Black Spruce-feathermoss Research Articles

Long-Term Simulated Nitrogen Deposition Has Moderate Impacts on Soil Microbial Communities across Three Bioclimatic Domains of the Eastern Canadian Forest

The soil microbiome plays major roles in the below-ground processes and productivity of forest ecosystems. Atmospheric nitrogen (N) deposition is predicted to increase globally and might create disturbances in soil microbial communities, essentially by modifying soil chemistry. However, the impacts of higher N deposition on the soil microbiome in N-limited northern forests are still unclear. For 16 years, we simulated N deposition by adding ammonium nitrate at rates of 3 and 10 times the ambient N deposition directly into soils located in three bioclimatic domains of the eastern Canadian forest (i.e., sugar maple–yellow birch, balsam fir–white birch, and black spruce–feather moss). We identified changes in the microbial communities by isolating the DNA of the L, F, and H soil horizons, as well as by sequencing amplicons of the bacterial 16S rRNA gene and the fungal ITS region. We found that long-term increased N deposition had no effect on soil microbial diversity, but had moderate impacts on the composition of the bacterial and fungal communities. The most noticeable change was the increase in ectomycorrhizal fungi ASV abundance, potentially due to increased tree root growth on fertilized plots. Our work suggests that, in N-limited northern forests, extra N is rapidly mobilized by vegetation, thus minimizing impacts on the soil microbiome.

Comparative Ecophysiology of Black Spruce between Lichen Woodlands and Feathermoss Stands in Eastern Canada

Climate change is likely to affect the growth, development and regeneration of the black spruce stands across the boreal forest. Regeneration failures cause gaps in the dense black spruce-feathermoss (SM) mosaic increasing the landscape proportion of open lichen woodland (LW). The aims of the study are to determine whether the contrasting characteristics of SM and LW induce different maximum carboxylation rate (Vcmax), maximum electron transport rate (Jmax) and light-saturated maximum photosynthesis (Amax) in black spruce trees across a latitudinal or seasonal gradient. Results show that the Vcmax and Jmax were higher in SM than in LW in western Quebec, at the ecotone of the closed-crown and open forest. Vcmax and Jmax were different between SM and LW mainly because nutrient acquisition seems different between stand types. Latitude affects values of Vcmax and Jmax, but the effect could be explained by soil and vegetation composition between experimental plots rather than by latitude. Physiological capacities do not match Amax values for stand types and latitude. Indeed, Amax rates suggest that black spruce in LWs perform as well as those in SMs at the needle scale because Amax would be limited by CO2 concentration which prevents saturation of Rubisco. Despite the lack of difference between the Amax of SM and LW stands, future increases in CO2 concentration and temperature could induce a gap between their respective photosynthesis rates because of their different physiological capacities.

Regional Climate, Edaphic Conditions and Establishment Substrates Interact to Influence Initial Growth of Black Spruce and Jack Pine Planted in the Boreal Forest

In eastern Canada, spruces (Picea spp.) and pines (Pinus spp.) are among the main commercial species being logged for their lumber or wood fiber. Annually, about 175 million seedlings are planted in areas totaling ~100,000 ha. Appropriate microsite selection is essential during reforestation operations, given that it can improve the chances of survival and initial growth of the seedlings. In fir (Abies spp.) and spruce forests of eastern Canada, the optimal characteristics of establishment microsites have yet to be identified; these would be determined by different physical and climatic variables operating at several scales. Our study determined the influence of climatic (regional-scale), edaphic (stand-scale), local (microsite-scale) and planting conditions on the establishment substrate and initial growth of black spruce (Picea mariana Britton, Sterns and Poggenb.) and jack pine (Pinus banksiana Lamb.). Substrate characterization and growth monitoring (three growing seasons) for the two species were conducted on 29 planted cutblocks that were distributed over an east–west climatic gradient (precipitation and temperature) in the balsam fir and black spruce–feather moss forests of Quebec (Canada). Linear mixed models and multivariate analyses (PCAs) determined the effects of climatic, edaphic and micro-environmental variables and their interactions on the establishment substrate and seedling initial growth. The predictive models explained, respectively, 61% and 75% of the growth variability of black spruce and jack pine. Successful establishment of black spruce and jack pine depended upon regional conditions of precipitations and temperature, as well as on their interactions with stand-scale edaphic variables (surface deposit, drainage and slope) and local variables (micro-environmental) at the microsite-scale (establishment substrate types and substrate temperature). Mineral, organo-mineral and organic establishment substrates exerted mixed effects on seedling growth according to regional precipitation and temperature conditions, as well as their interactions with edaphic and local variables at the stand and microsite-scales, respectively.

Comparing soil profiles of adjacent forest stands with contrasting tree densities: lichen woodlands vs. black spruce–feathermoss stands in the continuous boreal forest

We studied mature and adjacent open lichen–spruce woodlands (LWs) and closed-canopy spruce–feathermoss stands (FMs) growing under similar edaphic conditions in the continuous boreal forest zone in Quebec (Canada). A total of six pairs of stands were investigated by profile sampling. Stem density, basal area, and biomass were about four times greater in FMs than in LWs on an area basis. In the humus layer, total stocks of C and N and of exchangeable K, Ca, Mg, Al, and Na were 1.4–2.3 times larger in FM than in LW soils. The first 30 cm and the first metre of mineral soils in LWs and FMs displayed similar available nutrient pools except for total C stocks, which were more than twice as large in FM as in LW soils in these soil layers. For the whole profile, total stocks of C and N and stocks of exchangeable Ca and Mg were 1.3–2.6 times larger in FM than in LW soils. These results highlight the low intrinsic fertility of LW soils, primarily due to the humus layer, but also the importance of the biological control of C, N, and mineral nutrients in these boreal soils.

Range Extensions of 35 Bryophyte Species in the Black Spruce–Feather Moss Forest of Western Quebec, Canada

Although the North American bryophyte flora are relatively well known, bryophytes of the Black Spruce–feather moss forest in the Nord-du-Québec administrative region, especially its southern portion (49–51°N, 74–79°W), remain under-sampled. Here, we report 169 bryophyte taxa for this region, of which 35 (14 true mosses, 20 liverworts, one sphagnum) represent noteworthy records, including 20 taxa new for the region. These new occurrences close several gaps in distribution in the study area and, more broadly, in the boreal Black Spruce (Picea mariana) forest of adjacent ontario. Microhabitat preferences of the species are also documented. This work represents a substantial contribution to knowledge of the bryophyte flora, which will help refine protection priority ranks of species of Quebec and Labrador.

Boreal bryophyte response to natural fire edge creation

AbstractQuestionChanges in species richness and composition in plant communities as a result of edge creation are well documented in anthropogenically modified landscapes, but what happens after natural disturbance? We investigate the responses of boreal bryophytes to edge creation in a post‐wildfire residual forest patch system.LocationBoreal black spruce–feather moss forest, western Québec, Canada.MethodsBryophyte community was sampled in 50‐m2 plots: 117 plots in 39 undisturbed forest cores (control) and 108 plots in 30 residual forest patches from wildfire, divided into 48 core plots, 30 north‐facing edge plots and 30 south‐facing edge plots. Temporal, severity, spatial and structural characteristics of the stands were also recorded to explain bryophyte community composition.ResultsResidual edges and cores were more species‐rich than undisturbed cores, particularly north‐facing edges, but community composition differed between undisturbed and residual cores. Indicator species of undisturbed cores and residual edges differed in their species traits. Spatial variables explained variations in composition, with a larger difference between core and edge communities in patches over 3 ha compared to smaller patches.ConclusionWe observed community composition change in response to natural edge creation. While edge influence is reduced in large patches, the bryophyte community remains distinct from undisturbed cores. This community change is a ‘control’ in studies of boreal bryophyte response to anthropogenic edge creation, and suggests that we should consider the natural range of variability. Finally, this study emphasizes that forest remnants do not substitute for continuous and undisturbed forests.

Afforestation of Boreal Open Woodlands: Early Performance and Ecophysiology of Planted Black Spruce Seedlings

Open lichen woodlands (LWs) are degraded stands that lack the ability to regenerate naturally due to a succession of natural and/or anthropogenic disturbances. As they represent both interesting forest restoration and carbon sequestration opportunities, we tested disc scarification and planting of two sizes of containerized black spruce (Picea mariana Mill. (BSP)) seedlings for their afforestation. We compared treatment of unproductive LWs to reforestation of harvested, closed-crown black spruce-feathermoss (BSFM) stands. After one year, seedling survival and nutritional status were equivalent among stand types but despite higher root elongation index (REI), planted seedlings in LWs had lower relative growth rate, smaller total biomass and stem diameter than those in BSFM stands. Soil fertility variables, soil temperature, nor seedling water potential, helped at explaining this early growth response. Disc scarification significantly improved seedling first-year survival, biomass and foliar nutrient concentrations of P, Ca, and Mg. Smaller planting stock showed higher REI, higher shoot water potential, and higher foliar nutrient concentration of all but one of the measured nutrients (N, P, K and Mg). Hence, preliminary results suggest that planting of smaller containerized black spruce stock, combined with disc scarification, shows potential for afforestation of unproductive LWs. The impact of the lichen mat and other potential growth limiting factors on afforestation of these sites requires further investigation.

Lichen-Spruce Woodland Early Indicators of Ecological Resilience Following Silvicultural Disturbances in Québec’s Closed-Crown Forest Zone

Lichen woodlands (LW) located in the closed-crown boreal forest are not a successional stage moving towards a closed black spruce feathermoss stand (FM), but an alternative stable state, due to their previous forest history, and the occurrence of LWs located nearby closed-crown FM stands. Therefore, afforestation in those LWs through site preparation and plantation could shift back LW into FM stands. We implemented an experimental design with different combinations of silvicultural treatments in both site types (LW, FM). We monitored the evolution of plant diversity and the physiology of three bio-indicators (Picea mariana, Kalmia angustifolia, Rhododendron groenlandicum) in different microsites created by the silvicultural treatments. The return to the initial composition was noticed only two years after treatments, especially in the LW stands, thus indicating a higher level of early ecosystem resilience in LWs compared to FM stands. Mean species cover, especially in the FM stands, decreased the most in the skid trails created by logging, probably due to a lack of acclimation of bryophytes to open stand conditions. Conversely, ericaceous shrubs and lichens found in the LWs were already acclimated to open stand conditions, which give to LWs a restructuring advantage compared to FM plant communities after silvicultural treatments. Overall, FM and LW short-term resilience was similar, indicating equally efficient ecosystem reorganization in both stands. The comparable early resilience in managed LW and FM stands, in terms of plant biodiversity, contradicts the presumed fragility of LW stands, especially in this case where LWs are assumed to be an alternative stable state created by compound disturbances. Silvicultural treatments maintained the functional group diversity in LWs, a key element for ecosystem resilience. Therefore, this study support the idea that plantation following site preparation in LWs could be a valuable management strategy to reach several objectives, such as increasing forest carbon sinks.

Is the closed‐crown boreal forest resilient after successive stand disturbances? A quantitative demonstration from a case study

AbstractQuestionRecent studies revealed by indirect evidence that lichen–spruce woodlands in the closed‐crown boreal forest are an alternative stable state of black spruce–feathermoss stands. This paper aims to demonstrate and quantify black spruce forest cover regression following successive wildfire events in the closed‐crown forest zone during the 20th century. A case study in north‐eastern North American closed‐crown forest zone is presented.LocationThe study area is at the centre of the closed‐crown boreal forest zone of eastern Canada (49°59′ N, 71°59′ W).MethodsA burned area of 25 ha was selected in which three tree cohorts were visible. The first cohort contains partly buried trunks and stumps of a stand destroyed by a forest fire in 1920, the second contains standing dead trees from a 1995 forest fire and the third contains seedlings established after the 1995 fire. Fires were dated by scar analyses of living trees at the edge of the site. An array of sample plots was established in autumn 1997 to evaluate each cohort in terms of stocking and tree density.ResultsBlack spruce stocking decreased significantly between cohorts, from 43.5% in Cohort 1 (all coniferous trees) to 25.6% in Cohort 2 and 11.5% in Cohort 3 (2011). Jack pine showed an inverse tendency, from 0% in Cohort 1 (not determined) to 0% in Cohort 2 and 3.8% in Cohort 3. Black spruce stems per hectare also dropped significantly from a minimum of 1322 in Cohort 1 to 1088 in Cohort 2 and 976 in Cohort 3.ConclusionsThis study is the first quantitative demonstration that lichen–spruce woodlands in the closed‐crown boreal forest zone are an alternative stable state of black spruce–feathermoss stands. Stocking levels and densities markedly declined after each of the two fires (1920 and 1995) that both occurred after spruce budworm outbreaks (1910 and 1980) and are most likely responsible for the shift from a closed‐crown to an open woodland.

Estimating the net carbon balance of boreal open woodland afforestation: A case-study in Québec’s closed-crown boreal forest

Black spruce ( Picea mariana (Mill.) B.S.P.) is the dominant tree species in the Canadian province of Québec’s boreal ecosystem, particularly in the black spruce-feathermoss (BSFM) domain (between the 49th and the 52nd parallels). While black spruce is generally well adapted to regenerate after wildfires, regeneration failure can sometimes occur, resulting in the irreversible conversion of closed-crown BSFM to open black spruce-lichen woodlands (OW). With OWs representing approximately 7% (1.6 M ha) of Québec’s BSFM domain, the afforestation of OWs carries significant theoretical potential for carbon (C) sequestration, which has not yet been evaluated. The main objectives of the study were then: (i) to estimate the theoretical C balance of OW afforestation within the closed-crown BSFM domain in Québec’s boreal forest; (ii) to calculate, using the life cycle analysis (LCA) method, all the GHG emissions related to black spruce OW afforestation in the closed-crown BSFM domain of Québec. The CO2FIX v. 3.1 model was used to calculate the biological C balance between the baseline (natural OW of site index 9 at age 50) and afforestation (black spruce plantation of site index 6 at age 25) scenarios, using the best estimates available for all five recommended C compartments (aboveground biomass, belowground biomass, litter, deadwood, and soil). The simulation revealed a biological C balance of 77.0 t C ha −1, 70 years following afforestation, for an average net sequestration rate of 1.1 t C ha −1 year −1. Biological C balance only turns positive after 27 years. When integrating the uncertainties related to both the plantation growth yield and the wildfire disturbance, the average sequestration rate varies between 0.2 and 1.9 t C ha −1 year −1. GHG emissions are 1.3 t CO 2 equiv. ha −1 for all afforestation-related operations, which is less than 0.5% of the biological C balance after 70 years. Thus, GHG emissions do not significantly affect the net C balance of the afforestation project simulated. Several recommendations are made, mostly centered on the factors influencing the growth rate of carbon stocks and the impact of natural disturbances, to minimize the range of uncertainties associated to the sequestration potential and maximize the mitigation benefits of an OW afforestation project.

Growth response and water relations of 3-year-old planted black spruce and jack pine seedlings in site prepared lichen woodlands

Black spruce-lichen woodlands are low tree density stands within the closed crown, North American boreal forest that represent a diverging post-fire type of black spruce forest. As natural densification of lichen woodlands has never been observed, afforestation remains the only way of shifting these stands to closed canopy stands. The objective of the study was to evaluate site preparation effects on growth and water relations of black spruce ( Picea mariana (Mill.) B.S.P.) and jack pine ( Pinus banksiana Lamb.) seedlings in black spruce-lichen woodlands (LW), compared to managed black spruce-feathermoss stands (BSFM). Site preparation treatments in LW were no preparation (LWno), patch scarification (LWps), and disk scarification (LWds). The operationally managed BSFM stands stood for the control. Results from the third growing season indicate that soil water availability in intact or lightly prepared patch scarified LW is a limiting variable for seedling growth for both black spruce and jack pine seedlings. However, once LW are prepared with disk scarification, this planting check factor is significantly reduced, to the point of being equal to BSFM in terms of water availability and seedling water relations. The significant seedling growth difference, in favor of BSFM, might be a consequence of the lower level of pre-plantation disturbance in LW, compared to BSFM. Our results also suggest that jack pine, through higher water stress tolerance, could constitute a wise silvicultural choice over black spruce, especially with LW established on coarse drought-prone material.

Influence of Aspen on Forest Floor Properties in Black Spruce-dominated Stands

In the absence of fire in black spruce-feathermoss stands, a thick forest floor layer dominated by bryophytes and sphagnum accumulates. This layer is associated with wet, cool and nutrient-poor soil conditions conducive to the paludification process and pushing the ecosystem towards an unproductive open black spruce forest. The presence of Populus tremuloides in theses stands may halt this process because this species has a high nutrient cycling rate and a litter that represses moss cover. The main hypothesis of this study is that, despite similar abiotic conditions (slope and drainage), the presence of Populus tremuloides in a stand dominated by Picea mariana affects surface soil nutrient availability, total N, pH as well as the decomposition process. The abundance of Populus tremuloides trees was associated with higher exchangeable cations, cationic exchangeable capacity and pH of the forest floor layer on all sites. A decrease in organic matter thickness with increasing aspen presence was also found on all sites, suggesting that this species affects the decomposition process by the quality of its litter as well as by a general improvement of soil physical and chemical properties. The decomposition rate of a standard substrate as well as in vitro potential net nitrogen mineralization were positively related to Populus tremuloides on only one of the three sites, and non-significant on the other sites. Strong immobilization of added nitrogen during incubation was observed on all sites and was not related to aspen, which suggested that in these stands, the soil microbial community is uniformly and strongly nitrogen limited. The zone of influence of Populus tremuloides was evaluated in areas around the soil sampling plot ranging from 3 to 7 m. The results revealed that this zone varies with soil properties. The results suggest that the presence of Populus tremuloides accelerate nutrient cycling, which could affect stand productivity to some extent.

SOIL CARBON, NITROGEN, AND PHOSPHORUS STOCKS AND DYNAMICS UNDER DISTURBED BLACK SPRUCE FORESTS

In the North American boreal forest, black spruce (Picea mariana) forests have historically regenerated after the passage of large-scale wildfires. Over the past 30 years, tree harvesting has replaced wildfire as the predominant agent of disturbance in black spruce–feathermoss communities in the Lac St. Jean–Chibougamau region of Quebec, Canada. This study addressed how natural and anthropogenic disturbances altered soil and fine root carbon (C), nitrogen (N), and phosphorus (P) stocks and dynamics under four disturbance types in central Quebec including: (1) recently burned sites, (2) sites recently harvested using a practice called “cutting with protection of tree regeneration and soils” (CPRS), (3) sites that were burned 75–85 yr ago, and (4) sites that were horse-logged 55 yr ago. Soil N contents in the surface organic layer of the recently burned sites (91 g/m2) were significantly lower than under the old burn sites (146 g/m2). Using equivalent soil masses, total N stores in the mineral soils under the recently harvested stands (76 g/m2) were significantly lower than N stores under the old burn sites (114 g/m2). In field incubations from June to October 1997, net N mineralization rates in the organic horizons ranged from 3.3 kg/ha (recent burn) to 17.1 kg/ha (old harvest). The largest pools of labile inorganic-P (Bray 2 extractable) were in the organic horizons of the old harvest and old burn sites (59.1 and 55.3 mg/kg, respectively), and these pools increased up to 24% from the beginning to the end of the growing season. Pools of dissolved organic nitrogen in surface organic horizons in all of the disturbance types were highest just after snowmelt and declined steadily through the growing season. The older disturbance types had higher N stores in fine roots compared to the recently disturbed sites, and fine root length in the old harvest sites (1623 m/m2) was higher than in the stands burned 75–85 yr ago (917 m/m2).

Soil Carbon, Nitrogen, and Phosphorus Stocks and Dynamics under Disturbed Black Spruce Forests

In the North American boreal forest, black spruce (Picea mariana) forests have historically regenerated after the passage of large-scale wildfires. Over the past 30 years, tree harvesting has replaced wildfire as the predominant agent of disturbance in black spruce–feathermoss communities in the Lac St. Jean–Chibougamau region of Quebec, Canada. This study addressed how natural and anthropogenic disturbances altered soil and fine root carbon (C), nitrogen (N), and phosphorus (P) stocks and dynamics under four disturbance types in central Quebec including: (1) recently burned sites, (2) sites recently harvested using a practice called “cutting with protection of tree regeneration and soils” (CPRS), (3) sites that were burned 75–85 yr ago, and (4) sites that were horse-logged 55 yr ago. Soil N contents in the surface organic layer of the recently burned sites (91 g/m2) were significantly lower than under the old burn sites (146 g/m2). Using equivalent soil masses, total N stores in the mineral soils under the recently harvested stands (76 g/m2) were significantly lower than N stores under the old burn sites (114 g/m2). In field incubations from June to October 1997, net N mineralization rates in the organic horizons ranged from 3.3 kg/ha (recent burn) to 17.1 kg/ha (old harvest). The largest pools of labile inorganic-P (Bray 2 extractable) were in the organic horizons of the old harvest and old burn sites (59.1 and 55.3 mg/kg, respectively), and these pools increased up to 24% from the beginning to the end of the growing season. Pools of dissolved organic nitrogen in surface organic horizons in all of the disturbance types were highest just after snowmelt and declined steadily through the growing season. The older disturbance types had higher N stores in fine roots compared to the recently disturbed sites, and fine root length in the old harvest sites (1623 m/m2) was higher than in the stands burned 75–85 yr ago (917 m/m2).

Classification, Description, and Dynamics of Upland Plant Communities within a Minnesota Wilderness Area

The major upland plant community types of the Boundary Waters Canoe Area (BWCA) of northeastern Minnesota, identified by multivariate analyses (clustering and canonical and discriminant analysis) of 68 stands disturbed by logging and 106 stands undisturbed by logging, include the following: lichen, jack pine—oak, red pine, jack pine—black spruce, jack pine—fir, black spruce—feather moss, maple—oak, aspen—birch, aspen—birch—white pine, maple—aspen—birch, maple—aspen—birch—fir, fir—birch, and white cedar. Each of these types is based on a complex of 53 common species, through the name may incorrectly imply that one or two dominant overstory species are indicative of the type. Other forest stands from the BWCA are quantitatively related to the regional vegetation through discriminant analysis. Succession on the uplands in the area, without disturbance, leads to fir—birch and ultimately to the white—cedar community type. Whitetail deer may have had an impact on restricting the occurrence and reproduction of the white cedar type.