its width (32). On physical grounds, the thin gas gap suggested by our measurements should also be expected to possess soft modes with fluctuations whose wavelength ranges from small to large. From this perspective, we then expect that the experimental geometry of a Janus-type water film, selected for experimental convenience, was incidental to the main physical effect.These conclusions have evident connections to understanding the long-standing question of the structure of aqueous films near a hydrophobic surface and may have a bearing on understanding the structure of water films near the patchy hydrophilichydrophobic surfaces that are so ubiquitous in nature.Note added in proof: We have recently been made aware of neutron reflectivity experiments that indicate the existence of a nanometer-thick vapor-like coating that forms on an extended hydrophobic surface when it is immersed in water (33, 34). The high alpha-diversity of tropical forests has been amply documented, but beta-diversity-how species composition changes with distance-has seldom been studied. We present quantitative estimates of beta-diversity for tropical trees by comparing species composition of plots in lowland terra firme forest in Panama, Ecuador, and Peru. We compare observations with predictions derived from a neutral model in which habitat is uniform and only dispersal and speciation influence species turnover. We find that beta-diversity is higher in Panama than in western Amazonia and that patterns in both areas are inconsistent with the neutral model. In Panama, habitat variation appears to increase species turnover relative to Amazonia, where unexpectedly low turnover over great distances suggests that population densities of some species are bounded by as yet unidentified processes. At intermediate scales in both regions, observations can be matched by theory, suggesting that dispersal limitation, with speciation, influences species turnover. References and NotesBeta-diversity is central to concepts about what controls diversity in ecological communities. Species turnover can reflect deterministic processes, such as species' adaptations to differences in climate or substrate, or it can result from limited dispersal coupled with speciation, delayed response to climatic change, or other historical effects. Perhaps more important, beta-diversity is as important as alpha-diversity for conservation, because species turnover influences diversity at large scales. Recently, Hubbell (1) and Harte et al. (2, 3) have derived theories relating species turnover with distance to species-area relations and total species richness. In very rich forests of the neotropics, these theories may allow us to interpolate species turnover and estimate species distributions and diversity at scales relevant to conservation even with the sparse data from forest plots that are currently available.To measure beta-diversity and test factors influencing it, we identified all trees in 34 plots near the Panama Canal, 16 plots in Ecuador's Yasuní National Park, and 1...
Light gap disturbances have been postulated to play a major role in maintaining tree diversity in species-rich tropical forests. This hypothesis was tested in more than 1200 gaps in a tropical forest in Panama over a 13-year period. Gaps increased seedling establishment and sapling densities, but this effect was nonspecific and broad-spectrum, and species richness per stem was identical in gaps and in nongap control sites. Spatial and temporal variation in the gap disturbance regime did not explain variation in species richness. The species composition of gaps was unpredictable even for pioneer tree species. Strong recruitment limitation appears to decouple the gap disturbance regime from control of tree diversity in this tropical forest.
A trade-off between growth and mortality rates characterizes tree species in closed canopy forests. This trade-off is maintained by inherent differences among species and spatial variation in light availability caused by canopy-opening disturbances. We evaluated conditions under which the trade-off is expressed and relationships with four key functional traits for 103 tree species from Barro Colorado Island, Panama. The trade-off is strongest for saplings for growth rates of the fastest growing individuals and mortality rates of the slowest growing individuals (r2 = 0.69), intermediate for saplings for average growth rates and overall mortality rates (r2 = 0.46), and much weaker for large trees (r2 < or = 0.10). This parallels likely levels of spatial variation in light availability, which is greatest for fast- vs. slow-growing saplings and least for large trees with foliage in the forest canopy. Inherent attributes of species contributing to the trade-off include abilities to disperse, acquire resources, grow rapidly, and tolerate shade and other stresses. There is growing interest in the possibility that functional traits might provide insight into such ecological differences and a growing consensus that seed mass (SM), leaf mass per area (LMA), wood density (WD), and maximum height (H(max)) are key traits among forest trees. Seed mass, LMA, WD, and H(max) are predicted to be small for light-demanding species with rapid growth and mortality and large for shade-tolerant species with slow growth and mortality. Six of these trait-demographic rate predictions were realized for saplings; however, with the exception of WD, the relationships were weak (r2 < 0.1 for three and r2 < 0.2 for five of the six remaining relationships). The four traits together explained 43-44% of interspecific variation in species positions on the growth-mortality trade-off; however, WD alone accounted for > 80% of the explained variation and, after WD was included, LMA and H(max) made insignificant contributions. Virtually the full range of values of SM, LMA, and H(max) occurred at all positions on the growth-mortality trade-off. Although WD provides a promising start, a successful trait-based ecology of tropical forest trees will require consideration of additional traits.
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