Shade tolerance

Among 13 tropical tree species on Barro Colorado Island, species with high seedling mortality rates during the first year in shade had higher reltive growth rates (RGR) from germination to 2 months in both sun (23% full sun) and shade [2%, with and without lowered red: far red (R:FR) ratio] than shade tolerant species. Species with higher RGR in sun also had higher RGR in shade. These interspecific trends could be explained by differences in morphological traits and allocation paterns among species. Within each light regime, seedlings of shade-intolerant species had lower root: shoot ratios, higher leaf mass per unit area, and higher leaf area ratios (LAR) than shade tolerant species. In contrast, leaf gas exchange characteristics, or acclimation potential in these traits, had no relationship with seedling mortality rates in shade. In both shade tolerant and intolerant species, light saturated photosynthesis rates, dark respiration, and light compensation points were higher for sungrown seedlings than for shade-grown seedlings. Differences in R:FR ratio in shade did not affect gas exchange, allocation patterns, or growth rates of any species. Survival of young tree seedlings in shade did not depend on higher net photosynthesis or biomass accumulation rates in shade. Rather, species with higher RGR died faster in shade than species with lower RGR. This trend could be explained if survival depends on morphological characteristics likely to enhance defense against herbivores and pathogens, such as dense and tough leaves, a well-established root system, and high wood density. High construction costs for these traits, and low LAR as a consequence of these traits, should result in lower rates of whole-plant carbon gain and RGR for shade tolerant species than shade-intolerant species in shade as well as in sun.

Large-scale patterns of Amazonian biodiversity have until now been obscured by a sparse and scattered inventory record. Here we present the first comprehensive spatial model of tree α-diversity and tree density in Amazonian rainforests, based on the largest-yet compilation of forest inventories and bolstered by a spatial interpolation technique that allows us to estimate diversity and density in areas that have never been inventoried. These data were then compared to continent-wide patterns of rainfall seasonality. We find that dry season length, while only weakly correlated with average tree α-diversity, is a strong predictor of tree density and of maximum tree α-diversity. The most diverse forests for any given DSL are concentrated in a narrow latitudinal band just south of the equator, while the least diverse forests for any given DSL are found in the Guayana Shield and Amazonian Bolivia. Denser forests are more diverse than sparser forests, even when we used a measure of diversity that corrects for sample size. We propose that rainfall seasonality regulates tree α-diversity and tree density by affecting shade tolerance and subsequently the number of different functional types of trees that can persist in an area.

Maximum Rubisco activities (V(cmax)), rates of photosynthetic electron transport (J(max)), and leaf nitrogen and chlorophyll concentrations were studied along a light gradient in the canopies of four temperate deciduous species differing in shade tolerance according to the ranking: Populus tremula L. < Fraxinus excelsior L. < Tilia cordata Mill. = Corylus avellana L. Long-term light environment at the canopy sampling locations was characterized by the fractional penetration of irradiance in the photosynthetically active spectral region (I(sum)). We used a process-based model to distinguish among photosynthesis limitations resulting from variability in fractional nitrogen investments in Rubisco (P(R)), bioenergetics (P(B), N in rate-limiting proteins of photosynthetic electron transport) and light harvesting machinery (P(L), N in chlorophyll and thylakoid chlorophyll-protein complexes). On an area basis, V(cmax) and J(max) (V(a) (cmax) and J(a) (max)) increased with increasing ...

1Water availability is the main determinant of species’ distribution in lowland tropical forests. Species’ occurrence along water availability gradients depends on their ability to tolerate drought.2To identify species’ traits underlying drought-tolerance we excavated first year seedlings of 62 dry and moist forest tree species at the onset of the dry season. We evaluate how morphological seedling traits differ between forests, and whether functional groups of species can be identified based on trait relations. We also compare seedling traits along independent axes of drought and shade-tolerance to assess a hypothesized trade-off.3Seedlings of dry forest species improve water foraging capacity in deep soil layers by an increased below-ground biomass allocation and by having deep roots. They minimize the risk of cavitation by making dense stems, and reduce transpiration by producing less leaf tissue. Moist forest seedlings have large leaf areas and a greater above-ground biomass, to maximize light interception, and long, cheap, branched root systems, to increase water and nutrient capture.4Associations among seedling traits reveal three major drought strategies: (i) evergreen drought-tolerant species have high biomass investment in enduring organs, minimize cavitation and minimize transpiration to persist under dry conditions; (ii) drought-avoiding species maximize resource capture during a limited growing season and then avoid stress with a deciduous leaf habit in the dry season; (iii) drought-intolerant species maximize both below- and above-ground resource capture to increase competitiveness for light, but are consequently precluded from dry habitats.5We found no direct trade-off between drought- and shade-tolerance, because they depend largely on different morphological adaptations. Drought-tolerance is supported by a high biomass investment to the root system, whereas shade-tolerance is mainly promoted by a low growth rate and low SLA.6Synthesis. We conclude that there are three general adaptation strategies of drought-tolerance, which seemingly hold true across biomes and for different life forms. Drought- and shade-tolerance are largely independent from one another, suggesting a high potential for niche differentiation, as species’ specialization can occur at different combinations of water and light availability.Water availability is the main determinant of species’ distribution in lowland tropical forests. Species’ occurrence along water availability gradients depends on their ability to tolerate drought.To identify species’ traits underlying drought-tolerance we excavated first year seedlings of 62 dry and moist forest tree species at the onset of the dry season. We evaluate how morphological seedling traits differ between forests, and whether functional groups of species can be identified based on trait relations. We also compare seedling traits along independent axes of drought and shade-tolerance to assess a hypothesized trade-off.Seedlings of dry forest species improve water foraging capacity in deep soil layers by an increased below-ground biomass allocation and by having deep roots. They minimize the risk of cavitation by making dense stems, and reduce transpiration by producing less leaf tissue. Moist forest seedlings have large leaf areas and a greater above-ground biomass, to maximize light interception, and long, cheap, branched root systems, to increase water and nutrient capture.Associations among seedling traits reveal three major drought strategies: (i) evergreen drought-tolerant species have high biomass investment in enduring organs, minimize cavitation and minimize transpiration to persist under dry conditions; (ii) drought-avoiding species maximize resource capture during a limited growing season and then avoid stress with a deciduous leaf habit in the dry season; (iii) drought-intolerant species maximize both below- and above-ground resource capture to increase competitiveness for light, but are consequently precluded from dry habitats.We found no direct trade-off between drought- and shade-tolerance, because they depend largely on different morphological adaptations. Drought-tolerance is supported by a high biomass investment to the root system, whereas shade-tolerance is mainly promoted by a low growth rate and low SLA.Synthesis. We conclude that there are three general adaptation strategies of drought-tolerance, which seemingly hold true across biomes and for different life forms. Drought- and shade-tolerance are largely independent from one another, suggesting a high potential for niche differentiation, as species’ specialization can occur at different combinations of water and light availability.

1To survive in forest understoreys, seedlings must depend on carbohydrate reserves when they experience negative carbon balance imposed by occasional light reduction and tissue loss to herbivores and diseases. We present the first experimental evidence in support of this hypothesis, using seven woody neotropical species.2We transplanted seedlings that had recently expanded their first photosynthetic cotyledon or leaf to the forest understorey (1% of full sun) and quantified initial biomass and total non-structural carbohydrate (TNC) in stems, roots and storage cotyledons. We then randomly assigned seedlings to control and two stress treatments: light reduction (0.08% of full sun for 8 weeks) and complete defoliation.3First-year survival of control seedlings, a comparative measure of shade tolerance, differed widely among species. The two stress treatments reduced survival and relative growth rates (RGR) of all species. Shade-tolerant species were little impacted by the stress treatments, whereas the two least shade-tolerant species experienced 100% mortality.4In all treatments, 8-week and first-year survival was positively correlated with initial TNC pool size in stems and roots. By contrast, survival was generally not correlated with initial TNC concentration in any organ, TNC pools in cotyledons, seed mass or seedling biomass.5TNC in stems and roots, but not in cotyledons, decreased in response to light reduction and defoliation over 8 weeks. Leaf area recovery of defoliated seedlings was positively correlated with initial TNC pools in stems and roots.6First-year survival in each treatment was negatively correlated with 0–8 week RGR of control seedlings, suggesting higher stress tolerance of species with inherently slow growth rates in shade. RGR of control seedlings from 0 to 8 weeks was negatively correlated with initial TNC pools, but not concentrations, in stems and roots. After 8 weeks, RGR was positive for all species, without clear relationships with survival or TNC.7We conclude that carbohydrate storage in stems and roots enhances long-term survival in shade by enabling seedlings to cope with periods of biotic and abiotic stress. Carbohydrate storage is a key functional trait that can explain species differences in growth and survival that lead to species coexistence through niche assembly processes and life-history trade-offs.To survive in forest understoreys, seedlings must depend on carbohydrate reserves when they experience negative carbon balance imposed by occasional light reduction and tissue loss to herbivores and diseases. We present the first experimental evidence in support of this hypothesis, using seven woody neotropical species.We transplanted seedlings that had recently expanded their first photosynthetic cotyledon or leaf to the forest understorey (1% of full sun) and quantified initial biomass and total non-structural carbohydrate (TNC) in stems, roots and storage cotyledons. We then randomly assigned seedlings to control and two stress treatments: light reduction (0.08% of full sun for 8 weeks) and complete defoliation.First-year survival of control seedlings, a comparative measure of shade tolerance, differed widely among species. The two stress treatments reduced survival and relative growth rates (RGR) of all species. Shade-tolerant species were little impacted by the stress treatments, whereas the two least shade-tolerant species experienced 100% mortality.In all treatments, 8-week and first-year survival was positively correlated with initial TNC pool size in stems and roots. By contrast, survival was generally not correlated with initial TNC concentration in any organ, TNC pools in cotyledons, seed mass or seedling biomass.TNC in stems and roots, but not in cotyledons, decreased in response to light reduction and defoliation over 8 weeks. Leaf area recovery of defoliated seedlings was positively correlated with initial TNC pools in stems and roots.First-year survival in each treatment was negatively correlated with 0–8 week RGR of control seedlings, suggesting higher stress tolerance of species with inherently slow growth rates in shade. RGR of control seedlings from 0 to 8 weeks was negatively correlated with initial TNC pools, but not concentrations, in stems and roots. After 8 weeks, RGR was positive for all species, without clear relationships with survival or TNC.We conclude that carbohydrate storage in stems and roots enhances long-term survival in shade by enabling seedlings to cope with periods of biotic and abiotic stress. Carbohydrate storage is a key functional trait that can explain species differences in growth and survival that lead to species coexistence through niche assembly processes and life-history trade-offs.

Water availability is the most important factor determining tree species distribution in the tropics, but the underlying mechanisms are still not clear. In this study, we compared functional traits of 38 tropical tree species from dry and moist forest, and quantified their ability to survive drought in a dry-down experiment in which wilting and survival were monitored. We evaluated how seedling traits affect drought survival, and how drought survival determines species distribution along the rainfall gradient. Dry forest species tended to have compound leaves, high stem dry matter content (stem dry mass/fresh mass), and low leaf area ratio, suggesting that reduction of transpiration and avoidance of xylem cavitation are important for their success. Three functional groups were identified based on the seedling traits: (1) drought avoiders with a deciduous leaf habitat and taproots; (2) drought resisters with tough tissues (i.e., a high dry matter content); and (3) light-demanding moist forest species with a large belowground foraging capacity. Dry forest species had a longer drought survival time (62 d) than moist forest species (25 d). Deciduousness explained 69 percent of interspecific variation in drought survival. Among evergreen species, stem density explained 20 percent of the drought survival. Drought survival was not related to species distribution along the rainfall gradient, because it was mainly determined by deciduousness, and species with deciduous seedlings are found in both dry and moist forests. Among evergreen species, drought survival explained 28 percent of the variation in species position along the rainfall gradient. This suggests that, apart from drought tolerance, other factors such as history, dispersal limitation, shade tolerance, and fire shape species distribution patterns along the rainfall gradient.

We compared historical (1909-1913) and contemporary (1997-1999) forest struc- ture and composition on 15 permanent plots in ponderosa pine (Pinus ponderosa Dougl. ex Laws.) forests of Arizona and New Mexico. We used the same sampling methods as in the early 1900s and compared stand density, diameter distributions, species composition, and broad age classes from the two periods. Stand density (trees