Photosynthetic responses of Miconia species to canopy openings in a lowland tropical rainforest

We examined the photosynthetic acclimation of three tropical species of Miconia to canopy openings in a Costa Rican rainforest. The response of photosynthesis to canopy opening was very similar in Miconia affinis, M. gracilis, and M. nervosa, despite differences in growth form (trees and shrubs) and local distributions of plants (understory and gap). Four months after the canopy was opened by a treefall, photosynthetic capacity in all three species had approximately doubled from closed canopy levels. There were no obvious signs of high light damage after treefall but acclimation to the gap environment was not immediate. Two weeks after treefall, A_max, stomatal conductance, apprarent quantum efficiency, and dark respiration rates had not changed significantly from understory values. The production of new leaves appears to be an important component of light acclimation in these species. The only variables to differ significantly among species were stomatal conductance at A_max and the light level at which assimilation was saturated. M. affinis had a higher stomatal conductance which may reduce its water use efficiency in gap environments. Photosynthesis in the more shade-tolerant M. gracilis saturated at lower light levels than in the other two species. Individual plant light environments were assessed after treefall with canopy photography but they explained only a small fraction of plant variation in most measures of photosynthesis and growth. In conclusion, we speculate that species differences in local distribution and in light requirements for reproduction may be more strongly related to species differences in carbon allocation than in carbon assimilation.     

香果树实生苗的光合特性及其与环境因子的关系

为确定香果树实生苗的适生环境并为其自然更新提出有针对性策略,研究了不同生境(冠下、冠缘、林窗和林缘)中2年生香果树实生苗的净光合速率、水分利用效率、叶绿素含量、苗高、基径、生物量等的变化及其与生态因子之间的关系.结果表明: 4种生境中的光合有效辐射最大值为50～1380 μmol·m^-2·s^-1,冠下和冠缘中香果树实生苗的净光合速率日变化呈单峰型,而林窗和林缘实生苗的净光合速率日变化呈双峰型;香果树实生苗为耐阴植物,但耐阴能力较弱,其功能叶的光饱和点、补偿点和暗呼吸在4种生境中大小顺序为: 林缘>林窗>冠缘>冠下,表观量子效率的变化规律与之相反;林窗和冠缘2种生境中香果树实生苗的适应能力较强,叶片的蒸腾速率、气孔导度、水分利用效率和净光合速率较高;林窗中香果树实生苗叶片的叶绿素含量较低,但实生苗的生长速度最快,生物量最大;香果树实生苗的净光合速率与光合有效辐射和气孔导度呈显著正相关.对于冠下生境,需降低林冠层密度,增加透光率,以利于香果树实生苗的光合作用;对于林缘生境,则需要增加植被盖度,降低光照强度,以利于其快速生长.     

Arbuscular mycorrhizal colonization of giant sequoia (Sequoiadendron giganteum) in response to restoration practices

Interactions with soil microbiota determine the success of restoring plants to their native habitats. The goal of our study was to understand the effects of restoration practices on interactions of giant sequoia Sequoiadendron giganteum with arbuscular mycorrhizal (AM) fungi (Glomeromycota). Natural regeneration of Sequoiadendron is threatened by the absence of severe fires that create forest canopy gaps. Generating artificial canopy gaps offers an alternative tool for giant sequoia restoration. We investigated the effect of regeneration practices, including (i) sapling location within gaps, (ii) gap size and (iii) soil substrate, on AM fungal colonization of giant sequoia sapling roots in a native giant sequoia grove of the Sierra Nevada, California. We found that the extent of AM fungal root colonization was positively correlated with sapling height and light availability, which were related to the location of the sapling within the gap and the gap size. While colonization frequency by arbuscules in saplings on ash substrate was higher relative to saplings in mineral soil, the total AM fungal root colonization was similar between the substrates. A negative correlation between root colonization by Glomeromycota and non-AM fungal species indicated antagonistic interactions between different classes of root-associated fungi. Using DNA genotyping, we identified six AM fungal taxa representing genera Glomus and Ambispora present in Sequoiadendron roots. Overall, we found that AM fungal colonization of giant sequoia roots was associated with availability of plant-assimilated carbon to the fungus rather than with the AM fungal supply of mineral nutrients to the roots. We conclude that restoration practices affecting light availability and carbon assimilation alter feedbacks between sapling growth and activity of AM fungi in the roots.     

Leaf Gas Exchange in Canopy Species of a Venezuelan Cloud Forest

Tropical cloud forests are considered humid ecosystems with frequent cloud cover down to the ground surface. However, seasonal variation in precipitation may induce short-term water stress. For canopy leaves, this water stress may also be a consequence of large atmospheric vapor pressure deficits. The objective of this work was to study five canopy cloud forest species to determine if there are restrictions to leaf gas exchange as a consequence of seasonality in precipitation and to daily water deficit due to air evaporative demand mainly during maximum incoming radiation hours. Seasonal daily courses of microclimatic variables (air temperature, relative humidity, photosynthetic photon flux density) and plant responses (leaf water potential, stomatal conductance, CO₂ assimilation rates, leaf nitrogen concentration) were measured at 2400 m asl in Monterrey, an intermontane valley of the Venezuelan Andes. A gradient in terms of responses to water stress conditions was observed between the species, with Clusia multiflora (a 46% reduction in stomatal conductance between seasons) as the most affected and Miconia resimoides (increased stomatal conductance) responding more favorably to slight water stress conditions. If we consider the limitations of water stress and/or light conditions on CO₂ assimilation we may arrange the species into those in which water stress conditions have a greater impact on leaf carbon gain, those where light conditions are determinant and one in which both water stress and light conditions may affect leaf carbon assimilation.     

Composition and Dynamics of Functional Groups of Trees During Tropical Forest Succession in Northeastern Costa Rica

We compared the functional type composition of trees ≥10 cm dbh in eight secondary forest monitoring plots with logged and unlogged mature forest plots in lowland wet forests of Northeastern Costa Rica. Five plant functional types were delimited based on diameter growth rates and canopy height of 293 tree species. Mature forests had significantly higher relative abundance of understory trees and slow-growing canopy/emergent trees, but lower relative abundance of fast-growing canopy/emergent trees than secondary forests. Fast-growing subcanopy and canopy trees reached peak densities early in succession. Density of fast-growing canopy/emergent trees increased during the first 20 yr of succession, whereas basal area continued to increase beyond 40 yr. We also assigned canopy tree species to one of three colonization groups, based on the presence of seedlings, saplings, and trees in four secondary forest plots. Among 93 species evaluated, 68 percent were classified as regenerating pioneers (both trees and regeneration present), whereas only 6 percent were classified as nonregenerating pioneers (trees only) and 26 percent as forest colonizers (regeneration only). Slow-growing trees composed 72 percent of the seedling and sapling regeneration for forest colonizers, whereas fast-growing trees composed 63 percent of the seedlings and saplings of regenerating pioneers. Tree stature and growth rates capture much of the functional variation that appears to drive successional dynamics. Results further suggest strong linkages between functional types defined based on adult height and growth rates of large trees and abundance of seedling and sapling regeneration during secondary succession.
Abstract in Spanish is available at http://www.blackwell-synergy.com/loi/btp     

柞蚕林冠层结构,林内光照分布与叶生物量研究

柞蚕林是以养蚕为目的的萌生栎林,林冠经剪伐而形成不同的冠型结构。本文对3种冠型的叶面积系数,冠层内叶量分布及消光系数等冠层结构特点进行了测定。结果表明,不同冠形的冠层结构特点明显影响林内光照环境及叶生物量。现有的3种冠型中,“阶梯”型结构有较高的叶面积系数和较合理的叶量分布,因而具有较高的叶生物量。“中干”型叶生物量与无干型虽无明显差异,但“中干”型消光系数较低,林内空间较大,林内光照环境比“无干”型结构优越。     

Diversity of woody species in forest,treefall gaps,and edge in Kibale National Park,Uganda

If specialization influences species presence, then high tropical tree and shrub diversity should correspond with high environmental heterogeneity. Such heterogeneity may be found among different successional communities (i.e., canopy types). We explore species associations in three forest-dominated canopy types, forest, gap, and edge, in Kibale National Park, Uganda and determine environmental, soil and light, differences among canopy types. To determine the strength of differences among forested canopy types, they are also compared to grasslands. Tree and shrub density and species richness using rarefaction analysis were determined based on data from 24 small plots (5 × 5 m) in all four canopy types and 16 large plots (10 × 50 m) in forest and grassland canopy types. Environmental variables were determined along 10 (20 m) transects in the four canopy types. Using analysis of variance and principal components analysis, we demonstrate that forest and gap environments had similar soils, but forest had lower light levels than gap. We also found that grassland and edge were more similar to one another than to forest and gap, but differed in a number of important biotic and abiotic factors controlling soil water availability (e.g., edge had higher root length density of small roots < 2 mm diameter in the top 20 cm than grassland). Using principal components analysis to assess similarities in community composition, we demonstrate that gap and forest had indistinguishable communities and that edge was similar to but distinct from both communities. Complete species turnover only occurred between grassland and the three forested canopy types. Even though overall community composition was similar in the three forested canopy types, in analyses of individual species using randomization tests, many common species were most frequently found in only one canopy type; these patterns held across size classes. These results suggest that despite differences among environments, community composition was similar among forested canopy types, which are likely intergrading into one another. Interestingly, individual species are more frequently found in a single canopy type, indicating species specialization.     

Drought-induced shifts in the floristic and functional composition of tropical forests in Ghana

The future of tropical forests under global environmental change is uncertain, with biodiversity and carbon stocks at risk if precipitation regimes alter. Here, we assess changes in plant functional composition and biomass in 19 plots from a variety of forest types during two decades of long-term drought in Ghana. We find a consistent increase in dry forest, deciduous, canopy species with intermediate light demand and a concomitant decrease in wet forest, evergreen, sub-canopy and shade-tolerant species. These changes in composition are accompanied by an increase in above-ground biomass. Our results indicate that by altering composition in favour of drought-tolerant species, the biomass stocks of these forests may be more resilient to longer term drought than short-term studies of severe individual droughts suggest.     

Gap disturbances in northern old‐growth forests of British Columbia,Canada

Abstract. We characterized the abundance, size and spatial patterning of canopy gaps, as well as gap‐forming processes and light availability in boreal, sub‐boreal, northern temperate and subalpine old‐growth forests of northwestern British Columbia. The proportion of area in canopy gaps ranged from 32% in northern temperate forests to 73% in subalpine forests. Evenly distributed developmental gaps were dominant but permanent openings created by edaphic components and by shrub communities were also common, particularly in subboreal forests. Abundant gaps, large gap sizes, high numbers of gap makers per gap and frequent gap expansion events suggest that gaps have long tenure in these forests. Snapped stems and standing dead mortality were the most common modes of mortality in all forest types resulting in little forest floor disturbance, creating few germination sites for seedling establishment. We found high mean light levels (16–27% full sun) and little difference between non‐gap and gap light environments. Our results suggest that gap dynamics in these forests differ fundamentally from those in temperate and tropical forest ecosystems.     

亚高山暗针叶林不同林冠环境下华西箭竹的克隆可塑性

以亚高山暗针叶林3种林冠环境中以及暗针叶林林缘的华西箭竹（Fargesia nitida）为对象,对其无性系数量特征、无性系根茎特征、分株生物量以及分株形态特征进行了对比研究。结果表明：（1）林冠环境的差异导致了不同种群的基株密度和每基株分株数的显著差异,但林冠环境差异不影响分株密度。林冠郁闭度愈大,每基株分株数愈少,分株分布愈均匀。（2）不同林冠环境间。分株生物量、分株构件生物量和分株构件的生物量分配百分率均有显著差异。开敞的林冠环境有利于华西箭竹的生长和生物量积累。（3）随着林冠郁闭度的增加,华西箭竹通过增大分枝角度、叶生物量分配百分率、比叶面积和叶面积率以提高光能利用效率,有效适应弱光环境。（4）隔离者长度、隔离者直径和分枝强度在林缘和林窗环境中要显著大于林内环境;同级隔离者分枝角度随林冠郁闭度的增加而最大,其值在林下显著大于林窗和林缘,而异级隔离者分枝角度的变化则正好相反。研究表明,华西箭竹种群在不同的林冠环境中发生了明显的可塑性变化,这些可塑性变化是种群对林冠郁闭度差异的适应性反应的结果,有利于增强种群对环境中有限光资源的利用。     

Phenotypic plasticity of lianas in response to altered light environment

The growth, morphology and biomass allocation of 11 liana species (six light-demanding and five shade-tolerant) were investigated by growing plants in three contrasting light environments (i.e., field, forest edge and forest interior). Our objectives were to determine: (1) changes in plant traits at the species level; and (2) differences in light-demanding and shade-tolerant species in response to altered light environment. We found that all seedlings of liana species increased in total biomass, total leaf area, relative growth rate (RGR), net assimilation rate (NAR), height, basal diameter, root length, leaf number, root mass/total plant mass (RMR) and root-to-shoot dry biomass (R/S ratio), and decreased in leaf area ratio (LAR), specific leaf area (SLA), leaf size, stem mass-to-total plant mass ratio (SMR) and leaf mass-to-total plant mass ratio (LMR) with increasing light availability. Under the three light environments, the two types of species differed significantly in total biomass, total leaf area, RGR, NAR, LAR, SLA and leaf number, and not in leaf area. Only light-demanding species differed significantly in height, root length, basal diameter, RMR, SMR, LMR and R/S ratio. The mean plasticity index of growth and biomass allocation were relatively higher than the morphological variables, with significant differences between the two groups. Our results showed that liana species respond differently to changing light environments and that light-demanding species exhibit higher plasticity. Such differences may affect the relative success of liana species in forest dynamics.     

阔叶红松林林隙结构与树种多样性关系研究

对阔叶红松林林隙与林下不同层次树种多样性进行比较,探讨林隙大小及发育阶段与树种多样性间的关系.结果表明,林隙与林下树种多样性存在显著差异（P<0.01）.与林下相比,林隙更新层树种多样性、丰富度和均匀度增大,生态优势度减小（P<0.01）;演替层树种多样性、丰富度和均匀度减小,生态优势度增大（P<0.0001）.林隙中不同层次树种多样性随着林隙大小呈现出相反的变化趋势.长白山阔叶红松林林隙演替层树种多样性和丰富度总体上呈单峰形变化,中等大小林隙（100～250 m²）中树种多样性和丰富度较高,生态优势度较小.而更新层树种多样性和均匀度在≥250 m²和<100 m²的林隙中最大,在200～250 m²的林隙中最小;生态优势度在200～250 m²的林隙中达到最大.更新层和演替层树种对林隙面积大小反应不同,有利于更新层幼苗建立的林隙面积并不是演替层幼树发育和成活的最适面积;随林隙年龄的增加,更新层和演替层树种多样性在不同层次上呈互补关系.     

Age‐related Crown Thinning in Tropical Forest Trees

Gap dynamics theory proposes that treefall gaps provide high light levels needed for regeneration in the understory, and by increasing heterogeneity in the light environment allow light‐demanding tree species to persist in the community. Recent studies have demonstrated age‐related declines in leaf area index of individual temperate trees, highlighting a mechanism for gradual changes in the forest canopy that may also be an important, but less obvious, driver of forest dynamics. We assessed the prevalence of age‐related crown thinning among 12 tropical canopy tree species sampled in lowland forests in Panama and Puerto Rico (total N = 881). Canopy gap fraction of individual canopy tree crowns was positively related to stem diameter at 1.3 m (diameter at breast height) in a pooled analysis, with 10 of 12 species showing a positive trend. Considered individually, a positive correlation between stem diameter and canopy gap fraction was statistically significant in 4 of 12 species, all of which were large‐statured canopy to emergent species: Beilschmiedia pendula, Ceiba pentandra, Jacaranda copaia, and Prioria copaifera. Pooled analyses also showed a negative relationship between liana abundance and canopy gap fraction, suggesting that lianas could be partially obscuring age‐related crown thinning. We conclude that age‐related crown thinning occurs in tropical forests, and could thus influence patterns of tree regeneration and tropical forest community dynamics.     

Genetic variation in circadian regulation of nocturnal stomatal conductance enhances carbon assimilation and growth

Circadian resonance, whereby a plant's endogenous rhythms are tuned to match environmental cues, has been repeatedly shown to be adaptive, although the underlying mechanisms remain elusive. Concomitantly, the adaptive value of nocturnal transpiration in C₃ plants remains unknown because it occurs without carbon assimilation. These seemingly unrelated processes are interconnected because circadian regulation drives temporal patterns in nocturnal stomatal conductance, with maximum values occurring immediately before dawn for many species. We grew individuals of six Eucalyptus camaldulensis genotypes in naturally lit glasshouses and measured sunset, predawn and midday leaf gas exchange and whole‐plant biomass production. We tested whether sunrise anticipation by the circadian clock and subsequent increases in genotype predawn stomatal conductance led to rapid stomatal opening upon illumination, ultimately affecting genotype differences in carbon assimilation and growth. We observed faster stomatal responses to light inputs at sunrise in genotypes with higher predawn stomatal conductance. Moreover, early morning and midday stomatal conductance and carbon assimilation, leaf area and total plant biomass were all positively correlated with predawn stomatal conductance across genotypes. Our results lead to the novel hypothesis that genotypic variation in the circadian‐regulated capacity to anticipate sunrise could be an important factor underlying intraspecific variation in tree growth.     

Ecological outcomes of agroforests and restoration 15 years after planting

Large‐scale forest restoration relies on approaches that are cost‐effective and economically attractive to farmers, and in this context agroforestry systems may be a valuable option. Here, we compared ecological outcomes among (1) 12–15‐year‐old coffee agroforests established with several native shade trees, (2) 12–15‐year‐old high‐diversity restoration plantations, and (3) reference old‐growth forests, within a landscape restoration project in the Pontal do Paranapanema region, in the Atlantic Forest of southeastern Brazil. We compared the aboveground biomass, canopy cover, and abundance, richness, and composition of trees, and the regenerating saplings in the three forest types. In addition, we investigated the landscape drivers of natural regeneration in the restoration plantations and coffee agroforests. Reference forests had a higher abundance of trees and regenerating saplings, but had similar levels of species richness compared to coffee agroforests. High‐diversity agroforests and restoration plantations did not differ in tree abundance. However, compared to restoration plantations, agroforests showed higher abundance and species richness of regenerating saplings, a higher proportion of animal‐dispersed species, and higher canopy cover. The abundance of regenerating saplings declined with increasing density of coffee plants, thus indicating a potential trade‐off between productivity and ecological benefits. High‐diversity coffee agroforests provide a cost‐effective and ecologically viable alternative to high‐diversity native tree plantations for large‐scale forest restoration within agricultural landscapes managed by local communities, and should be included as part of the portfolio of reforestation options used to promote the global agenda on forest and landscape restoration.     

Using network connectance and autonomy analyses to uncover patterns of photosynthetic responses in tropical woody species

Daily courses of leaf gas exchange and chlorophyll fluorescence in forest gap and understorey environments were used to build photosynthetic networks in two pioneers and two late-successional species. Photochemical and gas exchange networks were linked to each other by the relationship between electron transport rate and net CO₂ assimilation. Global network connectance (Cg), which represents the mean strength of connections within a given network, was calculated in the photochemical and gas exchange networks for both functional groups and environments. Autonomy in relation to environmental fluctuations was estimated considering the mean correlation between environmental and physiological data. Cg was consistently higher in plants under gap condition. High daily-amplitude of environmental variables in the gap induced strong connectance in photochemical and gas exchange networks regardless of functional group. Gap scenario demands network modulation with higher level of control than understorey, which would be attained by strong connections among components of photochemical and gas exchange networks. This would allow fine and fast tuning adjustments when facing highly variable and demanding environmental conditions throughout a day. As a consequence of this highly variable environment, both functional groups showed lower autonomy in the gap, where higher coupling between leaf physiology and environmental fluctuations was evident. Our results suggest that high plant–environment coupling demands high network connectance. Contrastingly, Cg was lower (especially in photochemical network) under forest understorey, promoting autonomy in a more stable environment. Our results indicate that there is a conservative pattern of photosynthesis control based on network modulation and environmental coupling. This suggests that changes in network connectance may not be specific of a functional group but rather a more general response to environmental fluctuations, strongly related to system stability. We consider this information crucial in understanding how complex adaptive systems deal with environmental fluctuations.     

Effect of canopy gap light environment on evaporative load and stomatal conductance in the temperate forest understory herb Aster macrophyllus (Asteraceae)

Aster macrophyllus, a temperate forest understory species of the northeastern United States, inhabits a broad range of light habitats. Plants receiving several minutes of direct sun in canopy gap and forest edge habitats occasionally wilt, a response indicative of water stress. We compared two alterative scenarios for patterns of evaporative load and stomatal conductance for plants in large (0.15 ha) tree canopy gaps and small (3 m²) herbaceous subcanopy gaps: 1) evaporative loads are typically moderate and stomatal conductance is largely governed by light intensity; or 2) evaporative loads are often substantial, mandating stomatal closure to prevent excessive transpiration. In all cases evaporative loads were elevated by light intensity above 25% of full sun. This was accompanied by substantial stomatal closure. Transitions from low to moderate light intensity (<13% full sun) caused little increase in leaf evaporative load, and stimulated increases in stomatal conductance. Very brief periods of high light also stimulated stomatal opening. Light environments in the small herbaceous subcanopy gaps differ greatly in their patterns of evaporative load from day to day.     

Herbivory in canopy gaps created by a typhoon varies by understory plant leaf phenology

1. Availabilities of light and soil nitrogen for understory plants vary by extent of canopy gap formation through typhoon disturbance. We predicted that variation in resource availability and herbivore abundance in canopy gaps would affect herbivory through variation in leaf traits among plant species. We studied six understory species that expand their leaves before or after canopy closure in deciduous forests. We measured the availabilities of light, soil nitrogen, soil water content, and herbivore abundance in 20 canopy gaps (28.3–607.6 m²) formed by a typhoon and in four undisturbed stands. We also measured leaf traits and herbivory on understory plants. 2. The availabilities of light and soil nitrogen increased with increasing gap size. However, soil water content did not. The abundance of herbivorous insects (such as Lepidoptera and Orthoptera) increased with increasing gap size. 3. Concentrations of condensed tannins, total phenolics, and nitrogen in leaves and the leaf mass per area increased in late leaf expansion species with increasing gap size, whereas none of the leaf traits varied by gap size in early leaf expansion species. 4. Herbivory increased on early leaf expansion species with increasing gap size, but decreased on late leaf expansion species. In these late leaf expansion species, total phenolics and C : N ratio had negative relationships with herbivory. 5. These results suggested that after typhoon disturbance, increased herbivory on early leaf expansion species can be explained by increased herbivore abundance, whereas decreased herbivory on late leaf expansion species can be explained by variation in leaf traits.     

Mycorrhizal,Endophytic and Ecomorphological Status of Tree Roots in the Canopy of a Montane Rain Forest

Our investigation provides data on the symbiotic status and the functional morphology of tree fine roots in canopy soils of an upper montane forest in Costa Rica. Canopy roots completely lacked mycorrhizal colonization, but showed a high abundance of endophytic fungi suggesting a reduced ability for nutrient acquisition compared with terrestrial roots.     

Growth responses to arbuscular mycorrhizae by rain forest seedlings vary with light intensity and tree species

Light intensity and root colonization by arbuscular mycorrhizal (AM) fungi are considered important factors affecting the performance of rain forest plants, yet few studies have examined how these two factors interact. Whether AM colonization promoted growth or caused shifts in biomass allocation in seedlings of four species of Australian rain forest tree (Flindersia brayleana, Acmena resa, Cryptocarya mackinnoniana and Cryptocarya angulata), grown in a glasshouse under light conditions that mimicked the shaded understory (3% PAR) and small light gaps (10% PAR), was examined. Seedlings were grown in sterilized field soil and either inoculated with AM fungi or provided sterile inoculum. Four major findings emerged. First, in all species, seedlings grown in small gap light intensities were larger than seedlings grown in understory light intensities. Second, when seedling biomass was included as a covariate, variation in light intensity was associated with significant shifts in biomass allocation. In all species, leaf area ratio was lower at 10% PAR than at 3% PAR, while root-to-shoot ratio showed the opposite pattern in one of the four species (C. mackinonniana). Third, although percentage root length colonized by AM fungi was greater at 10% PAR than 3% PAR in all species, this difference could be accounted for by variation in seedling size in all species except C. angulata. Fourth, growth and biomass allocation responses to AM colonization varied with light intensity and plant species. AM colonization promoted growth in both light regimes only in F. brayleana, while it had no effect on growth in C. mackinnoniana and C. angulata in either light regime and promoted growth only under high light in A. resa. AM colonization had no effect on leaf area ratio or root-to-shoot ratio in any of the species, and significantly altered specific root length in only one of the four species (C. mackinnoniana). These findings suggest that rain forest seedlings are highly variable in their growth responses to AM colonization and that some of this variability is related to the light intensity of the environment. Given that seedlings may spend many years in the shaded understory, these differences among species could have important effects on long-term seedling performance and seedling community dynamics.