Influence of light availability on growth, leaf morphology and plant architecture of beech ( Fagus sylvatica L.), maple ( Acer pseudoplatanus L.) and ash ( Fraxinus excelsior L.) saplings

In a field study, we measured saplings of beech, ash and maple growing in a fairly even-aged mixed-species thicket established by natural regeneration beneath a patchy shelterwood canopy with 3–60% of above canopy radiation reaching the saplings. Under low light conditions, maple and ash showed a slight lead in recent annual length increment compared with beech. With increasing light, ash and maple constantly gained superiority in length increment, whereas beech approached an asymptotic value above 35% light. A suite of architectural and leaf morphological attributes indicated a more pronounced ability of beech to adapt to shade than ash and maple. Beech displayed its leaves along the entire tree height (with a concentration in the middle crown), yielding a higher live crown ratio than ash and maple. It allocated biomass preferentially to radial growth which resulted in low height to diameter ratios, and expressed marked plagiotropic growth in shade indicating a horizontal light-foraging strategy. In addition, beech exhibited the highest specific leaf area, a greater total leaf area per unit tree height, a slightly greater leaf area index, and a greater plasticity to light in total leaf area. Ash and maple presented a “gap species” growth strategy, characterized by a marked and constant response in growth rates to increasing light and an inability to strongly reduce their growth rates in deep shade. In shade, they showed some plasticity in displaying most of their leaf area at the top of the crown to minimize self-shading and to enhance light interception. Through this, particularly, maple developed an “umbrella” like crown. These species-specific responses may be used for controlling the development of mixed-species regeneration in shelterwood systems.     

Growth and mortality in underplanted tree seedlings in response to variations in canopy closure of Norway spruce stands

Six broadleaved tree species and Picea abies (L.) Karst. wereplanted under spruce plantations of varying densities, in Sweden.Treatments included control (994 stems ha^–1), dense (538stems ha^–1), sparse (294 stems ha^–1) and gap (0stems ha^–1) overstory treatments. There was an increasein height and diameter growth from control to sparse overstorytreatment of all underplanted tree species except for ash (Fraxinusexcelsior L.) and Norway maple (Acer platanoides L.). Site conditionsmay have hampered the growth of these species, as well as wildcherry (Prunus avium L.). Both oak (Quercus robur L.) and sprucehad greater growth in the gap treatment, relative to the othertreatments. Insecticide application did not influence seedlinggrowth or survival. The performance of beech (Fagus sylvaticaL.), lime (Tilia cordata Mill.), spruce and oak was consistentwith shade tolerance ranking. Beech and lime had a very highsurvival rate, even under the densest canopy. The growth andmortality of ash, maple and wild cherry differed significantlyfrom what was expected. This experiment demonstrated significantvariation in interspecific growth and mortality between sevenplanted tree species in relation to canopy density. Correctsite and species selection is crucial when underplanting inshelterwood systems.     

Development of canopy stratification during early succession in northern hardwoods

Canopy development on a 6-year-old strip cut was analyzed by measuring the heights to terminal buds and bud scale scars of the tallest individuals of each species present on 50 plots of radius 6 m. Phenology of height growth was monitored during the following growing season. Pin cherry (Prunus pensylvania L.), an intolerant short-lived tree, had the fastest growth rate and was on the average the tallest species from the second to the sixth year of regrowth. Although advance regeneration of sugar maple (Acer saccharum Marsh.) and beech (Fagus grandifoloa Ehrh.) were the tallest trees during the first growing season, their slower growth rate insured that they would not keep up with the pin cherry. Trembling aspen (Populus tremuloides Michx.), striped maple (Acer pensylvanicum L.) and yellow birch (Betula alleghaniensis Britt.) occupied an intermediate position in the canopy by the end of the sixth growing season, and showed relatively greater annual height increment than beech or sugar maple. Height growth phenology differed slightly for each species. Beech, ash (Fraxinus americana L.) and sugar maple commenced growth early, grew rapidly and set buds all by 1 August (beech by 15 June). Yellow birch, pin cherry and trembling aspen started growing as early as the others, grew more slowly at first but then grew for a longer period of time. Striped maple seemed to be somewhat intermediate. Growth phenology and growth rate are related to the tolerance and growth form type (e.g. determinate or indeterminate) of the species. The most tolerant species tend to be determinate in growth form, have slower growth rates and complete height growth earlier. The intolerant species tend to be indeterminate, have a faster growth rate and continue to grow for a longer period. These may be mechanisms by which many species can grow together and avoid adverse effects such as suppression.     

Seasonal variation in biomass and carbohydrate partitioning of understory sugar maple (Acer saccharum) and yellow birch (Betula alleghaniensis) seedlings

We investigated seasonal patterns of biomass and carbohydrate partitioning in relation to shoot growth phenology in two age classes of sugar maple (Acer saccharum Marsh.) and yellow birch (Betula alleghaniensis Britt.) seedlings growing in the understory of a partially harvested forest. The high root:shoot biomass ratio and carbohydrate concentration of sugar maple are characteristic of species with truncated growth patterns (i.e., cessation of aboveground shoot growth early in the growing season), a conservative growth strategy and high shade tolerance. The low root:shoot biomass ratio and carbohydrate concentration of yellow birch are characteristic of species with continuous growth patterns, an opportunistic growth strategy and low shade tolerance. In both species, starch represented up to 95% of total nonstructural carbohydrates and was mainly found in the roots. Contrary to our hypothesis, interspecific differences in shoot growth phenology (i.e., continuous versus truncated) did not result in differences in seasonal patterns of carbohydrate partitioning. Our results help explain the niche differentiation between sugar maple and yellow birch in temperate, deciduous understory forests.     

Effects of root trenching of overstorey Norway spruce (Piceaabies) on growth and biomass of underplanted beech (Fagussylvatica) and Douglas fir (Pseudotsugamenziesii) saplings

In Central Europe, the conversion of pure Norway spruce stands (Picea abies [L.] Karst.) into mixed stands with beech (Fagus silvatica L.) and other species like e.g. Douglas fir (Pseudotsuga menziesii [Mirb.] Franco) is accomplished mainly by underplanting of seedlings beneath the canopy of overstorey spruce trees after partial cutting treatments what means exposure to shade and below-ground root competition by the overstorey to the seedlings. Particularly about the second factor, our knowledge is limited. Therefore, we carried out a below-ground competition exclusion experiment by root trenching and investigated the effects on soil resources, growth, and biomass partitioning of underplanted beech and Douglas fir saplings under target diameter and strip cutting treatments. The exclusion of overstorey root competition by trenching increased the soil water potential in the second year that had a fairly dry growing season and led to significantly higher foliar concentrations of most nutrients, particularly in Douglas fir, indicating an amended nutrient supply. Both improvements were accompanied by an increase in length and diameter increment of the underplanted saplings, appearing in both species only after having surpassed a species-specific threshold light value (Douglas fir 16% of above canopy radiation, beech 22%). We also found significant interactions between trenching and light for specific fine root length and further biomass and morphological parameters. Judged by the much steeper increase in height and diameter growth with increasing light after release from below-ground competition, Douglas fir saplings appeared to be more sensitive to root competition than beech saplings what conforms to older findings for beech. According to our results, a strip cutting seems to be more appropriate than a target diameter cutting treatment to replace a pure spruce stand by a mixed stand with beech and Douglas fir.     

Covariation between tree size and shade tolerance modulates mixed-forest productivity

Key message

In tree communities, tree size inequality reduces productivity and interacts with tree shade tolerance to modulate stand productivity, with a higher productivity in stands where shade-intolerant species dominate shade-tolerant species in size.

Context

Positive diversity–productivity relationships have been reported in different plant communities, including tree communities. These effects may be strongly related to both structural diversity and functional diversity, but also to their interactions if there is a non-random distribution of species functional characteristics among canopy layers.

Aims

We explore the relative effects on forest productivity of tree species diversity, tree size inequality, and species shade tolerance diversity, as well as the effect of the distribution of tree shade tolerance in the canopy.

Methods

We used 11,054 mixed-species forest plots from the French Forest Inventory (IGN) distributed throughout France (2006–2011). We analyzed the effects of species richness, shade tolerance diversity, and height inequality on forest plot productivity, represented by basal area annual increment over a period of 5 years, while controlling for first-order structure characteristics (basal area and quadratic mean diameter) and environmental factors (soil water budget and sum of growing degree days). Using the covariance between tree height and shade tolerance in mixed species canopies, we also explored the effect of the distribution of species’ shade tolerance among canopy layers.

Results

The results showed a positive effect of species richness (effect size, 0.02) and a negative effect of height inequality (??0.05) on mixed-forest productivity. We also showed that a negative covariance between shade tolerance and height (e.g., higher proportion of shade-tolerant species in lower height classes) increased productivity (0.01). Shade tolerance diversity did not affect productivity.

Conclusion

In tree communities, as shown previously in monospecific forest stands, tree size inequality reduces productivity. This effect is modulated by the distribution of shade tolerance among canopy layers. Previous studies on species diversity effect have generally overlooked the importance of the size structure and the size hierarchy of functional characteristics. These effects are, however, crucial and deserve to be explored in greater detail.

     

Survival and growth of upland oak and co-occurring competitor seedlings following single and repeated prescribed fires

Studies within and outside the U.S. indicate recurring oak (Quercus spp.) regeneration problems. In deciduous forests of the eastern U.S., a prevailing explanation for this trend is fire suppression leading to high competitor abundance and low understory light. In response, prescribed fire is increasingly used as a management tool to remedy these conditions and encourage future oak establishment and growth. Within eastern Kentucky, we implemented single and repeated (3×) prescribed fires over a 6-yr period (2002–2007). Pre- and post-burn, we quantified canopy cover and oak seedling survival and growth compared to other woody seedlings deemed potential competitors, primarily red maple (Acer rubrum L.) and sassafras (Sassafras albidum (Nutt.) Nees.). Burning temporarily decreased canopy cover 3–10%, but cover rebounded the subsequent growing season. Repeated burning ultimately produced canopy cover about 6% lower than sites unburned and burned once, suggesting a cumulative effect on understory light. Red maple exhibited low survival (∼40%) following single and repeated burns, but growth remained similar to unburned seedlings. Burning had little impact on sassafras survival and led to total height and basal diameters 2× greater than unburned seedlings. A single burn had no impact on red oak (Erythrobalanus spp.) survival and increased height and basal diameters 25–30%, but this positive growth response was driven by seedlings on several plots which experienced high burn temperatures and consequently high overstory mortality. White oaks (Leucobalanus spp.), however, exhibited twice as high mortality compared to those unburned, with no change in growth parameters. Repeated burning negatively impacted survival and growth of both oak groups compared to unburned seedlings. With both burn regimes, oaks with smaller pre-burn basal diameters exhibited the lowest post-burn survival. Thus, despite the ability of prescribed burns to temporarily increase understory light and reduce red maple survival, neither single or repeated burns placed oaks in an improved competitive position. These findings result from a combination of highly variable yet interdependent factors including the (1) life history traits of oaks compared to their co-occurring competitors, (2) pre-burn stature of pre-existing oak seedlings, and (3) variability in fire temperature and effects on understory light.     

Spatial coexistence of American beech and sugar maple regeneration in post-harvest northern hardwood forests

Context

Fine scale regeneration patterns of coexistent species are influenced by regeneration mechanisms and microsite requirements. Spatial patterns may be either disjunct or overlapping, which will determine competitive effects and microsite dominance, and future forest composition.

Aims

Using American beech (Fagus grandifolia Ehrh.) and sugar maple (Acer saccharum Marshall) as an example, three hypotheses were tested: (1) random beech spatial patterns, (2) clumped spatial patterns of small sugar maple seedlings, and (3) disjunct beech and sugar maple patterns.

Methods

Individual stems were sampled in a contiguous grid of 1-m² quadrats across a 576-m² area at three sites. Densities were separated into three height classes (≤30 cm, 30–90 cm, and?>?90 cm, ≤4 cm diameter at breast height). Spatial statistics and regression were used to analyze spatial patterns and correlations.

Results

Beech and seedling sugar maple patterns were patchy, rejecting the first and not rejecting the second hypotheses. Hypothesis three was rejected because patches of the two species overlapped with advance regeneration beech overtopping sugar maple.

Conclusion

Patchy patterns of advance regeneration beech and post-harvest sugar maple establishment suggest spatiotemporal niche partitioning. Beech had a competitive height advantage following harvest, but sugar maple still occurred in beech-free patches and beneath overtopping beech at a fine scale. Self-replacing beech patterns will ensure the species will continue dominance unless a selective chemical or manual treatment is applied that removes beech and releases sugar maple.     

Photosynthesis and biomass allocation of beech (Fagus crenata) and dwarf-bamboo (Sasa kurilensis) in response to contrasting light regimes in a Japan Sea-Type beech forest

Regeneration of beech (Fagus crenata) forests depends on the formation of canopy gaps. However, in Japan Sea-type beech forests, a dwarf bamboo (Sasa kurilensis) conspicuously occupies sunny gaps. Therefore,F. crenata seedlings must escape the severe interference ofS. kurilensis in the gaps and persist beneath a closed canopy of the beech forest. We hypothesized that the growth ofF. crenata seedlings in the understory would be favored by their being more plastic thanS. kurilensis in photosynthetic and morphological traits, which would support the matter production ofF. crenata seedlings in a wide range of light availabilities. To examine this hypothesis, the photosynthetic-light response of individual leaves and the biomass allocation in aboveground parts (i.e., the culm/foliage ratio) were surveyed at sites with contrasting light availabilities in a Japan Sea-type beech forest in central Japan. InF. crenata, photosynthetic light utilization efficiency at relatively low light was greater, and the dark respiration rate was smaller in the leaves of seedlings (10 cm in height) beneath the closed canopy than in the leaves of saplings at the sunny forest edge. The culm/foliage (C/F) ratio of theF. crenata seedlings at the shady site was small, suggesting effective matter-production beneath the beech canopy. On the other hand,S. kurilensis both in the gap and beneath the beech canopy showed low plasticity in photosynthesis and the culm/foliage ratio. Because the shoot density ofS. kurilensis was smaller beneath the beech canopy than in the gap, the light availability at the bottom of theS. kurilensis layer was greater beneath the beech canopy. These results suggest that the photosynthetic productivity of theF. crenata seedlings would be enough for the seedlings to survive in the understory with a low density ofS. kurilensis shoots beneath the closed beech canopy.     

Influence of browsing damage and overstory cover on regeneration of American beech and sugar maple nine years following understory herbicide release in central Maine

Northern hardwood stands, notably those with American beech (Fagus grandifolia Ehrh), sugar maple (Acer saccharum Marsh.), and yellow birch (Betula alleghaniensis Britton), are abundant across the forested landscapes of northeastern USA and southeastern Canada. Recent studies have reported an increasing dominance of American beech in the understory and midstory of these forests. Beech is a commercially less desirable tree species due to its association with beech-bark disease, and because it commonly interferes with the regeneration of other more desirable tree species. We examined hardwood regeneration characteristics nine years after application of a 3 × 4 factorial combination of glyphosate herbicide (0.56, 1.12, and 1.68 kg ha^?1) and surfactant concentrations (0.0, 0.25, 0.5, and 1.0% v v^?1) to release sugar maple regeneration from beech-dominated understories using three stands that received shelterwood seed cutting in central Maine. Measurements nine years after treatment showed that glyphosate rate increased both absolute (AD) and relative density (RD) of sugar maple regeneration, but not its height (HT). In contrast, beech AD, RD, and HT were all significantly reduced with increasing glyphosate rate. Post-release browsing by ungulates and a high residual overstory basal area resulted in reduced sugar maple HT. Our results indicated that glyphosate herbicide applied in stands that have been recently shelterwood seed cut can significantly increase the abundance of sugar maple regeneration. However, subsequent browsing damage combined with the negative influence of the residual overstory cover can limit the longer-term benefit of understory herbicide treatments. Subsequent removal of the overstory and browsing-control measures may be needed to promote sugar maple regeneration over beech in similar northern hardwood stands.     

Growth and photosynthetic responses of four Virginia Piedmont tree species to shade

To determine the effects of shade on biomass, carbon allocation patterns and photosynthetic response, seedlings of loblolly pine (Pinus taeda L.), white pine (Pinus strobus L.), red maple (Acer rubrum L.), and yellow-poplar (Liriodendron tulipifera L.) were grown without shade or in shade treatments providing a 79 or 89% reduction of full sunlight for two growing seasons. The shade treatments resulted in less total biomass for all species, with loblolly pine showing the greatest shade-induced growth reduction. Yellow-poplar was the only species to show increased stem height growth in the 89% shade treatment. The shade treatments increased specific leaf area of all species. Quantum efficiency, dark respiration and light compensation point were generally not affected by the shade treatments. Quantum efficiency, dark respiration, maximum photosynthesis and light compensation point did not change consistently between the first and second growing seasons. We conclude that differences in shade tolerance among these species are not the result of changes in the photosynthetic mechanism in response to shade.     

Seedling populations and regeneration of woody species in dry Afromontane forests of Ethiopia

The composition and density of seedlings, as well as height class distribution of woody species, were studied at two dry Afromontane forests in Ethiopia. A total of 40 species were recorded at Gara Ades and 41 at Menagesha forests. The density was 16 290 and 32 650 seedlings/ha at Gara Ades and Menagesha, respectively. A few species were not represented in the seedling bank, although mature trees were present. This may suggest that the species are under threat of local extinction or ‘that they recruit after major disturbances’. The trend in the height class distribution varied from species to species. However, about 50% of the species showed a negative exponential distribution at both sites, while there were indications of intermittent regeneration in the others. Seedling survival and growth of four species were also investigated, both under shade and in the open, at Gara Ades. The number of surviving seedlings declined progressively during the study period in all the species. Damage and uprooting by animals and people, drought, defoliation by insects and burial by siltation are among the apparent causes of seedling mortality. Mean annual height increment was generally low (< 15 cm) in the four species. Only one of the species showed more rapid height growth in gaps than under shade while the other three species had almost similar growth rates under shade and in gaps. The seedling growth analysis does not support the view that climax species quickly commence rapid height growth after canopy gap develops above them. The results show that in dry Afromontane forests, formation of seedling banks under the forest canopy is the major regeneration route of woody plants, especially climax species.     

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.     

Conspecific negative density dependence in a long-lived conifer,yew <Emphasis Type="Italic">Taxus baccata</Emphasis> L.

Conspecific negative density dependence in tree species can have major consequences for structuring of communities, yet in temperate forests this phenomenon remains largely unexplored. We investigated density-dependent recruitment in the long-lived conifer yew (Taxus baccata L.), for which low levels of regeneration have been linked with reduced light availability beneath conspecific canopies and speculative autotoxic effects. We combined in situ assessment of yew regeneration in a temperate forest in southwest Ireland with ex situ pot experiments to assess whether light and/or beneath conspecific canopy conditions inhibit the germination and early growth of yew. In field experiments, recruitment was related to distance from adult conspecifics, with higher levels of regeneration found in areas not beneath yew canopies. However, reduced light availability beneath the dense yew canopy was not responsible for this pattern, with abundant seedlings found in areas of equivalent or even lower light availability. Pot experiments showed that seedling survival was highest in deep shade treatments. Experiments also demonstrated that the addition of yew needles negatively affected seedling growth, but not seed germination. Together, our results suggest that under natural forest conditions, the absence of regeneration beneath mature yew canopies, often attributed to low light availability, may at least be partly related to substances in yew canopy or leaf leachates.     

Effect of gap size on mid-rotation stand structure and species composition in a naturally regenerated mixed broadleaf forest

This paper is an assessment of the effect of gap size on stand structure and species composition 48 years following treatment in a mixed broadleaf upland forest. Established in 1960, the study tests three circular openings, 15.2 m (0.02 ha), 45.7 m (0.16 ha), and 76.2 m (0.46 ha). Forty-eight years following treatment (2008) basal area, top height, and quadratic mean diameter were significantly lower in 15.2 m openings. Maple (Acer spp.) species had the highest mean importance value across treatments (0.40). Trends suggest that species composition of dominant and codominant trees among opening sizes may have been influenced by shade tolerance adaptations of the species groups present. Whereas 15.2 m openings were dominated by shade tolerant maple species, 45.7 and 76.2 m openings produced a mixture of commercial species including shade intolerant species such as yellow-poplar (Liriodendron tulipifera L.), trees of intermediate shade tolerance like oak (Quercus spp.), and shade tolerant maple. Data further suggest the density of overstory oak was highest in the intermediate opening size (45.7 m), while yellow-poplar increased in the larger opening sizes. Evaluation of species shifts between 1981 and 2008 showed that relative basal area of maple increased across all treatments. Relative basal areas from 45.7 to 76.2 m openings suggest declines in yellow-poplar and other non-commercial species were balanced by increases in oak and maple.     

Using competition and light estimates to predict diameter and height growth of naturally regenerated beech seedlings growing under changing canopy conditions

In a naturally regenerated beech (Fagus sylvatica L.) standin northeastern France, five regeneration patches were selectedwith different degrees of canopy closure; the percentage ofabove-canopy light (PACL) ranged between 5 and 45 per cent.The canopy was removed by a storm 2 years later. In order toanalyse the immediate and delayed effects of PACL on seedlingdiameter and height growth, the effects of the light conditionsprevailing before canopy opening on seedling growth measuredbefore and after canopy opening were tested. The effects oflocal density were examined using neighbourhood analysis. Fourdifferent competition indices (CIs) and neighbourhood radiiranging from 10 to 200 cm were tested. Models including PACLand CI accounted for between 56 and 64 per cent of the variationin individual seedling annual diameter or height growth. Localdensity had a strong negative influence on diameter growth anda much smaller influence on height growth. PACL was positivelycorrelated with diameter and height growth before canopy opening.Delayed effects of PACL on height growth were observed justafter canopy opening, but disappeared after 2 years. No delayedeffects of PACL on diameter growth after canopy opening wereobserved.     

Determining the shade tolerance of American chestnut using morphological and physiological leaf parameters

American chestnut (Castanea dentata (Marsh.) Borkh.) was once a principal component of the eastern deciduous forest until it became functionally extinct as a result of the invasive fungus Cryphonectria parasitica (Murr.) Barr. Restoration efforts are underway by means of a blight-resistant American-Chinese chestnut hybrid, and detailed silvicultural information, such as the shade tolerance of American chestnut and optimum site selection for restoration, is critical for planting success. In the present study, the physiological and morphological leaf characteristics of pure American chestnut seedlings, saplings, and mature trees were investigated in one of the few remaining stands of oak-chestnut vegetation (West Salem, WI) to determine shade tolerance. Seedlings, saplings, and mature trees had high maximum rates of photosynthesis, similar to shade intolerant species, and low light compensation points (LCPs), quantum efficiency, leaf mass per area (LMA), and percent nitrogen content, reminiscent of shade tolerant species. Dark respiration rates of seedlings and saplings were low, but increased in mature trees. LMA was found to significantly increase with height in the canopy, indicating a high level of light-induced plasticity. The results of this study suggest that American chestnut should be classified as intermediate in shade tolerance.     

Leaf weight per area and leaf size of 85 Estonian woody species in relation to shade tolerance and light availability

Leaf weight per area (LWA) and leaf size were examined in 85 species of woody plants representing 83% of Estonian native flora.

Average values of 68.6 g m⁻² for LWA and 6.62 cm² for leaf size were estimated for the medium light availability. Mean LWA values can be used to correct for the bias in estimating foliage functional properties in forest stand models and for converting leaf biomass into foliage area. LWA increased with relative light availability (1 minus stand canopy cover) and species light demand, indicating that LWA was higher at the same light availability for more intolerant species. Leaf size is a canopy parameter that should be considered when canopy radiative balance and distribution of solar radiation is of interest. Mean leaf size was not so closely related to light availability as LWA, decreasing with species light demand and increasing with total plant height. Variation in leaf size is discussed in terms of species shade tolerance and height growth.     

Development of Quercus robur advance regeneration following canopy reduction in an oak woodland

Observations to follow the development of 2-year-old oak seedlingstook place over a 6-year period at a fenced site within a maturestand of oak in southern England. Bramble and bracken were abundant,but the cover of other species was low. Although seedlings doubledin height, only 13 per cent survived and few were taller thaneither bramble or bracken. The initial number of seedlings wasinversely related to both canopy cover and distance to the nearestparent tree. The final number of seedlings could be predictedfrom the initial percentage covers of bramble and bracken, andeither the overstorey canopy cover or the initial number ofseedlings. The average annual rate of seedling mortality, whichwas 25–50 per cent, was significantly influenced by thevegetation present. Grass was positively related to survivalwhereas there was an inverse relationship with both brambleand bracken cover. Models estimated 80 per cent annual seedlingsurvival at 25 per cent bramble cover but only 50 per cent at60 per cent cover. Although grasses can be competitive species,results indicate that oak seedling establishment may be betterin areas with a ground flora comprising grass.