Effects of partial throughfall exclusion on the phenology of Coussarea racemosa (Rubiaceae) in an east-central Amazon rainforest
Severe droughts may alter the reproductive phenology of tropical tree species, but our understanding of these effects has been hampered by confounded variation in drought, light and other factors during natural drought events. We used a large-scale experimental reduction of throughfall in an eastern-central Amazon forest to study the phenological response to drought of an abundant subcanopy tree, Coussarea racemosa. We hypothesized that drought would alter the production and the timing of reproduction, as well as the number of viable fruits. The study system comprised two 1-ha plots in the Tapajos National Forest, Para, Brazil: a dry plot where 50% of incoming precipitation (80% throughfall) was diverted from the soil during the six-month wet season beginning in January 2000, and a wet plot that received natural rainfall inputs. Fruit production of C. racemosa was quantified every 15 days using 100 litter traps (0.5 m(2)) in each plot. The production of new leaves and flowers was recorded monthly for C. racemosa individuals. Soil water, pre-dawn leaf water potential and solar radiation were measured to help interpret phenological patterns. Over the -3.5-year period (April 2000 through December 2003), total fruit production remained similar between plots, declining by 12%. In 2003, production was four times higher in both plots than in previous years. In the dry plot, fruit fall shifted 40 and 60 days later into the dry season in 2002 and 2003, respectively. Total fruit fall dry mass production was variable across the study period. Foliage and flower production coincided with peak irradiance early in the dry season until delays in flowering appeared in the dry plot in 2002 and 2003. Plant water stress, through its influence on leaf developmental processes and, perhaps, inhibition of photosynthesis, appears to have altered both the timing of fruit fall and the quality and number of seeds produced.