HETEROPHYLLY AND NEOFORMATION OF LEAVES IN SUGAR MAPLE (ACER SACCHARUM)

Variation in leaf form and timing of leaf initiation were investigated in vigorous leader shoots of open-grown saplings and larger forest trees of sugar maple (Acer saccharum Marsh.). Winter buds of leader shoots usually contained 6 or 8 leaf primordia and embryonic leaves, whereas 12 to 18 leaves typically expanded along the shoots each year. Preformed (early) leaves differ in form from neoformed (late) leaves. As in some other Acer species, the first-formed late leaves have large angles of secondary lobe divergence and deeply indented sinuses. This pattern of heterophylly contributes to the multilayered nature of open-grown saplings and leader shoots of forest trees of sugar maple.     

14C transfer between the spring ephemeral Erythronium americanum and sugar maple saplings via arbuscular mycorrhizal fungi in natural stands

We investigated in the field the carbon (C) transfer between sugar maple (Acer saccharum) saplings and the spring ephemeral Erythronium americanum via the mycelium of arbuscular mycorrhizal (AM) fungi. Sugar maple saplings and E. americanum plants were planted together in pots placed in the ground of a maple forest in 1999. Ectomycorrhizal yellow birches (Betula alleghaniensis) were added as control plants. In spring 2000, during leaf expansion of sugar maple saplings, the leaves of E. americanum were labelled with ¹⁴CO₂. Seven days after labelling, radioactivity was detected in leaves, stem and roots of sugar maples. Specific radioactivity in sugar maples was 13-fold higher than in yellow birches revealing the occurrence of a direct transfer of ¹⁴C between the AM plants. The quantity of ¹⁴C transferred to sugar maple saplings was negatively correlated with the percentage of ¹⁴C allocated to the storage organ of E. americanum. A second labelling was performed in autumn 2000 on sugar maple leaves during annual growth of E. americanum roots. Radioactivity was detected in 7 of 22 E. americanum root systems and absent in yellow birches. These results suggest that AM fungi connecting different understorey species can act as reciprocal C transfer bridges between plant species in relation with the phenology of the plants involved.     

Norway Maple (Acer platanoides) Invasion of a Natural Forest Stand: Understory Consequence and Regeneration Pattern

Norway maple (Acer platanoidesis) is invasive in a natural stand in suburban Ithaca, NY. To determine the understory pattern and consequences of a Norway maple invasion, I compared density and species richness under Norway maples and sugar maples (Acer saccharum). Mean sapling density was significantly lower (P<0.0027) under Norway maples (3.64/100 m²±1.6 SE) than under sugar maples (19.4/100 m²±4.4 SE). Mean sapling species richness was significantly lower (P<0.0018) under Norway maples (0.7/32 m²±0.18 SE) than under sugar maples (2.6/32 m²±0.48 SE). Likewise, Norway maple regeneration is more frequent under sugar maples than sugar maple regeneration: 57% of sugar maple plots had Norway maple saplings while 0% of Norway maple plots had sugar maple saplings. Two significant plot effects were found for presence–absence: Norway maple saplings grow under Norway maples with a significantly lower frequency (P<0.03) than under sugar maples; sugar maple saplings grow under Norway maples with a significantly lower frequency (P<0.000) than under sugar maples. Across the site, Norway maple saplings were the most abundant (29 saplings for 480 m²). The success of Norway maple regeneration and the reductions in total stem density beneath Norway maples is most likely the result of its strong competitive abilities, notably its high shade tolerance and abundant seed crops.     

Diversity of canopy and understorey spiders in north-temperate hardwood forests

1 We characterized and compared diversity patterns of canopy and understorey spiders (Arachnida: Araneae) on sugar maple ( Acer saccharum Marsh.) and American beech ( Fagus grandifolia Ehrh.) in hardwood forests of southern Québec, Canada.
2 We sampled canopies of 45 sugar maple and 45 American beech trees and associated understorey saplings in mature protected forests near Montréal. Samples were obtained by beating the crown foliage at various heights and by beating saplings around each tree.
3 Eighty-two species were identified from 13 669 individuals. Forty-eight species and 3860 individuals and 72 species and 9809 individuals were collected from the canopy and the understorey, respectively.
4 Multivariate analyses (NMDS ordination and NPMANOVA) showed the composition of canopy and understorey assemblages differed significantly, and canopy assemblages differed between tree species. Rank-abundance distribution models fitted to the canopy and understorey data indicated that different mechanisms structure the assemblages in both habitats. Three abundant spider species were significantly more common in the canopy; ten species were collected significantly more often in the understorey.
5 The forest canopy was shown to be an important reservoir for spider diversity in north-temperate forests.     

Differential drought responses between saplings and adult trees in four co-occurring species of New England

Tree-ring characteristics in four species were examined to address whether co-occurring mature trees of different successional status respond differently to drought, and whether saplings of these species have a greater response to drought than mature trees. We examined saplings and mature trees of paper birch, yellow birch, red maple and sugar maple, which varied in successional status (shade-tolerance) and co-occurred at Harvard Forest, Petersham, Mass., USA. Three drought events in 1964–1966, 1981 and 1995 were identified using climate data. For mature trees, there was no significant interspecific difference in relative changes in ring-width index (RWI) during the 1964–1966 and 1995 drought events. However, the interspecific difference was significant in the 1981 drought event. Response function analysis for mature trees showed that the radial growth of sugar maple was mainly controlled by spring and summer precipitation, red maple by spring and summer precipitation and temperature, yellow birch by winter and summer precipitation, and spring and summer temperature, and paper birch by spring and summer precipitation and spring temperature. Saplings of sugar maple and yellow birch, but not red maple and paper birch, showed significant positive correlations between RWI and annual total precipitation. In the 1995 drought event, saplings and mature trees of red maple and paper birch differed significantly in drought responses, but this was not true in sugar maple and yellow birch. Our results do not support a generally greater response in saplings than in mature trees, nor an early- versus late successional difference in drought responses.     

Application of the Functional-Structural Tree Model LIGNUM to Sugar Maple Saplings (Acer saccharum Marsh) Growing in Forest Gaps

LIGNUM is a functional-structural model that represents a treeusing four modelling units which closely resemble the real structureof trees: tree segments, tree axes, branching points and buds.Metabolic processes are explicitly related to the structuralunits in which they take place. Here we adapt earlier versionsof LIGNUM designed to model growth of conifers for use withbroad-leaved trees. Two primary changes are involved. First,the tree segment for broad-leaved trees consists of enclosedcylinders of heartwood, sapwood and bark. Leaves consistingof petioles and blades are attached to the segments. Secondly,axillary buds and rules governing their dormancy are includedin the model. This modified version of LIGNUM is used to simulatethe growth and form of sugar maple saplings in forest gaps.The annual growth of the model tree is driven by net productionafter respiration losses are taken into account. The productionrate of each leaf depends on the amount of photosyntheticallyactive radiation it receives. The radiation regime is trackedexplicitly in different parts of the tree crown using a modelof mutual shading of the leaves. Forest gaps are representedby changing the radiation intensity in different parts of themodel sky. This version of LIGNUM modified for use with broad-leaf,deciduous trees and parameterized for sugar maple, yields goodsimulations of growth and form in saplings from different forestgap environments. Copyright 2001 Annals of Botany Company LIGNUM, functional-structural tree model, tree architecture, sugar maple, modelling growth and form     

Patterns of tree replacement: canopy effects on understory pattern in hemlock-northern hardwood forests

The effect of canopy trees on understory seedling and sapling distribution is examined in near-climax hemlock-northern hardwood forests in order to predict tree replacement patterns and assess compositional stability. Canopy trees and saplings were mapped in 65 0.1-ha plots in 16 tracts of old-growth forests dominated by Tsuga canadensis, Acer saccharum, Fagus grandifolia, Tilia americana, and Betula lutea in the northeastern United States. Seedlings were tallied in sub-plots. Canopy influence on individual saplings and sub-plots was calculated, using several indices for canopy species individually and in total. For each species sapling and seedling distributions were compared to those distributions expected if saplings were located independently of canopy influence. Non-random distributions indicated that sapling and seedling establishment or mortality were related to the species of nearby canopy trees. Hemlock canopy trees discriminate against beech and maple saplings while sugar maple canopy favors beech saplings relative to other species. Basswood canopy discourages growth of saplings of other species, but produces basal sprouts. Yellow birch saplings were rarely seen beneath intact canopy. Since trees in these forests are usually replaced by suppressed seedlings or saplings, canopy-understory interactions should influence replacement probabilities and, ultimately, stand composition. I suggest that hemlock and basswood tend to be self-replacing, maple and beech tend to replace each other, and birch survives as a fugitive by occupying occasional suitable gaps. This suggests that these species may co-exist within stands for long periods with little likelihood of successional elimination of any species. There is some suggestion of geographical variation in these patterns.     

Tree demography suggests multiple directions and drivers for species range shifts in mountains of Northeastern United States

Climate change is expected to lead to upslope shifts in tree species distributions, but the evidence is mixed partly due to land‐use effects and individualistic species responses to climate. We examined how individual tree species demography varies along elevational climatic gradients across four states in the northeastern United States to determine whether species elevational distributions and their potential upslope (or downslope) shifts were controlled by climate, land‐use legacies (past logging), or soils. We characterized tree demography, microclimate, land‐use legacies, and soils at 83 sites stratified by elevation (~500 to ~1200 m above sea level) across 12 mountains containing the transition from northern hardwood to spruce‐fir forests. We modeled elevational distributions of tree species saplings and adults using logistic regression to test whether sapling distributions suggest ongoing species range expansion upslope (or contraction downslope) relative to adults, and we used linear mixed models to determine the extent to which climate, land use, and soil variables explain these distributions. Tree demography varied with elevation by species, suggesting a potential upslope shift only for American beech, downslope shifts for red spruce (more so in cool regions) and sugar maple, and no change with elevation for balsam fir. While soils had relatively minor effects, climate was the dominant predictor for most species and more so for saplings than adults of red spruce, sugar maple, yellow birch, cordate birch, and striped maple. On the other hand, logging legacies were positively associated with American beech, sugar maple, and yellow birch, and negatively with red spruce and balsam fir – generally more so for adults than saplings. All species exhibited individualistic rather than synchronous demographic responses to climate and land use, and the return of red spruce to lower elevations where past logging originally benefited northern hardwood species indicates that land use may mask species range shifts caused by changing climate.     

Dendroclimatology of sugar maple (Acer saccharum): Climate-growth response in a late-successional species

Sugar maple (Acer saccharum) is a shade-tolerant, late successional dominant species in the North American eastern deciduous forest. The objective of this study was to quantify the relationship between climate and radial growth in sugar maple and to identify spatial and temporal patterns in dendroclimatic response. We used a combination of archived sugar maple tree-ring chronologies and newly sampled sites to calculate dendroclimatic response of sugar maple at 13 sites in the United States and Canada. At all sites, sugar maple growth was significantly correlated to monthly temperature, precipitation, or Palmer Drought Severity Index. However, there were no generalizable patterns in sugar maple’s growth response to climate. Individual sites had unique dendroclimatic responses with respect to: a) which climatic variables were correlated to radial growth; b) what months had significant correlations between climate and radial growth; and c) what years had significant correlations between climate and radial growth. The individualistic dendroclimatic response of sugar maple appears to reflect a plastic response of the species to changes in climate perhaps related to its status as a strong competitor in late-successional forests. This ability to survive a wide range of environmental conditions may bode well for the species persistence under variable future climatic conditions. It also points to the need for more research on late-successional species in examining forest response to potential climate change scenarios because these species may be more resilient than early-successional species.     

Patterns of rhizosphere carbon flux in sugar maple (Acer saccharum) and yellow birch (Betula allegheniensis) saplings

Despite its importance in the terrestrial C cycle rhizosphere carbon flux (RCF) has rarely been measured for intact root–soil systems. We measured RCF for 8‐year‐old saplings of sugar maple (Acer saccharum) and yellow birch (Betula allegheniensis) collected from the Hubbard Brook Experimental Forest (HBEF), NH and transplanted into pots with native soil horizons intact. Five saplings of each species were pulse labeled with ¹³CO₂ at ambient CO₂ concentrations for 4–6 h, and the ¹³C label was chased through rhizosphere and bulk soil pools in organic and mineral horizons for 7 days. We hypothesized yellow birch roots would supply more labile C to the rhizosphere than sugar maple roots based on the presumed greater C requirements of ectomycorrhizal roots. We observed appearance of the label in rhizosphere soil of both species within the first 24 h, and a striking difference between species in the timing of ¹³C release to soil. In sugar maple, peak concentration of the label appeared 1 day after labeling and declined over time whereas in birch the label increased in concentration over the 7‐day chase period. The sum of root and rhizomicrobial respiration in the pots was 19% and 26% of total soil respiration in sugar maple and yellow birch, respectively. Our estimate of the total amount of RCF released by roots was 6.9–7.1% of assimilated C in sugar maple and 11.2–13.0% of assimilated C in yellow birch. These fluxes extrapolate to 55–57 and 90–104 g C m^?2 yr^?1 from sugar maple and yellow birch roots, respectively. These results suggest RCF from both arbuscular mycorrhizal and ectomycorrhizal roots represents a substantial flux of C to soil in northern hardwood forests with important implications for soil microbial activity, nutrient availability and C storage.     

Sapling dynamics in a southeastern Texas floodplain forest

Abstract. How much temporal variation in recruitment, mortality and change in size class occurs in the sapling layer of mature temperate forests in the absence of large‐scale exogenous disturbance? Using 15 years of data from a flood‐plain forest in Big Thicket National Preserve, we found that year‐to‐year variation in demographic parameters was greater than we originally expected. Death rates were generally more variable than recruitment rates, and were much more variable for large saplings than for small ones. Small saplings of the 10 most common species had at least one year when they experienced two to eight times their long‐term mean recruitment and death rates. Large saplings had at least one year when they experienced three to 10 times their long‐term mean death rates and at least one year with two to seven times the long‐term mean recruitment rate. Temporal patterns in sapling death rates were related to flooding patterns, while temporal patterns in recruitment were related to the Palmer Drought Severity Index, an indicator of drought severity and soil moisture availability. We also identified apparently synchronous patterns of demographic response among less flood‐tolerant species which differed from the responses of more flood‐tolerant species. We demonstrated the effects of both climatic variation and light variation in affecting stand‐wide sapling demographics in a forest where canopy gaps are important for regeneration, and where chronic understorey disturbance favours growth over survivorship as a sapling strategy.     

Japanese maple (Acer palmatum var. Matsumurae, Aceraceae) recruitment patterns: seeds, seedlings, and saplings in relation to conspecific adult neighbors

We analyzed the spatial patterns among seeds, seedlings, saplings, and conspecific adult trees of the cool-temperate tree species Acer palmatum var. Matsumurae in a conifer-hardwood mixed forest in northern Japan, using two models that consider the influence of each adult within the neighborhood of the offspring. The results showed that recruitment patterns of each stage could be characterized and that significant shifts occur between successive stages. Sound seeds were more widely dispersed than unsound seeds; the mean dispersal distance (MDD) was 41.5 m for sound seeds, but only 12.6 m for unsound seeds. Most seedlings were located near conspecific adult trees, with a MDD of 14.3 m. Saplings, however, were more dispersed away from conspecific adult trees, with an MDD of more than 35 m. Light and gap distributions did not strongly affect the spatial distribution of the offspring; most saplings were located under nonconspecific canopies. These results suggest that the recruitment pattern of Japanese maple offspring is strongly affected by conspecific adult neighbors, rather than by light and gap distributions, with close proximity to conspecific adult trees reducing the growth and survival of seedlings during the transition to saplings.     

A gall-inducing arthropod drives declines in canopy tree photosynthesis

Mature forest canopies sustain an enormous diversity of herbivorous arthropods; however, with the exception of species that exhibit large-scale outbreaks, canopy arthropods are thought to have relatively little influence on overall forest productivity. Diminutive gall-inducing mites (Acari; Eriophyoidae) are ubiquitous in forest canopies and are almost always highly host specific, but despite their pervasive occurrence, the impacts of these obligate parasites on canopy physiology have not been examined. We have documented large declines in photosynthetic capacity (approx. 60%) and stomatal conductance (approx. 50%) in canopy leaves of mature sugar maple (Acer saccharum) trees frequently galled by the maple spindle gall mite Vasates aceriscrumena. Remarkably, such large impacts occurred at very low levels of galling, with the presence of only a few galls (occupying approx. 1% of leaf area) compromising gas-exchange across the entire leaf. In contrast to these extreme impacts on the leaves of adult trees, galls had no detectible effect on the gas-exchange of maple saplings, implying large ontogenetic differences in host tolerance to mite galling. We also found a significant negative correlation between canopy tree radial increment growth and levels of mite galling. Increased galling levels and higher physiological susceptibility in older canopy trees thus suggest that gall-inducing mites may be major drivers of “age-dependent” reductions in the physiological performance and growth of older trees.     

Inadequate Cold Tolerance as a Possible Limitation to American Chestnut Restoration in the Northeastern United States

The American chestnut (Castanea dentata (Marshall) Borkh.), once a major component of eastern forests from Maine to Georgia, was functionally removed from the forest ecosystem by chestnut blight (an exotic fungal disease caused by Cryphonectria parasitica (Murr.) Barr), first identified at the beginning of the twentieth century. Hybrid‐backcross breeding programs that incorporate the blight resistance of Chinese chestnut (Castenea mollissima Blume) and Japanese chestnut (Castenea crenata Sieb. & Zuc.) into American chestnut stock show promise for achieving the blight resistance needed for species restoration. However, it is uncertain if limitations in tissue cold tolerance within current breeding programs might restrict the restoration of the species at the northern limits of American chestnut's historic range. Shoots of American chestnut and hybrid‐backcross chestnut (i.e., backcross chestnut) saplings growing in two plantings in Vermont were tested during November 2006, February 2007, and April 2007 to assess their cold tolerance relative to ambient low temperatures. Shoots of two potential native competitors, northern red oak (Quercus rubra L.) and sugar maple (Acer saccharum L.), were also sampled for comparison. During the winter, American and backcross chestnuts were approximately 5°C less cold tolerant than red oak and sugar maple, with a tendency for American chestnut to be more cold tolerant than the backcross chestnut. Terminal shoots of American and backcross chestnut also showed significantly more freezing damage in the field than nearby red oak and sugar maple shoots, which showed no visible injury.     

The effect of deforestation on the genetic diversity and structure in <Emphasis Type="Italic">Acer saccharum</Emphasis> (Marsh): Evidence for the loss and restructuring of genetic variation in a natural system

The level and distribution of genetic diversity can be influenced by species life history traits and demographic factors, including perturbations that might produce population bottlenecks. Deforestation and forest fragmentation are common sources of population disturbance in contemporary populations of forest ecosystems. Although the genetic effects of forest fragmentation and deforestation have been examined by assessing levels of genetic variation in forest fragments that remain after logging, few considerations have been made of the populations that re-colonize once-cleared areas. Here we examine the effects of human-mediated population bottlenecks on the level and distribution of genetic diversity in natural populations of the long-lived forest tree species, Acer saccharum (sugar maple). We compared genetic variation and structure for populations of sugar maple found within old-growth forested area and in area that has re-colonized since logging. In this study the percent polymorphic loci and allelic richness estimates were reduced in the logged populations compared to old-growth populations. Jackknifed estimates of population genetic differentiation showed significantly higher differentiation among logged populations, with this result being consistently seen when individuals within populations were grouped according to diameter at breast height. The result of decreased genetic variation and higher levels of genetic structure among logged populations suggests that even one extensive bout of logging can alter the level and distribution of genetic variation in this forest tree species.     

Light and tree size influence belowground development in yellow birch and sugar maple

The effects of light and tree size on the root architecture and mycorrhiza of yellow birch (Betula alleghaniensis Britton) and sugar maple (Acer saccharum Marsh) growing in the understory of deciduous forests in southern Québec, Canada were studied. At the study site, small (<50 m²), medium (101–200 m²) and large (201–500 m²) canopy gaps were investigated. From within these gaps, 17 yellow birch and 23 sugar maple saplings from 40 to 600 cm in height were sampled. In both species, root biomass and morphological traits were strongly correlated with tree size, but only weakly with light availability. Increased root biomass was primarily allocated to coarse roots and secondarily to fine roots. Yellow birch roots were longer, had a larger area, more endings and branches and grew more rapidly than sugar maple roots. Mycorrhizal colonization increased with available light and declined with tree age in sugar maple and was positively associated with tree size in yellow birch. The study demonstrates that tree size is a very important determinant of how belowground systems acclimate to understory conditions.     

Conifer Presence May Negatively Affect Sugar Maple’s Ability to Migrate into the Boreal Forest Through Reduced Foliar Nutritional Status

The discipline of ecology suffers from a lack of knowledge of non-climatic factors (for example, plant–soil, plant–plant and plant–insect interactions) to predict tree species range shifts under climate change. The next generation of simulation models of forest response to climate change must build upon local observations of species interactions and growth along climatic gradients. We examined whether sugar maple (Acer saccharum) seedlings were disadvantaged with respect to soil nutrient uptake under coniferous canopies, as this species would need to migrate northward into conifer-dominated forests in response to climate change. An experimental design was applied to 3 sites, forming the largest possible latitudinal/climatic gradient for sugar maple in Quebec (Canada) and isolating the effect of conifer presence on its seedling’s nutritional status. We tested whether: (1) both soil and climate and (2) presence of conifers affected foliar nutrient levels of sugar maple seedlings. Climate and soil (through pH) strongly affected nutrient availability for sugar maple seedlings and predicted 63.7% of their foliar nutrient variability. When controlling for site effects, we found a significant negative effect of conifers on foliar Ca and Mg levels of maple seedlings, which can adversely affect their overall health and vigour. When considering projected modifications of the forest environment due to climate change, we suggest that northward migration of sugar maple will be negatively affected by the presence of conifers through reduced foliar nutrition.     

Testing the enemy release hypothesis: a comparison of foliar insect herbivory of the exotic Norway maple (Acer platanoides L.) and the native sugar maple (A. saccharum L.)

Norway maple (Acer platanoides) is a Eurasian introduced tree species which has invaded the North American range of its native congener, sugar maple (A. saccharum). One hypothesis used to explain the success of an invasive species is the enemy release hypothesis (ERH), which states that invasive species are often particularly successful in their new range because they lack the enemies of their native range. In this study, we hypothesized that Norway maple would have less insect damage than sugar maple due to such enemy release. Autumn 2005 and summer 2006 leaves of Norway and sugar maple were collected from six sites in New Jersey and Pennsylvania to compare percent leaf area loss, gall damage, fungal damage, and specific leaf area (cm²/g). Although both species had low overall mean levels of leaf damage (0.4–2.5%), in both years/seasons Norway maple had significantly less leaf damage than sugar maple. Insects were also collected to compare insect assemblies present on each tree species. The numbers of insect taxa and individuals found on each species were nearly equivalent. Overall, the results of this study are consistent with the enemy release hypothesis for Norway maple. In addition, sugar maples when surrounded by Norway maples tended to show reduced herbivory. This suggests that the spread of Norway maple in North America, by reducing amounts of insect herbivory, may have further ecosystem-wide impacts.     

Changes in stand composition and structure between 1981 and 1996 in four Front Range plant communities in Colorado

We measured leaf distribution, crown shape, and biomass allocation patterns in open-grown saplings of three tree species (Cedrela odorata, Cordia alliodora, and Hyeronima alchorneoides) that differed in above-ground architecture and successional status. The objectives were to: (1) analyze the relationship between crown structure and biomass allocation in free-growing juveniles and (2) assess whether predictions that relate crown structure and biomass distribution of adult individuals can be applied to noncompeting saplings. Significant differences in crown structure and biomass allocation were found among the three species. Mean dry biomass was 990 g in Cedrela, 665 g, in Cordia, and 1281 g in Hyeronima. Cedrela allocated a greater proportion of biomass into roots than the other two species. We observed no distinct pattern of foliage angle, and no evidence of decrease in crown construction efficiency with tree size. Predictions stated by other authors for mature individuals partially explained the relationships between biomass allocation and morphological variables of non-competing saplings. We proposed two alternative biomass allocation patterns: (1) in open conditions saplings of early successional species exhibit structural characteristics more adapted for high light levels than gap-dependent species, and (2) early successional species have lower cost-benefit ratio for biomass construction and higher foliar efficiency than late successional ones. In general, crown morphometric characteristics and biomass distribution patterns of the three species studied corresponded with the two proposed patterns.     

Inter- and intraspecific relationships between shade tolerance and shade avoidance in temperate trees

Patterns of physiological and architectural adaptation and acclimation to decreasing light availability were investigated along a light gradient for saplings of 12 common species of temperate deciduous trees in southeastern Ontario, Canada. Physiological adaptation and acclimation (shade tolerance physiology) were quantified at the leaf level by measuring leaf mass per unit area (LMA), dark respiration per unit leaf nitrogen, chlorophyll per unit leaf nitrogen and the chlorophyll a:b ratio for the newest fully expanded leaf on the leader. Architectural adaptation and acclimation (shade avoidance) were quantified by measuring branching intensity and side shoot:main shoot length ratios for the most recent three years of growth on the leader and selected side branches. Within species, increases in LMA, chlorophyll a:b ratio and respiration per unit nitrogen and decreases in chlorophyll per unit nitrogen indicated that shade tolerance physiology generally increased with decreasing canopy openness. Increases in the branching intensity and side shoot:main shoot ratios of the leader and side branches indicated that shade avoidance also increased with decreasing canopy openness for the majority of species; however, in some species, stem bending under deep shade resulted in lateral growth. Interspecific variation in shade tolerance physiology was minimal when species were compared under equal amounts of canopy openness. In contrast, interspecific variation in shade avoidance variables was relatively high under equal canopy openness, with saplings of shade tolerant canopy species exhibiting higher shade avoidance than saplings of shade tolerant understory species.