Causes and consequences of resource heterogeneity in forests: interspecific variation in light transmission by canopy trees

We have analyzed the light transmission characteristics of the nine deciduous and coniferous species that dominate the transition oak–northern hardwood forests of southern New England. Maximum likelihood techniques were used to estimate species-specific light extinction coefficients, using fish-eye photography combined with data on the locations and geometry of trees in the neighborhood around each photo point. Quantum sensors were also used to quantify interspecific variation in the importance of sunflecks and beam enrichment. Variation in light extinction was closely correlated with shade tolerance and successional status of the species. The most shade-tolerant species (Fagusgrandifolia Ehrh. and Tsugacanadensis (L.) Carr.) cast the deepest shade (<2% of full sun), while earlier successional species such as Quercusrubra L. and Fraxinusamericana L. allowed greater light penetration (>5% full sun). These differences were more closely related to differences in crown depth than to differences in light extinction per unit depth of crown. Sunflecks contributed relatively little radiation beneath late successional species (<10% of total understory photosynthetically active radiation), but represented a major fraction (40–50%) of radiation beneath less shade-tolerant species. Using growth and mortality functions for the same species developed in a related study, our results indicate that saplings of all of the species have high survivorship in the shade cast by conspecific adults. However, only the three most shade-tolerant species have low rates of sapling mortality under the low light levels characteristic of stands dominated by late successional species. Our results are consistent with previously reported models, which propose that secondary succession is driven by interspecific differences in resource uptake and tolerance.