The effect of leaf age on photosynthetic capacity, a critical parameter in the theory of optimal leaf longevity, was studied for two tropical pioneer tree species, Cecropia longipes and Urera caracasana, in a seasonally dry forest in Panama. These species continuously produce short-lived leaves (74 and 93 d, respectively) during the rainy season (May-December) on orthotropic branches. However, they differ in leaf production rate, maximum number of leaves per branch, light environment experienced by the leaves, leaf mass per unit area, and nitrogen content. Light-saturated photosynthetic rates for marked leaves of known ages (±1 wk) were measured with two contrasting schemes (repeated measurements vs. chronosequence within branch), which overall produced similar results. In both species, photosynthetic rates and nitrogen use efficiency were negatively correlated with leaf age and positively correlated with light availability. Photosynthetic rates declined faster with leaf age in Cecropia than in Urera as predicted by the theory. The rate of decline was faster for leaves on branches with faster leaf turnover rates. Nitrogen per unit leaf area decreased with leaf age only for Urera. Leaf mass per unit area increased with leaf age, either partly (in Cecropia) or entirely (in Urera) due to ash accumulation.

SignificanceLeaf traits, such as photosynthetic capacity, nitrogen concentration, and leaf mass per area, strongly affect plant growth and nutrient cycles. Understanding relationships among leaf traits is, therefore, a fundamental challenge in plant biology, crop science, and ecology. Different groups of leaves exhibit distinct relationships among pairs of traits. For example, photosynthetic capacity per unit leaf area increases strongly with leaf mass per area from sun to shade within species, but these same traits are only weakly related across global species. Our analysis suggests that divergent trait relationships can be understood by partitioning leaf mass into photosynthetic and structural support components. Our paper clarifies the causes of relationships among traits and why those relationships differ among different groups of plants.

A severe outbreak of Lepidoptera followed the 1997–98 El Niño Southern Oscillation event, during which the climate in central Panama was unusually dry. The outbreak involved the larvae of at least 12 species of Lepidoptera and occurred at a seasonally dry, deciduous forest site, where extensive background data were available regarding climate, tree species and non-outbreak herbivory levels. Most Lepidoptera were associated with only one or two larval host plant species belonging to the same family, and the majority were monophagous during this study. During the outbreak, caterpillar densities for the major outbreak species averaged 1.6 larvae per young leaf and 0.18 larvae per leaf for leaves of all ages. For canopy trees and lianas, the mean level of leaf damage was 13.8%, ranging from 1–100%. Seven out of 20 tree species sustained most of the damage, with 21–37% of the leaf area consumed. Relative to non-outbreak years, damage levels increased by more than 250% during the outbreak. Single-species outbreaks were observed in other areas with a similar drought, but wetter forests in central Panama did not experience outbreaks during this period. Historically, fewer outbreaks have been reported from tropical forests than from temperate forests, however, similar El Niño-related outbreaks have occurred recently in several other locations throughout the tropics.

The importance of termites as decomposers in tropical forests has long been recognized. Studies on the richness and diversity of termite species and their ecological function have flourished in more recent times, but these have been mostly conducted in a thin stratum within a standing man's reach. Our aims were to evaluate the specific richness and composition of the termite assemblage in the canopy of a tropical rainforest and to determine its originality with respect to the sympatric ground-level fauna. We conducted systematic searches for canopy termites, together with conventional sampling of the sympatric ground-level fauna, in the San Lorenzo forest, Panama. We hypothesized that (1) the canopy accommodates two categories of wood-feeding termites (long-distance foragers and small-colony "one-piece" species) and possibly soil-feeders in suspended soil-like habitats; (2) due to the abundance of soil-feeders, the overall diversity of the ground fauna is higher than that of the canopy; (3) differences in microclimate and resource accessibility favour vertical stratification among wood-feeders. Sixty-three canopy samples yielded ten species of termites, all wood-feeders. Five of these were not found at ground level, although a total of 243 ground samples were collected, representing 29 species. In addition to long-distance foragers (Microcerotermes and Nasutitermes spp.) and small-colony termites (mostly Kalotermitidae), the canopy fauna included Termes hispaniolae, a wood-feeding Termitidae from an allegedly soil-feeding genus, living in large dead branches. Soil-feeders were absent from the canopy, probably because large epiphytes were scarce. As predicted, the ground fauna was much richer than that of the canopy, but the species richness of both habitats was similar when only wood-feeders were considered. Vertical stratification was strongly marked among wood-feeders, as all common species, apart from the arboreal-nesting Microcerotermes arboreus, could unequivocally be assigned to either a ground or a canopy group. The canopy, therefore, contributes significantly to the total species richness of the termite assemblage, and the diversity, abundance and ecological importance of canopy termites in tropical rainforests may be higher than previously recognized.

The Republic of Panama recently experienced a limited forest transition. After five decades of decline, the total forest cover increased by 0.36% yr-1 between 1992 and 2000; however, mature forest cover simultaneously decreased by 1.3% yr-1. This limited forest transition at the national scale comprised two distinctly different patterns of recent forest-cover change related to historical land use. Districts that were largely deforested when the first national survey of forest cover was completed in 1947 experienced a strong forest transition between 1992 and 2000. In these, the proportion of the population employed in agriculture decreased by an average of 31% and natural secondary forest succession increased the total forest cover by an average of 85% between 1992 and 2000. In contrast, no forest transition was evident for districts that were largely forested in 1947. In these, the absolute number of people employed in agriculture remained constant, old-growth forest cover decreased by 8% on average, and natural secondary forest succession increased, so that the total forest cover tended to be static between 1992 and 2000. Historical land use, an index of human poverty, and the population density of agricultural workers explained 61% of the among-district variation in forest cover in 2000, with forest concentrated in areas where populations were small and poor. Historical land use and gross income per hectare from agriculture explained 23.5% of the among-district variation in forest-cover change between 1992 and 2000. The early history of forest loss, an uneven distribution of people, and disparities in farm income contributed to the limited forest transition observed in Panama.