LIFE HISTORY DIVERSITY OF CANOPY AND EMERGENT TREES IN A NEOTROPICAL RAIN FOREST

To assess the diversity of tropical tree life histories, a conceptual framework is needed to guide quantitative comparative study of many species. We propose one such framework, which focuses on long—term performance through ontogeny and over the natural range of microsites. For 6 yr we annually evaluated survival, growth, and microsite conditions of six non—pioneer tree species in primary tropical wet forest at the La Selva Biological Station, Costa Rica. The species were: Lecythis ampla, Hymenolobium mesoamericanum, Dipteryx panamensis, Pithecellobium elegans, Hyeronima alchorneoides (all emergents), and Minquartia guianensis (a canopy species). The study was based on long—term measurement of individuals from all post—seedling size classes. Trees were sampled from 150 ha of primary forest spanning several watersheds and soil types. To evaluate individuals' microsites we recorded the number of overtopping crowns, forest phase (gap, building, mature), and crown illumination index (an estimate of the tree's light environment). For comparison, we also evaluated the microsites of three species that have been categorized as pioneers (Cecropia insignis, C. obtusifolia) or high—light demanders (Simarouba amara). For the six species of non—pioneers, mortality rates declined with increasing juvenile size class. As a group, these emergent and canopy trees showed a much lower exponential annual mortality rate (0.44%/yr at >10 cm diameter) than has been found for the La Selva forest as a whole. Growth rates increased with juvenile size class for all six species. As adults (trees >30 cm in diameter), all five emergent species showed substantial annual diameter increments (medians of 5—14 mm/yr). Small saplings and adults of all species had significant year—to—year variation in diameter growth, with much greater growth occurring in the year of lowest rainfall. Passage time analysis suggests that all six species require >150 yr for growth from small saplings to the canopy. Evaluation of all nine species revealed four patterns of microsite occupancy by juveniles. Among the non—pioneers, one species pair (Lecythis and Minquartia: Group A) was associated with low crown illumination and mature—phase forest in all juvenile stages. For two species (Dipteryx and Hymenolobium: Group B) the smallest saplings were in predominantly low—light, mature—forest sites, but crown illumination and association with gap— or building—phase sites increased with juvenile size (Simarouba also showed this pattern). Two species (Pithecellobium and Hyeronima: Group C) were strongly associated with gap or building phase as small juveniles (≤4 cm diameter) and again as subcanopy trees (>10—20 cm diameter), but were predominantly in mature—phase sites at intermediate sizes. Juveniles of the two pioneer species (Cecropia: Group D) showed the highest crown illumination and association with gap or building sites. Among the six non—pioneer species, only one aspect of juvenile performance clearly varied according to microsite group. The smallest saplings (≤1 cm diameter) of Groups B and C showed significant mortality differences across a small gradient in crown illumination; neither of the Group A species showed this pattern. Otherwise, juvenile performance was strikingly similar among the six species. All showed a capacity for growth responses to small increases in light, substantial height and diameter increments at higher light levels, equal ability to survive 4—yr periods of no growth, and very low mortality rates at intermediate—to—large juvenile sizes. Species differed significantly in growth rates, but relative differences shifted with tree size and were unrelated to microsite group. These findings do not support prevailing paradigms concerning trade—offs and correlated suites of traits. For non—pioneer tropical trees, life history classification based on generalized concepts such as gap dependence and shade tolerance is inadequate to describe the complex size—dependent patterns of life history differences and similarities that exist among species.