To study the acclimation responses of the leaves of saplings of six tree species when changed to low or high levels of irradiance, we carried out a light exposure experiment. Species representative of contrasting shade tolerance groups were identified across a light gradient in the understorey of a Venezuelan Andean cloud forest. Measured traits included gas exchange, chlorophyll fluorescence, and morphoanatomical, biochemical and optical properties. Saplings were grown for 6 months in a shade-house receiving 20% photosynthetic photon flux (PPF) of full sunlight. Plant samples were then moved to shade-houses receiving low PPF (4%) or high PPF (65%). A factorial model (species × PPF), with repeated measurements (0, 15 and 120 days) was designed. Our results showed that morphological and anatomical traits were more plastic to PPF changes than photosynthetic traits. All species were susceptible to photoinhibition (15 days): shade-intolerant species showed dynamic photoinhibition (120 days), whereas shade-tolerant species presented chronic photoinhibition and the consequent inability to increase C assimilation rates under high PPF. The partially shade-tolerant species showed mixed responses; nonetheless, they exhibited larger adjustments in morphoanatomical and optical properties. Thus the acclimation responses of these species when subject to contrasting light conditions could help to explain their distribution along the light gradient in the understorey.
Cloud forests represent a minor portion of the world forests, though outstanding in terms of biodiversity, endemisms, and environmental services provided. Understanding the factors that drive the regeneration and species composition of these forests, and in particular how light availability affects the patterns of juvenile tree distribution in the understory, is critical for conservation and restoration programs. In this study, we determined the range-size and overlap of the abundance distribution of juveniles for 20 tree species in an Andean tropical cloud forest in Venezuela along a gradient of percentage canopy openness (%CO) used as a surrogate of light availability. The observed distribution of %CO was then compared with a bounded null model of community structure in order to test light partitioning as a driver of tree species' coexistence. We measured %CO using hemispherical photography and the abundance and size of juvenile trees in 280 plots of 1-m radius spread over a 32 ha forest area. The distribution of sites was skewed towards the lower end of the %CO gradient (0.5 to 12.8%), while species abundance sharply diminished at both ends of the gradient. Nevertheless, 15 out of 20 species had a non-random distribution in relation to %CO, with many species concentrated near the lower side of the gradient. The observed pattern of species' overlap was within the 95% confidence limits for the average overlap expected under the bounded null model. These patterns indicate that low canopy openness is the rule in this forest, in spite of the scattered tree-fall gaps, and suggest that light partitioning does not determine the tree community structure at the juvenile stage. High redundancy in light requirements among juveniles of tree species may have a positive effect on species coexistence in cloud forests, thus maintaining a high species diversity. However, other factors such as recruitment limitation and differential growth/carbon-gain among species at the juvenile stage along the light gradient could contribute to the high diversity of these ecosystems.
Tropical cloud forests play a fundamental role in the hydrological cycle of mountain watersheds having the largest biodiversity per unit area. In Venezuela, cloud forests are subject to intense deforestation and fragmentation by farming and cattle-ranching causing soil erosion, water cycle alteration, and biodiversity loss. Reforestation projects used exotic species as Pines and Eucalyptus, native species were rarely planted by lacking knowledge on species requirements and management. We report the performance of 25 native cloud forest species differing in shade-tolerance, planted in mixed assemblies on degraded areas. Tree survival and the individual tree variables: total height, root-collar diameter, tree-slenderness, and crown-ratio were evaluated at 1, 2, 4.5 and 7 years-old. Data was analyzed with a repeated measures analysis of variance mixed model considering species shade-tolerance, light intensity at planting and age as explanatory factors. Survival was over 80%. Shade-intolerant species displayed faster height and root-collar diameter growth. Shade-tolerant species had larger crown ratios due to persistence of lower branches; whereas, shade-intolerant showed signs of crown recession at age 7. Slenderness values from age 4.5 were indicative of good trees stability and health across treatments. The positive results have motivated landowners to establish native species plantations in critical areas with our support.
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