Interannual variation in competitive interactions from natural and anthropogenic disturbances in a temperate forest tree species: Implications for ecological interpretation

Competition is a major determinant of plant growth and is often used in studies of tree growth and species coexistence. However, these approaches are usually temporally static, i.e., assessed at a single point or period in time. While constantly changing forest conditions due to natural and human-induced disturbances potentially alter competition among individuals, static approaches cannot qualify the temporal variability of competitive interactions. Here we present a longitudinal analysis of competitive interactions among trees and discuss the implication of our results for ecological interpretation.Spatially-explicit tree growth data were obtained from 18 study plots (0.4 ha each) in sugar maple (Acer saccharum Marsh.) stands in Quebec, Canada. During the studied period (1980-2003), these stands had been disturbed by insect outbreaks (forest tent caterpillar, Malacosoma disstria Hubner) and by commercial partial harvest. We analyzed radial growth rates (outcome of competition) on an annual basis and as a function of tree biology (bole diameter, crown position), competition (above- and belowground competition from neighbours) and environmental conditions (light availability, harvest disturbance).Competitive interactions changed throughout the studied period. Canopy disturbance from partial harvest interacted with defoliators and influenced competition symmetry by favoring smaller trees.Competitive interactions seemed to have switched from below- to above-ground following canopy recovery after harvest. Release from competition due to partial harvest increase neighbourhood size (radius of effective competition) and enhanced the competitive pressure from larger individuals.The temporal variability in parameter estimates may be used for setting confidence intervals on competitive success (growth rates), thereby yielding a more robust basis for ecological interpretation. Our results also show that temporal variability in competitive interactions could contribute to the maintenance of high tree species diversity and structural complexity in some ecosystems by temporally altering species-specific responses to environmental change and disturbance.     

Spatial impacts of soil disturbance and residual overstory on density and growth of regenerating aspen

We examined spatial aspects of harvesting impacts on aspen regeneration at 25 sites in northern Minnesota. These sites had been clearcut or partially harvested 4–11 years ago. At each site, residual overstory, which was composed of trees other than aspen, soil disturbance, and tree regeneration were determined along transects leading away from skid trails into the neighboring stand. We characterized spatial extent of soil disturbance as soil strength using an Eijkelkamp soil cone penetrometer. Soil disturbance dropped off very quickly at the edge of skid trails, suggesting that the impact of harvesting traffic on areas adjacent to skid trails is minor. On skid trails, disturbance levels were higher on sites harvested in summer than on sites harvested in winter. Even after adjustment for differences in soil disturbance, stands harvested in winter had higher regeneration densities and greater aspen height growth than stands harvested in summer, suggesting that aspen regeneration was more sensitive to a given level of soil disturbance on summer-harvested sites versus on winter-harvested sites. Soil disturbance and residual overstory interactively reduced aspen regeneration densities and height growth, indicating that avoidance of soil disturbance is even more critical in partially harvested stands. Predictions based in the spatial patterns of impact found in this study indicated that harvesting conditions may have a great impact in future productivity of a site.     

Evaluating tree competition indices as predictors of basal area increment in western Montana forests

Fire hazard reduction treatments are commonly applied to mixed-species coniferous forests in western Montana, USA, to modify fuels structures and alter the competitive environments of individual trees. An improved understanding of how competition can be measured and how it conditions individual tree growth is needed for projecting the development of these forests, with and without treatment. Numerous studies have evaluated how competition affects tree growth and many indices have been developed to quantify the competition an individual tree experiences. These studies suggest that no single competition index or a single class of indices is universally superior; indices perform differently according to forest type and forest conditions. We chose several widely used distance-independent and distance-dependent competition indices, and also derived anisotropic distance-dependent indices from estimates of light interception by tree crowns. We evaluated the effectiveness of these competition measures for predicting basal area increment (BAI) of Pinus ponderosa, Pseudotsuga menziesii, and Larix occidentalis in western Montana. The best distance-dependent competition indices explained a larger proportion of growth variation than the best distance-independent indices (64% vs. 56%). This result indicates that competition is an important growth determinant in these forests and that competition varies locally, with variable tree densities and relatively complex stand structures creating heterogeneous neighborhood conditions. Competition indices derived from light interception models were only weakly correlated with other indices and performed poorly in terms of predicting tree growth. This result accords with previous observations that competition for light is not the primarily growth limitation for trees in the semi-arid conditions of western Montana. More sophisticated light availability models could be used to better assess variability in light interception and its marginal contribution to predictive accuracy of radial tree growth. Diameter and distance-dependent BAI models were developed for growth prediction at the species level and for all species combined.     

Evaluation of competition and light estimation indices for predicting diameter growth in mature boreal mixed forests

A series of conventional distance-independent and distance-dependent competition indices, a highly flexible distance-dependent crowding index, and two light resource estimation indices were compared to predict individual tree diameter growth of five species of mature trees from natural-origin boreal mixed forests. The crowding index was the superior index for most species and ecosites. However, distance-independent indices, such as basal area of competing trees, were also effective. Distance-dependent light estimation indices, which estimate the fraction of seasonal photosynthetically-active radiation available to each tree, ranked intermediate to low. Determining separate competition indices for each competitor species accounted for more variation than ignoring species or classifying by ecological groups. Species’ competitive ability ranked (most competitive to least): paper birch ≈ white spruce ≈> trembling aspen > lodgepole pine > balsam poplar. Stratification by ecosite further improved model performance. However, the overall impact of competition on mature trees in these forests appears to be small.     

Nutrient concentrations in roots, leaves and wood of seedling and mature sugar maple and American beech at two contrasting sites

Differences in sensitivity to soil conditions across tree species and developmental stage are important to predicting forest response to environmental change. This study was conducted to compare elemental concentrations in leaves, stems, and roots of (1) sugar maple (Acer saccharum Marsh.) seedlings vs. mature trees and (2) mature sugar maple vs. mature American beech (Fagus grandifolia Ehrh.) in two sites that differ in soil base saturation and pH. Both sites are located in Huntington Forest, NY, USA; one site (hereafter ‘H’) has higher soil pH and Ca, Mg, and Mn concentrations than the other site (hereafter ‘L’). Sugar maple growth at H (14.8 cm² year⁻¹ per tree) was much greater than at L (8.6 cm² year⁻¹ per tree), but the growth of beech was not different between the two sites. Leaves, roots, and stem wood of mature beech trees and sugar maple seedlings and mature trees were sampled for nutrient analysis. Foliar Ca, K, and Al concentrations were positively correlated with soil elements, but Mn concentrations were negatively correlated. Sugar maple differed more than beech between sites in foliar K and Mn concentrations. Root Mg and P concentrations reflected soil chemistry differences, in contrast to foliar concentrations of Mg and P, which were indistinguishable between the sites. In sugar maple, seedlings differed more than in mature trees in nutrient concentrations in roots, especially for Mg and Mn. Although beech was not as responsive to nutrient availability as sugar maple in foliar and root nutrient concentrations, Ca and Mg concentrations in beech wood were higher in H (52% higher for Ca and 68% higher for Mg), while sugar maple did not differ between sites. Sugar maple regeneration failure on acidic soils in the same region is consistent with our finding that sugar maple seedlings were very sensitive to nutrient availability. This sensitivity could ultimately contribute to the replacement of sugar maple by American beech in regions of low pH and base cations if base cation leaching by anthropogenic deposition and tree harvesting continues.     

Long-term assessment of soil physicochemical properties and seedlings establishment after skidding operations in mountainous mixed hardwoods

One of the basic requirements for sustainable forest management in mixed broadleaved stands is to provide suitable conditions for natural regeneration of trees. These conditions include, but are not limited to, the frequency, quality and the composition of seedling species. Compacted soil layers caused by forest machinery traffic are the most common problem affecting seedling establishment and growth after skidding operations. In this study, we evaluated the frequency, quality, diversity of seedling species and physicochemical properties of soil on 10-, 20- and 30-year-old abandoned skid trails. Further comparison to the values of the mentioned above parameters in control areas allowed for the evaluation in terms of natural recovery processes, under varying traffic intensity and the longitudinal slope of the skid trails in a mixed broadleaved forest. Results showed that there is a significant positive trend of recovery for soil physicochemical properties and ecological characteristics (density and quality) of seedling growing on the skid trails. The time required to recover soil properties and ecological attributes of seedlings increased with increasing traffic intensity and slope of the skid trail. Our results showed that it takes 20 and 30 years to fully recover the chemical and physical properties of the soil, respectively. On the 30-year-old skid trails, density and species diversity indices were fully recovered, but the quality of the seedlings was not restored to the control conditions. The results suggest the importance of a proper planning of the skid trail network, avoiding in particular skid trails with a slope gradient higher than 20%.

     

Impacts of selective logging on above-ground forest biomass in the Monts de Cristal in Gabon

Selective logging is an important socio-economic activity in the Congo Basin but one with associated environmental costs, some of which are avoidable through the use of reduced-impact logging (RIL) practices. With increased global concerns about biodiversity losses and emissions of carbon from forest in the region, more information is needed about the effects of logging on forest structure, composition, and carbon balance. We assessed the consequences of low-intensity RIL on above-ground biomass and tree species richness in a 50 ha area in northwestern Gabon. We assessed logging impacts principally in 10 randomly located 1-ha plots in which all trees ?10 cm dbh were measured, identified to species, marked, and tagged prior to harvesting. After logging, damage to these trees was recorded as being due to felling or skidding (i.e., log yarding) and skid trails were mapped in the entire 50-ha study area. Allometric equations based on tree diameter and wood density were used to transform tree diameter into biomass.Logging was light with only 0.82 trees (8.11 m³) per hectare extracted. For each tree felled, an average of 11 trees ?10 cm dbh suffered crown, bole, or root damage. Skid trails covered 2.8% of the soil surface and skidding logs to the roadside caused damage to an average of 15.6 trees ?10 cm dbh per hectare. No effect of logging was observed on tree species richness and pre-logging above-ground forest biomass (420.4 Mg ha⁻¹) declined by only 8.1% (34.2 Mg ha⁻¹). We conclude from these data that with harvest planning, worker training in RIL techniques, and low logging intensities, substantial carbon stocks and tree species richness were retained in this selectively logged forest in Gabon.     

Individual-level analysis of damage to residual trees after single-tree selection harvesting in northern Japanese mixedwood stands

Quantifying tree damage and mortality caused by single-tree selection harvesting is critical to understanding postharvest forest dynamics and management. In this study, we quantified the effects of tree size and species and the distance from residual trees to felled trees and skid trails on damage to residual trees and mortality in mixed coniferous–broadleaved stands of Hokkaido, northern Japan. Among the 4,961 trees that we studied, 373 (7.5 %) were damaged, and 148 of these trees (3.0 %) died during or immediately after logging. Hierarchical Bayesian modeling showed that the risk of damage to residual trees increased with increasing size of the felled trees and with increasing proximity to felled trees and skid trails. Smaller residual trees had the greatest risk of damage. Species differed in their susceptibility to damage; Abies sachalinensis (Fr. Schm.) Masters and Picea jezoensis (Sieb. et Zucc.) Carr. were the most susceptible species in our sample plots. Smaller damaged trees had the highest risk of mortality. The damaged trees that did not die at the time of logging had a higher risk of postharvest mortality than undamaged trees. Our results indicate that, to minimize logging-induced damage and mortality: (1) the spatial arrangement of skid trails should be optimized and fixed, (2) the risk of skidding damage should be considered before marking the trees to be harvested and choosing the felling direction, and (3) logging operators should be instructed to avoid damaging small trees, especially those of the species that are most susceptible to damage.     

Individual-tree diameter growth model for sugar maple trees in uneven-aged northern hardwood stands under selection system

An individual-tree diameter model was developed for sugar maple (Acer saccharum Marsh.) in northern hardwood stands managed under selection system. We fitted long-term remeasurement data to a linear mixed model to account for the temporal autocorrelation of the remeasurements. The model was evaluated using independent data from two physiographic regions and representing a range of tree diameter classes, residual basal areas and years since cut. We compared our model to several individual-tree models based on data from stands with varied management histories. Several competition indices were also tested for an improvement in model fitting and prediction. Our model had lower bias and prediction error when compared to two previous models, as it better accounted for the increased diameter growth that occurred in trees from appropriately managed stands. The addition of a tree-specific competition index failed to improve model fit and predictive ability over stand-level basal area.     

Soil disturbance and post-logging forest recovery on bulldozer paths in Sabah, Malaysia

We examined the extent of soil disturbance associated with bulldozer yarding and the regrowth of woody vegetation on bulldozer paths (skid trails) in selectively logged dipterocarp forest. In an area logged in 1993, using conventional, i.e., uncontrolled, harvesting methods, about 17% of the area was covered by roads and skid trails. In contrast, in a 450-ha experimental area where reduced-impact logging guidelines were implemented, 6% of the area was similarly disturbed. Skid trails in the reduced-impact logging areas were less severely disturbed than those in conventional logging areas; the proportion of skid trails with subsoil disturbance was less than half that in conventional logging areas. Four years after logging, woody plant recovery on skid trails was greater in areas logged by reduced-impact than by conventional methods. Skid trails where topsoil had been bladed off had less woody vegetation than skid trails with intact topsoil. In a chronosequence of logging areas (3, 6, and 18 years after logging), species richness and stem densities of woody plants (>1 m tall, <5 cm dbh) were lower on skid trail tracks than on skid trail edges or in adjacent forest. Both richness and density increased with time since logging, but even 18 years after logging, abandoned skid trails were impoverished in small woody stems compared with adjacent forest. Minimizing soil and stand disturbance during logging appears to allow a more rapid recovery of vegetation on bulldozed soils, but the long-term fate of trees growing on compacted soils remains uncertain.     

Growth of young Pinus ponderosa and Pinus contorta on compacted soil in central Washington

Growth in height, diameter, and volume was measured on 9- to 18-year-old ponderosa pine (Pinus ponderosa) and 10- to 13-year-old lodgepole pine (Pinus contorta) trees growing on or near compacted skid trails in the Yakima Indian Reservation in south-central Washington. Soil bulk density of the 0- to 30.5-cm deep layer was measured with a single-probe nuclear densimeter on two sides of each sample tree and in adjacent undisturbed soil. On three ponderos pine sites logged 23 years before the study, average bulk density on skid trails was 15% greater than on adjacent undisturbed soil. On a lodgepole pine site logged 14 years before the study, soil on skid trails averaged 28% greater bulk density than undisturbed soil.Total growth of ponderosa pine and the last 5 years of growth were significantly related (P = 0.07) by regression analysis to age of trees, site index, basal area of the adjacent overstory, and the percentage of increase in soil bulk density. At the mean increase in soil bulk density, total height, diameter, and volume growth were reduced 5%, 8%, and 20%, respectively.Total growth of lodgepole pine and the last 5 years of height, diameter, and volume growth were significantly related to tree age and the percentage of soil organic matter. Increase in soil bulk density was not significantly related to growth of this species.     

Gap-, stand-, and landscape-scale factors contribute to poor sugar maple regeneration after timber harvest

Natural regeneration in canopy gaps is a key process affecting long-term dynamics of many forests, including northern hardwood forests. The density and composition of regenerating trees are often highly variable, reflecting sensitivity to a suite of driving factors operating at different scales (e.g., harvest gap to regional landscape), including production of seeds, physical characteristics of gaps and stands, competition with non-tree vegetation, and browsing by animals. Multivariate analyses over broad geographic areas provide insights into the relative effects of these factors and permit exploration of spatial patterns in regeneration. We examined the effects of gap-, stand-, and landscape-scale factors on densities of tree seedlings (<1 m tall) and saplings (1-2 m tall) in 59 selection-harvested northern hardwood stands located across a 4500 km² region of Michigan's Upper Peninsula. We used Bayesian multilevel modeling to account for the hierarchical structure of the data and assess uncertainty in parameter estimates. Sugar maple (Acer saccharum) saplings were absent from 61% of 154 m² plots centered in harvest gaps (n = 347) despite its high shade tolerance and overstory dominance, but densities were high in other gaps. Densities of sugar maple seedlings and/or saplings were negatively associated with a combination of greater stand-scale densities of white-tailed deer (Odocoileus virginianus), greater gap-scale cover of non-tree vegetation, and lower gap-scale light availability, with deer density having the greatest effect. Densities of unpalatable and commercially less valuable ironwood (Ostrya virginiana), the second most common regeneration species, were positively related to gap-scale seed-production potential but were unrelated to factors affecting sugar maple. Ironwood tended to replace sugar maple saplings in areas with high deer density. At the landscape scale, densities of sugar maple seedlings and saplings decreased with decreasing latitude and snow depth and increasing winter deer densities. These inverse spatial patterns suggest that deer herbivory can lead to landscape-scale variation in regeneration success. However, the spatial distribution of habitat types (a proxy for soil moisture and nutrient conditions) confound this observation, with higher densities of sugar maple generally located on stands with less nutrient-rich habitat types. Results demonstrate that combinations of factors operating at different scales, and with different relative magnitudes of impact, contribute to high variation in regeneration composition and density following timber harvest. Selection silvicultural practices, as currently applied, do not ensure regeneration of desirable species; practices might require modifications in general (e.g., increasing gap size) and to match them to regionally varying factors like deer density.     

Individual tree growth response to variable-density thinning in coastal Pacific Northwest forests

We examined 5-year basal area growth of nearly 2600 trees in stem-mapped plots at five locations differing in site characteristics, species composition, and management history on the Olympic Peninsula in Western Washington, USA. Our objectives were to determine if internal edges, the boundaries within the stand between components of the variable-density thinning, influenced individual tree growth, and whether incorporation of individual tree local competition indices in growth prediction models could account for treatment and edge effects. Treatment significantly affected tree growth at all sites, with trees in the thinned matrix displaying on average over 25% greater basal area growth than trees in unthinned patches. Proximity to canopy gaps created as part of the variable-density thinning increased basal area growth of trees in the thinned matrix by nearly 11%. In addition, growth of trees close to skid trails was 11% greater than trees located away from the trails. Past thinning history, and its effect on initial stocking rate, appeared to affect the magnitude of the edge effects. Blocks that had received earlier commercial thinnings, and thus had lower stocking at the onset of the study, displayed lower growth responses than previously unthinned blocks. Including local competition indices in the models generally reduced growth prediction error; however, the indices examined did not fully account for treatment or edge effects. Our results suggest that not accounting for internal edges in spatially complex stands could result in errors in projected growth of trees, although these edge effects are highly variable. Failure to account for the effects of internal edges could affect not just estimates of future stand yield, but also projections of future stand structure.     

The effect of competition on individual tree basal area growth in mature stands of Pinus cooperi Blanco in Durango (Mexico)

In this paper, we evaluated how well-selected distance-dependent and distance-independent competition indices explain individual tree basal area growth of trees, growing in mature and even-aged stands of Pinus cooperi Blanco. A total of 18 competition measures were analyzed of which six do not need tree location (distance-independent) and 12 that utilize tree location (distance-dependent). The competition situation of a stand and/or an individual tree was studied using 11 different competitor selection methods. The mean square error reduction relative to no-competition was used to judge the performance of each competition index. It was found that the best distance-independent competition indices performed as well as the best distance-dependent competition indices. It was concluded that the BALMOD-index would be a good competition index to be incorporated into further individual tree basal area growth models for the study area.

Species composition and dynamics in two hardwood stands in Vermont: a disturbance history

Stand composition and structure utilizing stem analysis was studied in two hardwood stands in Vermont. In a mixed hardwood stand with some white pine and hemlock, a major entry of new trees in the main canopy seems associated with harvesting coincident with land exchange. More recent partial cuttings have promoted establishment of new seedlings or development of suppressed advanced-growth shade-tolerant beech, (Fagus grandifolia, Ehrh.), hophornbeam (Ostrya virginiana, (Mill) K. Koch), and striped maple (Acer pensylvanicum, L.). Very few sugar maple and red oak seedlings and saplings are present.

In a northern hardwood stand some red spruce (Picea rubens, Sarg.), that were 240 to 306 years old, became established before any known harvest, and exhibited release following harvests of the mid-1800s. This major harvest, coupled with the differential growth between spruce and hardwoods, and seed/seedling availability, resulted in a major change in stand composition. Trees now in the main canopy of sampled stands appear to have either been released or newly established following various harvests. Harvests have been of such frequency that natural disturbances seem insignificant. Many of the competitive understory species have become abundant following harvests of the 1960s and 1980s and may have been present as advanced growth and responded to the release. Following the harvest of 1981–1982, abundant yellow birch became established on skid trails. Elsewhere in the stand, yellow birch seedlings and saplings are only in great abundance in areas that were possibly sizable gaps following earlier harvesting.

The dynamics of tree entry and growth in gaps of small or large size probably occur in a similar way in many other stands of the region. Though the sampling of this study is limited, there is no suggestion of continuous tree establishment at any particular location, the new age classes seem associated with either a gap or stand replacing disturbance attributed to harvesting.     

A model bridging distance-dependent and distance-independent tree models to simulate the growth of mixed forests

? It is widely believed that distance-independent tree models fail to take into account the complexity of mixed stands due to the fact that spatial structure often has a greater impact on growth and dynamics in mixed stands than in pure stands. On the other hand, distance-dependent tree models are difficult to use because they require a map of the stand, which is not only very costly but also impracticable in a routine management context.

? This paper reports the development of a model bridging distance-dependent and distanceindependent tree models, and that is designed to simulate the growth of a mixed forest. The model used distributions of the number of neighbours to reconstruct tree neighbourhoods and compute the competition indices needed as inputs to the growth model.

? Data were collected from a mixed forest of sessile oak and Scots pine in central France. The study showed that local competition indices explained a significant proportion of growth variability and that intraspecific competition was greater than interspecific competition. The model based on neighbourhood distributions gave consistent predictions compared to a distance-dependent model.

? This type of model could be used instead of distance-dependent models in management contexts.

     

Spatiotemporally interactive growth dynamics in selected South African forests: Edaphoclimatic environment, crowding and climate effects

Stand-level tree diameter growth patterns were explored for evergreen moist forests in the southern Cape, South Africa. Results of standard multiple regression analyses, involving 934 permanent sample plots with data spanning a 10-year interval, revealed that stand-level increment of canopy species in the canopy layer (>30 cm dbh) was significantly determined by inherent species-specific growth capacities (species composition of the stand), water availability, forest matrix crowding and tree condition impairment (age-related manifestations of reduced vitality indicated by signs of crown die-back, damage and stem rot). In contrast, stand-level increment of trees of canopy species in the subcanopy layer (10-20 cm dbh) was prominently shaped by light availability, as mainly determined by the degree of canopy-level disturbance (mortality rate of trees >30 cm dbh), crowding (canopy-level overhead and forest matrix crowding) and proximity to conspecific adults (within 6-8 m). In addition to species-inherent and resource factors, considerable variation in stand-level growth resulted from site-climate interactions. For 507 of the permanent sample plots, increment data was available for two consecutive 10-year intervals; permitting the analysis of spatiotemporal interactions of growth patterns (repeated measures ANOVA). In the Knysna forests higher canopy-level increment rates were associated with the moister southerly facing slope sites in comparison with the drier northerly facing and ridge sites during the first increment period. During the second increment period, increment rates on the drier, but better illuminated sites had increased disproportionately. In contrast, in the Tsitsikamma forests, higher increment rates during the second increment period were encountered on moister flat bottomland sites (with extended periods of subsoil wetness) than on the comparatively drier southerly facing slope sites (increment period × site-based water availability × forests interaction). In both forests relatively higher growth performance of subcanopy-level trees during the second increment period was associated with stands experiencing conditions of enhanced light availability. Atmospheric temperatures were higher during the second increment period (mean periodic T_max: + 0.64 °C). The detected spatiotemporal interactions were interpreted as site × climate interactions where site-related conditions of favourable light or water availability resulted in enhanced temperature-linked growth responses during the second increment period. A metabolic performance trade-off model provided a framework for the interpretation of these complex site-climate interactions by placing the patterns of forest growth into an ecophysiological explanatory context.     

Physiological, morphological and allocational plasticity in understory deciduous trees: importance of plant size and light availability

In a 4-year study, we investigated changes in leaf physiology, crown morphology and whole-tree biomass allocation in seedlings and saplings of shade-tolerant sugar maple (Acer saccharum Marsh.) and intermediate shade-tolerant yellow birch (Betula alleghaniensis Britt.) growing in natural understory light (0.5 to 35% of full sunlight) or in understory light reduced by 50% with shade nets to simulate the effect of gap closure. Leaf physiological parameters were mainly influenced by the light gradient, whereas crown morphological and whole-tree allocational parameters were mainly influenced by tree size. No single physiological, morphological or allocational trait was identified that could explain the difference in shade tolerance between the species. Yellow birch had higher growth rates, biomass allocation to branches and leaf physiological plasticity and lower crown morphological plasticity in unmodified understory light than sugar maple. Sugar maple did not display significant physiological plasticity, but showed variation with tree size in both crown morphology and whole-tree biomass allocation. When sugar maple was small, a greater proportion of whole-tree biomass was allocated to roots. However, physiological differences between the species decreased with decreasing light and most morphological and allocational differences tended to disappear with increasing tree size, suggesting that many species differences in shade-tolerance are expressed mainly during the seedling stage. Understory trees of both species survived for 4 years under shade nets, possibly because of higher plasticity when small and the use of stored reserves when taller.     

Assessment of sugar maple tree growth in relation to the partitioning of elements in xylem along a soil acidity gradient

Partitioning of elements in tree xylem is being increasingly studied, as it suggests that elements are potentially mobile within the xylem long after their uptake. A recent study revealed that only the most mobile xylem fraction (water-soluble) of base cations (calcium [Ca], magnesium [Mg], and potassium [K]) increased at higher soil acidity, while the two mobile fractions (water- and acid-soluble) of acidic metals—potentially phytotoxic aluminium (Al), cadmium (Cd) and manganese (Mn)—were significantly enhanced on very acid soils. The current paper presents an investigation of soil-wood chemistry relationships with basal area tree growth. It was hypothesized that the growth of sugar maple would be reduced by low base cation and high acidic metal concentrations in the xylem mobile fractions. Sugar maple trees (n = 55) from six watersheds in southern Quebec, Canada were analysed by sequential chemical extractions for the water-soluble, acid-soluble and residual fractions of base cations (Ca, K, Mg) and acidic metals (Al, Cd, Mn) in xylem. Generally, tree growth was positively correlated to concentrations of base cations in wood (ρ = 0.27-0.50) and soil (ρ = 0.41-0.67), and negatively correlated to concentrations of acidic metals in wood (ρ = −0.33 to −0.52) and soil (ρ = −0.67). However, these relations differed depending on the element fraction considered. Water- and acid-soluble xylem concentrations of base cations and Al were among the best predictors of growth trends (R² = 0.46-0.51). The relationship between acidic metals and tree growth is further discussed.     

Estimating the maple syrup production potential of American forests: an enhanced estimate that accounts for density and accessibility of tappable maple trees

This paper examines the maple syrup production potential of American forests by analyzing Forest Inventory & Analysis (FIA) data provided by the US Forest Service on the resource of sugar maple (Acer saccharum Marsh.) and red maple (Acer rubrum L.) trees in twenty states. The analysis is based on tree species and size (diameter at breast height, or dbh), ownership category, jurisdiction, the density of maple trees in a stand, and the distance of the stand to an access road. Although there are over 2 billion sugar and red maple trees of tappable size growing in US forests, when narrowed down according to the attributes of an optimal ‘sugarbush’, there are 100 million potential taps from sugar maples alone and 286 million potential taps with sugar and red maples combined. Overall, 45 % of the tappable-size maple trees are found in stands whose density is not high enough to support commercial sap extraction whereas only 6 % are found in stands that are at least 1.6 km from an access road. The ten states with commercial maple syrup industries have a much higher percentage of their maple trees occurring in stands of optimal density and also contain a higher percentage of sugar maple than red maple trees. States that are utilizing the highest percentage of their potential sugarbushes include Vermont and Maine, whereas states that have significant room for expansion include Michigan, New York, and Pennsylvania.