Mariana Stands Research Articles

Trees dying standing in the northeastern boreal old-growth forests of Quebec: spatial patterns, rates, and temporal variation

Spatial patterns, rates, and temporal variation of standing-tree mortality were studied in unmanaged boreal old-growth forests of northeastern Quebec. The study was carried out by sampling living and dead trees within 15 transects (400 m long, 40 m wide). The transects lay in stands that were classified according to their species composition in three types: dominated by black spruce, Picea mariana (Mill.) BSP; mixed P. mariana and balsam fir, Abies balsamea (L.) Mill.; and dominated by A. balsamea. Spatial patterns were analysed using Ripley's K function. The year of death was cross-dated using 190 sample discs extracted from dead standing A. balsamea and P. mariana to assess the rates and temporal variation of mortality. The spatial patterns of standing dead trees in P. mariana stands were predominantly clustered. The spatial patterns of large dead trees (>19 cm diameter at breast height (1.3 m height; DBH)) in mixed and A. balsamea-dominated stands were mainly random, with few stands showing clustered patterns. Small dead trees (9–19 cm DBH) in these stands were generally more clustered than larger trees. Tree mortality varied from year to year, though some mortality was observed in all the studied stand types for almost every year. Standing trees that had recently died accounted for 62%, 48%, and 51% of overall mortality in P. mariana-dominated, mixed, and A. balsamea-dominated stands, respectively. The results of this study indicate that mortality of standing trees outside of episodic mortality events (such as insect outbreaks) is an important process in the creation of structural complexity and habitat diversity in these stands.

Understorey vegetation in boreal Picea mariana and Populus tremuloides stands in British Columbia

Abstract. We compared the species composition and species density of vascular plants in the understorey vegetation of boreal forest between Picea mariana (Black spruce) and Populus tremuloides (Trembling aspen) stands in British Columbia, Canada, and related differences in species composition and species density between the two forest types to dominant canopy tree species as well as a wide variety of environmental factors. We analysed 231 stands, distributed in three different climatic regions representing drier, wetter, and milder variations of montane boreal climate. Of these stands 118 were dominated by P. mariana and 113 by P. tremuloides. P. tremuloides stands had higher species density than P. mariana stands in all climatic regions, but species density of each dominance type varied among climatic regions. The floristic composition of the understorey vegetation was markedly different for P. mariana and P. tremuloides dominated stands.A detailed study on the effect of canopy dominance and local environmental factors on the understorey vegetation of the boreal forest was conducted using 88 stands from one of the three climatic regions. Using a combination of ordination and variation partitioning by constrained ordination we demonstrated a small but unique effect of canopy dominance type on the understorey vegetation, while a larger amount of compositional variation was shared with other factors. Our results accord with a scenario in which differences in primary environmental factors and humus form properties, the latter accentuated by the canopy dominants themselves, are the most important causes of higher species density in P. tremuloides stands than in P. mariana stands, as well as differences in species composition among the two canopy dominance types. Processes and time scales involved in the small but significant direct and indirect effects of the canopy dominant on understo‐ rey species composition are discussed.

Understorey vegetation in boreal Picea mariana and Populus tremuloides stands in British Columbia

We compared the species composition and species density of vascular plants in the understorey vegetation of boreal forest between Picea mariana (Black spruce) and Populus tremuloides (Trembling aspen) stands in British Co- lumbia, Canada, and related differences in species composi- tion and species density between the two forest types to dominant canopy tree species as well as a wide variety of environmental factors. We analysed 231 stands, distributed in three different climatic regions representing drier, wetter, and milder variations of montane boreal climate. Of these stands 118 were dominated by P. mariana and 113 by P. tremuloides. P. tremuloides stands had higher species density than P. mariana stands in all climatic regions, but species density of each domi- nance type varied among climatic regions. The floristic compo- sition of the understorey vegetation was markedly different for P. mariana and P. tremuloides dominated stands. A detailed study on the effect of canopy dominance and local environmental factors on the understorey vegetation of the boreal forest was conducted using 88 stands from one of the three climatic regions. Using a combination of ordination and variation partitioning by constrained ordination we dem- onstrated a small but unique effect of canopy dominance type on the understorey vegetation, while a larger amount of com- positional variation was shared with other factors. Our results accord with a scenario in which differences in primary envi- ronmental factors and humus form properties, the latter accen- tuated by the canopy dominants themselves, are the most important causes of higher species density in P. tremuloides stands than in P. mariana stands, as well as differences in species composition among the two canopy dominance types. Processes and time scales involved in the small but significant direct and indirect effects of the canopy dominant on understo- rey species composition are discussed.

Environmental controls on soil CO2 flux following fire in black spruce, white spruce, and aspen stands of interior Alaska

Boreal forests contain large amounts of stored soil carbon and are susceptible to periodic disturbance by wildfire. This study evaluates the relationship between post-fire changes in soil temperature, moisture, and CO2 exchange in paired burned and control stands of three Alaskan forest systems: Picea mariana (Mill.) BSP, Picea glauca (Moench) Voss, and Populus tremuloides Michx. In these systems, the environmental factor that most directly controlled rates of carbon exchange varied depending upon burn status and soil drainage. In mature unburned stands, CO2 flux was highly correlated with seasonal patterns of soil temperature. Following fire, these soils became significantly warmer, and carbon exchange became more sensitive to fluctuations in surface moisture conditions. The effect of fire on soil climate was most pronounced in the P. mariana stands, which experienced a mean seasonal temperature increase of 5–8°C in the upper 1 m of the soil profile, a 200% increase in the rate of active layer thaw, and a reduction in mean surface moisture potential. Evidence from soil CO2 profiles suggests that these environmental changes may have resulted in enhanced decomposition of carbon previously immobilized by permafrost, potentially transforming a landscape that was once a net sink for carbon into a carbon source.

Implications of floristic and environmental variation for carbon cycle dynamics in boreal forest ecosystems of central Canada

Abstract. Species composition, detritus, and soil data from 97 boreal forest stands along a transect in central Canada were analysed using Correspondence Analysis to determine the dominant environmental/site variables that differentiate these forest stands. Picea mariana stands were densely clustered together on the understorey DCA plot, suggesting a consistent understorey species composition (feather mosses and Ericaceae), whereas Populus tremuloides stands had the most diverse understorey species composition (ca. 30 species, mostly shrubs and herbs). Pinus banksiana stands had several characteristic species of reindeer lichens (Cladina spp.), but saplings and Pinus seedlings were rare. Although climatic variables showed large variation along the transect, the CCA results indicated that site conditions are more important in determining species composition and differentiating the stand types. Forest floor characteristics (litter and humus layer, woody debris, and drainage) appear to be among the most important site variables. Stands of Picea had significantly higher average carbon (C) densities in the combined litter and humus layer (43530 kg‐C.ha‐1) than either Populus (25 500 kg‐C.ha‐1) or Pinus (19 400 kg‐C.ha‐1). The thick surface organic layer in lowland Picea stands plays an important role in regulating soil temperature and moisture, and organic‐matter decomposition, which in turn affect the ecosystem C‐dynamics. During forest succession after a stand‐replacing disturbance (e.g. fires), tree biomass and surface organic layer thickness increase in all stand types as forests recover; however, woody biomass detritus first decreases and then increases after ca. 80 yr. Soil C densities show slight decrease with ages in Populus stands, but increase in other stand types. These results indicate the complex C‐transfer processes among different components (tree biomass, detritus, forest floor, and soil) of boreal ecosystems at various stages of succession.

Implications of floristic and environmental variation for carbon cycle dynamics in boreal forest ecosystems of central Canada

Species composition, detritus, and soil data from 97 boreal forest stands along a transect in central Canada were analysed using Correspondence Analysis to determine the dominant environmental/site variables that differentiate these forest stands. Picea mariana stands were densely clustered together on the understorey DCA plot, suggesting a consistent understorey species composition (feather mosses and Erica- ceae), whereas Populus tremuloides stands had the most di- verse understorey species composition (ca. 30 species, mostly shrubs and herbs). Pinus banksiana stands had several charac- teristic species of reindeer lichens (Cladina spp.), but saplings and Pinus seedlings were rare. Although climatic variables showed large variation along the transect, the CCA results indicated that site conditions are more important in determining species composition and differentiating the stand types. Forest floor characteristics (litter and humus layer, woody debris, and drainage) appear to be among the most important site variables. Stands of Picea had significantly higher average carbon (C) densities in the combined litter and humus layer (43530 kg- C.ha-1) than either Populus (25 500 kg-C.ha-1) or Pinus (19400 kg-C.ha -1 ). The thick surface organic layer in lowland Picea stands plays an important role in regulating soil tem- perature and moisture, and organic-matter decomposition, which in turn affect the ecosystem C-dynamics. During forest succession after a stand-replacing disturbance (e.g. fires), tree biomass and surface organic layer thickness increase in all stand types as forests recover; however, woody biomass detri- tus first decreases and then increases after ca. 80 yr. Soil C densities show slight decrease with ages in Populus stands, but increase in other stand types. These results indicate the com- plex C-transfer processes among different components (tree biomass, detritus, forest floor, and soil) of boreal ecosystems at various stages of succession.

A multi-scale approach to mapping effective Leaf Area Index in Boreal Picea mariana stands using high spatial resolution CASI imagery

This study evaluates the performance of a simple geometric-optics reflectance model, used in combination with multi-spectral clustering, to map spatial patterns of effective Leaf Area Index ( L e ) within Boreal Picea mariana stands. Two metre resolution Compact Airborne Spectrographic Imager (CASI) images, acquired during the winter to minimize variability in understory reflectance, are used to map L e over BOREAS northern and southern Old Black Spruce tower flux sites. A combined multi-spectral clustering and ray-tracing approach is used to map open areas in each site at 2 m scale. A modified version of the Forest Light Interaction Model (FLIM) is then applied over canopy areas using 30 m scale red and near-infrared reflectance values derived from CASI images. Comparison of the combined FLIM and clustering approach (FLIMCLUS) with surface L e measurements in areas with overstory cover indicate a R 2 of 0.67 for the SSA-OBS site and 0.16 for the NSA-OBS site. The poor NSAOBS performance may be due to the low observed range of L e along the transect selected since additional measurements in a sparse canopy area closely match FLIM estimates. The relative standard error at both sites is under 10% and is close to the 5% precision error in surface L e estimates. Comparison of FLIMCLUS with FLIM and the simple ratio indicate substantial differences over open areas where the latter methods map-zero L e values. Further validation over other study sites, including surface data mapping edges between canopy and open areas, is proposed. The FLIM-CLUS L e maps may be useful for testing scale dependent assumptions within remote sensing algorithms and ecosystem flux models applied to the study sites and similar Picea mariana stands.

Carbon isotope composition of boreal plants: functional grouping of life forms.

We tested the hypothesis that life forms (trees, shrubs, forbs, and mosses; deciduous or evergreen) can be used to group plants with similar physiological characteristics. Carbon isotope ratios (δ13C) and carbon isotope discrimination (Δ) were used as functional characteristics because δ13C and Δ integrate information about CO2 and water fluxes, and so are useful in global change and scaling studies. We examined δ13C values of the dominant species in three boreal forest ecosystems: wet Picea mariana stands, mesic Populus tremuloides stands, and dry Pinus banksiana stands. Life form groups explained a significant fraction of the variation in leaf carbon isotope composition; seven life-form categories explained 50% of the variation in δ13C and 42% of the variation in Δ and 52% of the variance not due to intraspecific genetic differences (n=335). The life forms were ranked in the following order based on their values: evergreen trees<deciduous trees=evergreen and deciduous shrubs=evergreen forbs<deciduous forbs=mosses. This ranking of the life forms differed between deciduous (Populus) and evergreen (Pinus and Picea) ecosystems. Furthermore, life forms in the Populus ecosystem had higher discrimination values than life forms in the dry Pinus ecosystem; the Picea ecosystem had intermediate Δ values. These correlations between Δ and life form were related to differences in plant stature and leaf longevity. Shorter plants had lower Δ values than taller plants, resulting from reduced light intensity at lower levels in the forest. After height differences were accounted for, deciduous leaves had higher discrimination values than evergreen leaves, indicating that deciduous leaves maintained higher ratios of intracellular to ambient CO2 (c i/c a) than did evergreen leaves in a similar environment within these boreal ecosystems. We found the same pattern of carbon isotope discrimination in a year with above-average precipitation as in a year with below-average precipitation, indicating that environmental fluctuations did not affect the ranking of life forms. Furthermore, plants from sites near the northern and southern boundaries of the boreal forest had similar patterns of discrimination. We concluded that life forms are robust indicators of functional groups that are related to carbon and water fluxes within boreal ecosystems.

Responses ofPinus resinosain Newfoundland to wildfire

. Natural Pinus resinosa (red pine) stands in Newfoundland are restricted to 22 small, dry, nutrient-poor sites. A short wildfire cycle (15 - 30 yr) of both surface and crown fire regulates stand perimeters and is the main factor in regulating stand development. At the nucleus of current stands 200 yr occur, usually showing multiple fire scars. Stem char heights confirmed an increased flammability of the stand and tree mortality for fires moving in the upslope position, as well as for mixed Pinus resinosa-Picea mariana stands. All P. resinosa stands are severely nutrient-deficient. Leaf concentrations of N, P and K were below or near the reported critical values. Nutrient concentrations were highest three months after a surface fire, but dropped considerably one year later. A gradual increase to near post-fire levels is achieved four years after fire. Foliar nutrient concentrations were positively correlated with average width of the annual rings. Aspects of the ericaceous understory dynamics and its relation to P. resinosa regeneration are also discussed.