3–4 °C in late winter and 1–2 °C in summer. An atmospheric wetting effect in summer is indicated by a maximum increase in water vapour mixing ratios of 2 mmol mol^?1. At the same time, maximum boundary layer heights are reduced by about a third of the original height. In fall, simulated air temperature and atmospheric moisture between the two scenarios do not differ. Therefore, permafrost thaw‐induced boreal forest loss may modify regional precipitation patterns and slow down regional warming trends.     

Massive mortality of aspen following severe drought along the southern edge of the Canadian boreal forest

Drought‐induced, regional‐scale dieback of forests has emerged as a global concern that is expected to escalate under model projections of climate change. Since 2000, drought of unusual severity, extent, and duration has affected large areas of western North America, leading to regional‐scale dieback of forests in the southwestern US. We report on drought impacts on forests in a region farther north, encompassing the transition between boreal forest and prairie in western Canada. A central question is the significance of drought as an agent of large‐scale tree mortality and its potential future impact on carbon cycling in this cold region. We used a combination of plot‐based, meteorological, and remote sensing measures to map and quantify aboveground, dead biomass of trembling aspen (Populus tremuloides Michx.) across an 11.5 Mha survey area where drought was exceptionally severe during 2001–2002. Within this area, a satellite‐based land cover map showed that aspen‐dominated broadleaf forests occupied 2.3 Mha. Aerial surveys revealed extensive patches of severe mortality (>55%) resembling the impacts of fire. Dead aboveground biomass was estimated at 45 Mt, representing 20% of the total aboveground biomass, based on a spatial interpolation of plot‐based measurements. Spatial variation in percentage dead biomass showed a moderately strong correlation with drought severity. In the prairie‐like, southern half of the study area where the drought was most severe, 35% of aspen biomass was dead, compared with an estimated 7% dead biomass in the absence of drought. Drought led to an estimated 29 Mt increase in dead biomass across the survey area, corresponding to 14 Mt of potential future carbon emissions following decomposition. Many recent, comparable episodes of drought‐induced forest dieback have been reported from around the world, which points to an emerging need for multiscale monitoring approaches to quantify drought effects on woody biomass and carbon cycling across large areas.     

Insect‐induced tree mortality of boreal forests in eastern Canada under a changing climate

Forest insects are major disturbances that induce tree mortality in eastern coniferous (or fir-spruce) forests in eastern North America. The spruce budworm (SBW) (Choristoneura fumiferana [Clemens]) is the most devastating insect causing tree mortality. However, the relative importance of insect-caused mortality versus tree mortality caused by other agents and how this relationship will change with climate change is not known. Based on permanent sample plots across eastern Canada, we combined a logistic model with a negative model to estimate tree mortality. The results showed that tree mortality increased mainly due to forest insects. The mean difference in annual tree mortality between plots disturbed by insects and those without insect disturbance was 0.0680 per year (P < 0.0001, T-test), and the carbon sink loss was about 2.87t C ha⁻¹ year⁻¹ larger than in natural forests. We also found that annual tree mortality increased significantly with the annual climate moisture index (CMI) and decreased significantly with annual minimum temperature (T_min), annual mean temperature (T_mean) and the number of degree days below 0°C (DD0), which was inconsistent with previous studies (Adams et al. 2009; van Mantgem et al. 2009; Allen et al. 2010). Furthermore, the results for the trends in the magnitude of forest insect outbreaks were consistent with those of climate factors for annual tree mortality. Our results demonstrate that forest insects are the dominant cause of the tree mortality in eastern Canada but that tree mortality induced by insect outbreaks will decrease in eastern Canada under warming climate.     

Climate change effects on net carbon exchange of a boreal aspen–hazelnut forest: estimates from the ecosystem model ecosys

Although boreal forests are currently sinks for atmospheric C, there is some concern that they may not remain so under hypothesized warming of the boreal climate. The ecosystem model ecosys was used to evaluate possible changes in ecosystem C exchange and accumulation under changes in atmospheric CO₂ concentration (C_a) proposed in emissions scenario IS92a, and accompanying changes in air temperature and precipitation proposed by general circulation models running under IS92a. Ecosys was first tested under current climate by comparing modelled rates of C exchange and accumulation with those measured in a mixed aspen–hazelnut stand in central Saskatchewan. The model was then run with daily increments of C_a, temperature and precipitation, and differences in C exchange and accumulation between current and changing climates were evaluated. Model results indicated that over a 120‐y period, a mixed aspen–hazelnut stand currently accumulates about 14 kg C m^?2. Under the hypothesized changes in climate this stand would accumulate an additional 8.5 kg C m^?2, largely through higher rates of CO₂ fixation and longer growing seasons under higher C_a and temperature. This additional accumulation would be entirely as aspen wood, while soil organic matter would change little. This accumulation would therefore be vulnerable to losses from fire and insects.     

Differential declines in Alaskan boreal forest vitality related to climate and competition

Rapid warming and changes in water availability at high latitudes alter resource abundance, tree competition, and disturbance regimes. While these changes are expected to disrupt the functioning of boreal forests, their ultimate implications for forest composition are uncertain. In particular, recent site‐level studies of the Alaskan boreal forest have reported both increases and decreases in productivity over the past few decades. Here, we test the idea that variations in Alaskan forest growth and mortality rates are contingent on species composition. Using forest inventory measurements and climate data from plots located throughout interior and south‐central Alaska, we show significant growth and mortality responses associated with competition, midsummer vapor pressure deficit, and increased growing season length. The governing climate and competition processes differed substantially across species. Surprisingly, the most dramatic climate response occurred in the drought tolerant angiosperm species, trembling aspen, and linked high midsummer vapor pressure deficits to decreased growth and increased insect‐related mortality. Given that species composition in the Alaskan and western Canadian boreal forests is projected to shift toward early‐successional angiosperm species due to fire regime, these results underscore the potential for a reduction in boreal productivity stemming from increases in midsummer evaporative demand.     

Stand- and plot-level changes in a boreal forest understory community following wildfire

Background: Boreal forest understory plant communities are known to be resilient to fire – the species composition of stands after a fire is quite similar to the pre-fire composition. However, we know little about recovery of individual plants within particular locations in forest stands (i.e. plot-level changes) since we usually do not have pre-fire data for plots.

Aims: We wanted to determine whether species recruited into the same or different locations in a Pinus banksiana stand that experienced a severe wildfire.

Methods: We used pre-existing permanent plots to evaluate the cover of understory after an unplanned wildfire.

Results: Across the entire stand nine of 47 species showed a significant change in cover. The largest change in a plant functional group was in the mosses, with all species present before fire being eliminated. There was no change in species diversity or total cover. At the plot level, species composition showed a much greater change. An average of 47% of the species present in a plot before the fire were absent in the same plot after the fire, and the total species turnover in plots was 88% of the species present before the fire. The plots showed a similar shift in species composition.

Conclusions: These results confirm that boreal forest communities show a high degree of resilience to fire, but within a forest stand species will be found in different locations following fire, potentially exposing them to a different set of biotic and abiotic conditions in these new locations.