PLFA Profiling of Microbial Community Structure and Seasonal Shifts in Soils of a Douglas-fir Chronosequence

The impact and frequency of forest harvesting could significantly affect soil microbial community (SMC) structure and functioning. The ability of soil microorganisms to perform biogeochemical processes is critical for sustaining forest productivity and has a direct impact on decomposition dynamics and carbon storage potential. The Wind River Canopy Crane Research Forest in SW, WA, provided a unique opportunity to study a forest chronosequence and the residual effects of harvesting on the SMC in comparison to old-growth forests. The objective of this study was to determine the effect of clear-cutting and stand age on temporal dynamics of SMC and physiological stress markers using phospholipid fatty acid (PLFA) profiling. Soil microbial PLFA profiles were determined seven times over 22 months (Nov. 02 to Sep. 04) in old-growth coniferous forest stands (300–500 years) and 8 (CC8)- or 25 (CC25)-year-old replanted clear-cuts. PLFA patterns of the SMC shifted because of clear-cutting, but seasonal temporal changes had greater shifts than differences among stand age. The microbial biomass (total PLFA) and bacterial, fungal, and selected other PLFAs were significantly reduced in CC8 but not in CC25 sites relative to the old-growth sites. An increase in stress indicators [PLFA ratios of saturated/monsaturated and (cy17:0 + cy19:0)/(16:1ω7 + 18:1ω7)] in late summer was related to water stress. Although the canopy and litter input are quite different for a 25-year clear-cut compared to virgin old-growth forest, we conclude that the composition of the microbial communities, 25 years after clear-cutting, has recovered sufficiently to be much more similar to old-growth forests than a recent clear-cut at this Pacific Northwest forest site. The study shows the potential of PLFA analysis for profiling microbial communities and their stress status under field conditions, but wide temporal shifts emphasize the need for sampling over seasons to fully interpret ecosystem management impacts on microbial populations.     

Tree Mortality Following Boreal Forest Fires Reveals Scale-Dependant Interactions Between Community Structure and Fire Intensity

Fire disturbance patterns influence forest communities at a range of spatial scales. Forest community structure may also influence fire disturbance patterns, because tree species vary in their fuel value and in their tolerance to fire damage. However, the influence of community structure on fire disturbance likely depends on latent ecological differences between fires and on the spatial scale at which patterns are observed. Using data on fire intensity, community structure, and post-fire tree survival in four systematically sampled boreal forest fires, we tested the hypotheses that: (1) patterns in post-fire tree survival reflect interactions between fire intensity and community structure; (2) these relationships change with the spatial scale of observation. To test the first hypothesis, we used information theoretic methods to compare eight generalized linear mixed effects models describing the influence of community structure and fire intensity on tree survival in a 500 m² sample plot, accounting for latent fire-to-fire differences in response. To test the scaling hypothesis, we reaveraged the data at nine successively larger spatial resolutions up to approximately 2 km², at each resolution tracking the parameter values of the best model. When fit to the plot-level data, the dominant feature of the best model was a strong intensity–survival correlation which varied from fire to fire, and depended on plot-level community structure. In some fires, community structure and survival became more tightly coupled at larger scales, whereas fire intensity became less important. These results support the view that fire disturbance patterns are influenced by cross-scale interactions between community structure and fire intensity.     

Changes in Soil Fungal Communities, Extracellular Enzyme Activities, and Litter Decomposition Across a Fire Chronosequence in Alaskan Boreal Forests

Wildfires are a pervasive disturbance in boreal forests, and the frequency and intensity of boreal wildfires is expected to increase with climate warming. Boreal forests store a large fraction of global soil organic carbon (C), but relatively few studies have documented how wildfires affect soil microbial communities and soil C dynamics. We used a fire chronosequence in upland boreal forests of interior Alaska with sites that were 1, 7, 12, 24, 55, ~90, and ~100 years post-fire to examine the short- and long-term responses of fungal community composition, fungal abundance, extracellular enzyme activity, and litter decomposition to wildfires. We hypothesized that post-fire changes in fungal abundance and community composition would constrain decomposition following fires. We found that wildfires altered the composition of soil fungal communities. The relative abundance of ascomycetes significantly increased following fire whereas basidiomycetes decreased. Post-fire decreases in basidiomycete fungi were likely attributable to declines in ectomycorrhizal fungi. Fungal hyphal lengths in the organic horizon significantly declined in response to wildfire, and they required at least 24 years to return to pre-fire levels. Post-fire reductions in fungal hyphal length were associated with decreased activities of hydrolytic extracellular enzymes. In support of our hypothesis, the decomposition rate of aspen and black spruce litter significantly increased as forests recovered from fire. Our results indicate that post-fire reductions in soil fungal abundance and activity likely inhibit litter decomposition following boreal wildfires. Slower rates of litter decay may lead to decreased heterotrophic respiration from soil following fires and contribute to a negative feedback to climate warming.     

Diversity and Community Structure of Primary Wood-Inhabiting Bacteria in Boreal Forest

DNA-based pyrosequencing analysis of the V1- V3 16S rRNA gene region was used to identify bacteria community and shift during early stages of wood colonization in boreal forest soils. The dataset comprised 142,447 sequences and was affiliated to 11 bacteria phyla, 25 classes and 233 genera. The dominant groups across all samples were Proteobacteria, followed by Bacteroidetes, Acidobacteria, Actinobacteria, Amatimonadetes, Planctomycetes and TM7 group. The community structure of the primary wood-inhabiting bacteria differed between types of forest soils and the composition of bacteria remained stable over prolonged incubation time. The results suggest that variations in soil bacterial community composition have an influence on the wood-inhabiting bacterial structure.     

Boreal Forest Floor Greenhouse Gas Emissions Across a Pleurozium schreberi-Dominated,Wildfire-Disturbed Chronosequence

Ecosystems - The boreal forest is a globally critical biome for carbon cycling. Its forests are shaped by wildfire events that affect ecosystem properties and climate feedbacks including greenhouse...     

Spatiotemporal Variations of Fire Frequency in Central Boreal Forest

Determination of the direct causal factors controlling wildfires is key to understanding wildfire–vegetation–climate dynamics in a changing climate and for developing sustainable management strategies for biodiversity conservation and maintenance of long-term forest productivity. In this study, we sought to understand how the fire frequency of a large mixedwood forest in the central boreal shield varies as a result of temporal and spatial factors. We reconstructed the fire history of an 11,600-km² area located in the northwestern boreal forest of Ontario, using archival data of large fires occurring since 1921 and dendrochronological dating for fires prior to 1921. The fire cycle decreased from 295 years for the period of 1820–1920 to approximately 100 years for the period of 1921–2008. Spatially, fire frequency increased with latitude, attributable to higher human activities that have increased fragmentation and fire suppression in the southern portion of the study area. Fire frequency also increased with distance to waterbodies, and was higher on Podzols that were strongly correlated with moderate drainage and coniferous vegetation. The temporal increase of fire frequency in the central region, unlike western and eastern boreal forests where fire frequency has decreased, may be a result of increased warm and dry conditions associated with climate change in central North America, suggesting that the response of wildfire to global climate change may be regionally individualistic. The significant spatial factors we found in this study are in agreement with other wildfire studies, indicating the commonality of the influences by physiographic features and human activities on regional fire regimes across the boreal forest. Overall, wildfire in the central boreal shield is more frequent than that in the wetter eastern boreal region and less frequent than that in the drier western boreal region, confirming a climatic top-down control on the fire activities of the entire North American boreal forest.     

The Effects of Spatial Legacies following Shifting Management Practices and Fire on Boreal Forest Age Structure

Forest age structure and its spatial arrangement are important elements of sustainable forestry because of their effects on biodiversity and timber availability. Forest management objectives that include specific forest age structure may not be easily attained due to constraints imposed by the legacies of historical management and natural disturbance. We used a spatially explicit stochastic model to explore the synergetic effects of forest management and fire on boreal forest age structure. Specifically, we examined (1) the duration of spatial legacies of different management practices in the boreal forest, (2) how multiple shifts in management practices affect legacy duration and the spatial trajectories of forest age structure, and (3) how fire influences legacy duration and pattern development in combination with harvesting. Results based on 30 replicates of 500 years for each scenario indicate that (1) spatial legacies persist over 200 years and the rate at which legacies are overcome depends on whether new management targets are in synchrony with existing spatial pattern; (2) age specific goals were met faster after multiple management shifts due to the similar spatial scale of the preceding management types; (3) because large fires can erase the spatial pattern created by smaller disturbances, scenarios with fire had shorter lags than scenarios without fire. These results suggest that forest management goals can be accelerated by applying management at a similar spatial scale as existing spatial patterns. Also, management planning should include careful consideration of historical management as well as current and likely future disturbances.     

Effects of Forest Use on Aphyllophoraceous Fungal Community Structure in Sarawak, Malaysia

Aphyllophoraceous fungi are expected to reflect changes in the environmental conditions caused by forest use. To reveal the effects of forest uses on the fungal community structure, we performed a 3‐month survey of aphyllophoraceous species in five forest types (undisturbed primary forest, isolated patches of primary forest, old and young fallow forest, and rubber plantations) in Sarawak, Malaysia in 2005. We used a canonical correspondence analysis (CCA) to reveal the relationships between fungal community composition and the environmental variables (canopy openness, soil water potential, amount and composition of coarse woody debris, litter mass, basal area, plant species composition). A total of 155 samples from 67 species were collected during the study period. The fungal species density represented by the number of species in a transect differed significantly among forest types. The fungal species density increased significantly with increasing number of pieces of coarse woody debris (CWD), but decreased significantly with increasing the scores of second axis of principal component analysis (PCA) for plant species composition. In the CCA ordination, automatic forward selection revealed that only the number of pieces of CWD significantly affected the fungal species composition. The occurrences of Flabellophora licmophora, Coriolopsis retropicta, Microporus vernicipes, and Amauroderma subrugosum were positively correlated with the number of pieces of CWD. Our study clearly demonstrated that forest use negatively affected aphyllophoraceous fungal diversity and suggest that the quantity of CWD would be an important determinant of fungal diversity and composition.     

Structure and Function of Methanogens along a Short-Term Restoration Chronosequence in the Florida Everglades

The removal of plants and soil to bedrock to eradicate exotic invasive plants within the Hole-in-the-Donut (HID) region, part of the Everglades National Park (Florida), presented a unique opportunity to study the redevelopment of soil and the associated microbial communities in the context of short-term primary succession and ecosystem restoration. The goal of this study was to identify relationships between soil redevelopment and activity and composition of methanogenic assemblages in HID soils. Methane production potentials indicated a general decline in methanogenic activity with restoration age. Microcosm incubations strongly suggested hydrogenotrophic methanogenesis as the most favorable pathway for methane formation in HID soils from all sites. Culture-independent techniques targeting methyl coenzyme M reductase genes (mcrA) were used to assess the dynamics of methanogenic assemblages. Clone libraries were dominated by sequences related to hydrogenotrophic methanogens of the orders Methanobacteriales and Methanococcales and suggested a general decline in the relative abundance of Methanobacteriales mcrA with time since restoration. Terminal restriction fragment length polymorphism analysis indicated methanogenic assemblages remain relatively stable between wet and dry seasons. Interestingly, analysis of soils across the restoration chronosequence indicated a shift in Methanobacteriales populations with restoration age, suggesting genotypic shifts due to site-specific factors.     

Soil Bacterial Community Structure Responses to Precipitation Reduction and Forest Management in Forest Ecosystems across Germany

Soil microbial communities play an important role in forest ecosystem functioning, but how climate change will affect the community composition and consequently bacterial functions is poorly understood. We assessed the effects of reduced precipitation with the aim of simulating realistic future drought conditions for one growing season on the bacterial community and its relation to soil properties and forest management. We manipulated precipitation in beech and conifer forest plots managed at different levels of intensity in three different regions across Germany. The precipitation reduction decreased soil water content across the growing season by between 2 to 8% depending on plot and region. T-RFLP analysis and pyrosequencing of the 16S rRNA gene were used to study the total soil bacterial community and its active members after six months of precipitation reduction. The effect of reduced precipitation on the total bacterial community structure was negligible while significant effects could be observed for the active bacteria. However, the effect was secondary to the stronger influence of specific soil characteristics across the three regions and management selection of overstorey tree species and their respective understorey vegetation. The impact of reduced precipitation differed between the studied plots; however, we could not determine the particular parameters being able to modify the response of the active bacterial community among plots. We conclude that the moderate drought induced by the precipitation manipulation treatment started to affect the active but not the total bacterial community, which points to an adequate resistance of the soil microbial system over one growing season.     

Material Legacies and Environmental Constraints Underlie Fire Resilience of a Dominant Boreal Forest Type

Ecosystems - Resilience of plant communities to disturbance is supported by multiple mechanisms, including ecological legacies affecting propagule availability, species’ environmental...     

Species Composition of Saproxylic Fungal Communities on Decaying Logs in the Boreal Forest

Coarse woody debris supports large numbers of saproxylic fungal species. However, most of the current knowledge comes from Scandinavia and studies relating the effect of stand or log characteristics on the diversity and composition of decomposer fungi have not been conducted in Northeastern Canada. Logs from five tree species were sampled along a decomposition gradient in nine stands representing three successional stages of the boreal mixed forest of Northwestern Quebec, Canada. Using a molecular fingerprinting technique, we assessed fungal community Shannon–Weaver diversity index, richness, and composition. We used linear mixed models and multivariate analyses to link changes in fungal communities to log and stand characteristics. We found a total of 33 operational taxonomic units (OTUs) including an indicator species for balsam fir (similar to Athelia sp.) and one found only in aspen stands (similar to Calocera cornea). Spruce logs supported the highest fungal Shannon–Weaver diversity index and OTU number. Our results support the hypothesis that log species influences fungal richness and diversity. However, log decay class does not. Stand composition, volume of coarse woody debris, and log chemical composition were all involved in structuring fungal communities. Maintaining the diversity of wood-decomposing communities therefore requires the presence of dead wood from diverse log species.     

Decreases in Fire Spread Probability with Forest Age Promotes Alternative Community States, Reduced Resilience to Climate Variability and Large Fire Regime Shifts

The generalization that plant communities increase in flammability as they age and invariably lead to resilient self-organized landscape mosaics is being increasingly challenged. Plant communities often exhibit rapidly saturating or even hump-shaped age-flammability trajectories and landscapes often display strong non-linear behaviors, abrupt shifts, and self-reinforcing alternative community states. This plethora of fire-landscape interactions calls for a more general model that considers alternative age-flammability rules. We simulated landscape dynamics assuming communities that (1) increase in flammability with age and (2) gain flammability up to a certain age followed by a slight and moderate loss to a constant value. Simulations were run under combinations of ignition frequency and interannual climatic variability. Age-increasing fire probability promoted high resilience to changes in ignition frequency and climatic variability whereas humpbacked-shaped age-flammability led to strong non-linear behaviors. Moderate (20%) reductions in mature compared to peak flammability produced the least resilient behaviors. The relatively non-flammable mature forest matrix intersected by young flammable patches is prone to break up and disintegrate with slight increases in ignition/climate variability causing large-scale shifts in the fire regime because large fires were able to sweep through the more continuous young/flammable landscape. Contrary to the dominant perception, fire suppression in landscapes with positive feedbacks may effectively reduce fire occurrence by allowing less flammable later stage communities composed of longer lived, obligate seeders to replace earlier stages of light demanding, often more flammable resprouters. Conversely, increases in anthropogenic ignitions, a common global trend of many forested regions may, in synergism with increased climate variability, induce abrupt shifts, and large-scale forest degradation.     

Modelling Variable Fire Severity in Boreal Forests: Effects of Fire Intensity and Stand Structure

It is becoming clear that fires in boreal forests are not uniformly stand-replacing. On the contrary, marked variation in fire severity, measured as tree mortality, has been found both within and among individual fires. It is important to understand the conditions under which this variation can arise. We integrated forest sample plot data, tree allometries and historical forest fire records within a diameter class-structured model of 1.0 ha patches of mono-specific black spruce and jack pine stands in northern Québec, Canada. The model accounts for crown fire initiation and vertical spread into the canopy. It uses empirical relations between fire intensity, scorch height, the percent of crown scorched and tree mortality to simulate fire severity, specifically the percent reduction in patch basal area due to fire-caused mortality. A random forest and a regression tree analysis of a large random sample of simulated fires were used to test for an effect of fireline intensity, stand structure, species composition and pyrogeographic regions on resultant severity. Severity increased with intensity and was lower for jack pine stands. The proportion of simulated fires that burned at high severity (e.g. >75% reduction in patch basal area) was 0.80 for black spruce and 0.11 for jack pine. We identified thresholds in intensity below which there was a marked sensitivity of simulated fire severity to stand structure, and to interactions between intensity and structure. We found no evidence for a residual effect of pyrogeographic region on simulated severity, after the effects of stand structure and species composition were accounted for. The model presented here was able to produce variation in fire severity under a range of fire intensity conditions. This suggests that variation in stand structure is one of the factors causing the observed variation in boreal fire severity.     

Decomposition and Fragmentation of Coarse Woody Debris: Re-visiting a Boreal Black Spruce Chronosequence

We re-visited a seven-stand boreal chronosequence west of Thompson, Manitoba, Canada, in which coarse woody debris (CWD) and its instantaneous decomposition were measured in 2000. New CWD measurements were performed in 2007, and tree inventories updated to provide mortality and snag failure data. These data were used to model CWD changes, compare methods of estimating decomposition, and infer possible fragmentation rates. Measured CWD was between 9.7 (in both the 77- and 43-year-old stands) and 80.4 (in the 18-year-old stand) Mg ha⁻¹ in 2007. Spatial variability was high; at most stands CWD levels had not changed significantly from 2000 to 2007. Tree mortality was a significant flux only in older stands, whereas snag fall rate varied by an order of magnitude, from 2.9% y⁻¹ (0.2 Mg ha⁻¹ y⁻¹) in the 9-year-old stand to 9.8% y⁻¹ (2.3 Mg ha⁻¹ y⁻¹) in the 12-year-old stand. A one-pool model based on these inputs underestimated actual 2000–2007 CWD decomposition in the younger stands, suggesting that fragmentation could be an important part of the carbon flux exiting the CWD pool. We compared three independent measures of annual decomposition (k): direct measurements of CWD respiration, rates based on the 7-year re-sampling effort described here, and rates inferred from the chronosequence design itself. Mean k values arrived at via these techniques were 0.06 ± 0.03, 0.05 ± 0.04, and 0.05 ± 0.05 y⁻¹, respectively. The four-pool model suggested that the transition rate between decay classes was 0.14–0.19 y⁻¹; the model was most sensitive to initial CWD values. Although the computed k values implied a problem with chronosequence site selection for at least one site, the overall CWD trend was consistent with a larger number of sites surveyed in the region.     

Rapid Liana Colonization along a Secondary Forest Chronosequence

Lianas (woody vines) can have profound effects on tree recruitment, growth, survival, and diversity in tropical forests. However, the dynamics of liana colonization soon after land abandonment are poorly understood, and thus it is unknown whether lianas alter tree regeneration early in succession. We examined the liana community in 43 forests that ranged from 1 to 31 yr old in central Panama to determine how fast lianas colonize young forests and how the liana community changes with forest succession. We found that lianas reached high densities early in succession, commonly exceeding 1000 stems/ha within the first 5 yr of forest regeneration. Lianas also increased rapidly during early succession in terms of basal area but did not show evidence of saturation within the 30 yr of our chronosequence. The relative contribution of lianas to total woody plant community in terms of basal area and density increased rapidly and reached a saturation point within 5 yr (basal area) to 15 yr (density) after land abandonment. Our data demonstrate that lianas recruit early and in high density in tropical forest regeneration, and thus lianas may have a large effect on the way in which secondary forests develop both early and throughout succession.     

Forest Age and Plant Species Composition Determine the Soil Fungal Community Composition in a Chinese Subtropical Forest

Fungal diversity and community composition are mainly related to soil and vegetation factors. However, the relative contribution of the different drivers remains largely unexplored, especially in subtropical forest ecosystems. We studied the fungal diversity and community composition of soils sampled from 12 comparative study plots representing three forest age classes (Young: 10–40 yrs; Medium: 40–80 yrs; Old: ≥80 yrs) in Gutianshan National Nature Reserve in South-eastern China. Soil fungal communities were assessed employing ITS rDNA pyrotag sequencing. Members of Basidiomycota and Ascomycota dominated the fungal community, with 22 putative ectomycorrhizal fungal families, where Russulaceae and Thelephoraceae were the most abundant taxa. Analysis of similarity showed that the fungal community composition significantly differed among the three forest age classes. Forest age class, elevation of the study plots, and soil organic carbon (SOC) were the most important factors shaping the fungal community composition. We found a significant correlation between plant and fungal communities at different taxonomic and functional group levels, including a strong relationship between ectomycorrhizal fungal and non-ectomycorrhizal plant communities. Our results suggest that in subtropical forests, plant species community composition is the main driver of the soil fungal diversity and community composition.     

Modelling Spatial Interactions Among Fire,Spruce Budworm,and Logging in the Boreal Forest

In the boreal forest, fire, insects, and logging all affect spatial patterns in forest age and species composition. In turn, spatial legacies in age and composition can facilitate or constrain further disturbances and have important consequences for forest spatial structure and sustainability. However, the complex three-way interactions among fire, insects, and logging and their combined effects on forest spatial structure have seldom been investigated. We used a spatially explicit landscape simulation model to examine these interactions. Specifically, we investigated how the amount and the spatial scale of logging (cutblock size) in combination with succession, fire, and spruce budworm outbreaks affect area burned and area defoliated. Simulations included 30 replicates of 300 years for each of 19 different disturbance scenarios. More disturbances increased both the fragmentation and the proportion of coniferous species and imposed additional constraints on the extent of each disturbance. We also found that harvesting legacies affect fire and budworm differently due to differences in forest types consumed by each disturbance. Contrary to expectation, budworm defoliation did not affect area burned at the temporal scales studied and neither amount of logging nor cutblock size influenced defoliation extent. Logging increased fire size through conversion of more of the landscape to early seral, highly flammable forest types. Although logging increased the amount of budworm host species, spruce budworm caused mortality was reduced due to reductions in forest age. In general, we found that spatial legacies do not influence all disturbances equally and the duration of a spatial legacy is limited when multiple disturbances are present. Further information on post-disturbance succession is still needed to refine our understanding of long-term disturbance interactions.     

Fire and drought: Shifts in bark investment across a broad geographical scale for Neotropical savanna trees

Savanna tree communities occurring in confluence zones with other biomes likely experience different environmental pressures, resulting in shifts in the selection of individual traits, the combinations of such traits, and species composition. In seasonally dry fire-prone environments, plant survival is presumably associated with adaptive changes in bark properties related to fire protection and water storage. Here, we integrated the multiple functions of the bark to investigate whether different selective pressures could influence patterns of variation in bark structure and allocation across species in a broad geographical range. We measured thickness, density, and water content of the inner and outer bark in branches and the main stem of the 51 most abundant species in three savanna communities differing in climatic aridity, one located at the core region of Cerrado in Central Brazil and the other two at its periphery, in the transition zones with Amazonia and Atlantic forest biomes. We found no difference in outer bark thickness but markedly difference in inner bark thickness between the three plant communities. In the central region, where dry season is long and fire is frequent, branches and main stem showed thicker inner bark. Contrastingly, in the south periphery region, where dry season is short, species showed thinner inner bark in both branches and main stem. Species from the north periphery region, where mean annual precipitation is higher, but fire is frequent and the dry season is also long, showed similar main stem inner bark thickness, but thinner branch inner bark compared to core region species. Our findings support the idea that investing in inner bark thickness and bark moisture may be the most advantageous strategy in plant communities that suffer from high evaporative demand during a long period and are at a high risk of fire.