Responses of floodplain forest species to spatially condensed gradients: a test of the flood-shade tolerance tradeoff hypothesis.

Previous work in southeastern US floodplains led to the hypothesis that a tradeoff between flood and shade tolerance underlies species-specific responses to flooding and light, which drive forest regeneration. In systems where community turnover can occur with small-scale environmental changes, testing this hypothesis requires recognizing that turnover of species along the two gradients can be large relative to the spatial distances involved. We test the tradeoff hypothesis in an old-growth bottomland hardwood forest by (1) comparing shade and moisture profiles of woody juveniles versus random points and (2) using individual-based sampling of woody juveniles to model probability of occurrence in response to distance-to-water table and canopy openness gradients. We found that juveniles of all species combined occupied a similar range of distance-to-water table compared to measurements taken at random points, but average canopy openness above seedlings was significantly higher than at random points. On average, shade-tolerant species, with the exception of Acer rubrum, were found in shaded areas that were also drier, whereas less shade-tolerant taxa, plus A. rubrum, were found in wetter, more open areas, suggesting a tradeoff between flood and shade tolerance. Predictive models of species occurrence, which incorporate the availability of canopy and microtopographic conditions, indicated that three taxa (Fraxinus pennsylvanica, Quercus spp., and Ulmus americana) had patterns consistent with a flood-shade tolerance tradeoff. In contrast, Asimina triloba, Celtis laevigata, and Liquidambar styraciflua had positive responses when the joint stresses of flooding and shade were diminished. A. rubrum appeared to be the most tolerant to both stresses. Our work not only lends support to the flood-shade tradeoff hypothesis but also indicates that a more general model is needed that includes a "flood-shade release" component. We also suggest that responses to small-scale gradients should not be overlooked, particularly in systems where the gradients are ecologically steep.