JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org.. Ecological Society of America is collaborating with JSTOR to digitize, preserve and extend access to Ecology.Abstract. Many trees of tropical dry forests flower or form new shoots soon after leaf shedding during the dry season, i.e., during a period when trees are likely to be severely water stressed. To resolve this apparent paradox, phenology and seasonal changes in tree water status were monitored during two consecutive dry seasons in > 150 trees of 37 species growing at different sites in the tropical dry lowland forest of Guanacaste, Costa Rica.Tree development during the dry season varied considerably between species and between sites of different moisture availability. Leaf shedding, flowering, and shoot growth (flushing) were strongly correlated with seasonal changes in tree water status, measured by conventional and newly developed techniques. Tree water status varied with the availability of subsoil water and a variety of biotic factors such as structure and life-span of leaves, time of leaf shedding, wood density and capacity for stem water storage, and depth and density of root systems. Observed tree species differed widely in wood density (from 0.19 to 1.1 g/cm3) and stem water storage capacity (400-20% of dry mass), which was highly correlated with the degree of desiccation during drought. Only hardwood trees at dry upland sites, lacking stem water storage and access to subsoil water, desiccated strongly (stem water potential <-4 MPa) and remained inactive throughout the dry season. In all other trees elimination of transpirational water loss during leaf shedding in conjunction with utilization of residual water enabled rehydration of stem tissues and subsequent flowering or flushing during the dry season. Rate and degree of rehydration varied strongly with the availability of water stored in tree trunks or in the subsoil. Stored water buffers the impact of seasonal drought and enables flowering and flushing during the dry season. Climatic data are thus not sufficient to explain tree phenology in seasonally dry tropical forests.Phenology, seasonal changes in water status, and water storage capacity of tree species are highly correlated. Tree species cluster into a number of distinct functional types ranging from deciduous hardwood trees and water-storing lightwood trees in dry upland forests to evergreen light-and softwood trees confined to moist lowland sites. Seasonal variation in tree water status thus appears to be the principal determinant of both phenology and distribution of tree species in tropical dry forests.
In many conspecific trees of >50 species highly synchronous bud break with low inter-annual variation was observed during the late dry season, around the spring equinox, in semideciduous tropical forests of Argentina, Costa Rica, Java and Thailand and in tropical savannas of Central Brazil. Bud break was 6 months out of phase between the northern and southern hemispheres and started about 1 month earlier in the subtropics than at lower latitudes. These observations indicate that "spring flushing", i.e., synchronous bud break around the spring equinox and weeks before the first rains of the wet season, is induced by an increase in photoperiod of 30 min or less. Spring flushing is common in semideciduous forests characterized by a 4-6 month dry season and annual rainfall of 800-1,500 mm, but rare in neotropical forests with a shorter dry season or lower annual precipitation. Establishment of new foliage shortly before the wet growing season is likely to optimize photosynthetic gain in tropical forests with a relatively short growing season.
In tropical rainforests, 30-65% of tree species grow at densities of less than one individual per hectare. At these low population densities, successful cross-pollination relies on synchronous flowering. In rainforests with low climatic seasonality, photoperiodic control is the only reliable mechanism for inducing synchronous flowering. This poses a problem because there is no variation in day length at the Equator. Here we propose a new mechanism of photoperiodic timekeeping based on the perception of variation in sunrise or sunset time, which explains and predicts the annually repeated, staggered, synchronous and bimodal flowering of many tree species in Amazonian rainforests near the Equator.
The seasonal time course of vegetative phenology and cambium growth is compared for tree species from Central America and Asia growing in tropical climates with a long, severe dry season. Although the inhibition of plant growth by water stress is weH established, responses to seasonal drought vary widely among such trees, and their annual development is not weH synchronized by climatic seasonality. In deciduous trees growing at microsites with low soil moisture storage, phenology and cambium growth are weH correlated with each other and with seasonal rainfall, and most trees have distinct annual rings. Phenology and cambium growth are progressively uncoupled from climatic seasonality in brevideciduous and evergreen trees growing at microsites with large soil water reserves which buffer trees against seasonal drought and thus may prevent the formation of distinct annual rings. There is some experimental evidence conceming the control of growth initiation in apical meristems and the cambium, but little is known about the mechanisms which arrest growth and deterrnine qualitative changes in organ development and cambium cell differentiation.
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