Following damage caused by Hurricane Hugo (September 1989) we monitored inorganic nitrogen availability in soil twice in 1990, leaf area index in 1991 and 1993, and litter production from 1990 through 1992 in subtropical wet forest of eastern Puerto Rico. Experimental removal of litter and woody debris generated by the hurricane (plus any standing stocks present before the hurricane) increased soil nitrogen availability and above-ground productivity by as much as 40% compared to unmanipulated control plots. These increases were similar to those created by quarterly fertilization with inorganic nutrients. Approximately 85% of hurricane-generated debris was woody debris >5 cm diameter. Thus, it appeared that woody debris stimulated nutrient immobilization, resulting in depression of soil nitrogen availability and productivity in control plots. This was further suggested by simulations of an ecosystem model (CENTURY) calibrated for our site that indicated that only the large wood component of hurricane-generated debris was of sufficiently low quality and of great enough mass to cause the observed effects on productivity. The model predicted that nutrient immobilization by decaying wood should suppress net primary productivity for 13 yr and total live biomass for almost 30 yr following the hurricane. Our findings emphasize the substantial influence that woody debris has upon nutrient cycling and productivity in forest ecosystems through its effects on the activity of decomposers. We suggest that the manner in which woody debris regulates ecosystem function in different forests is significantly affected by disturbance regime.

Although soil resources are widely considered as a major factor that reduces the productivity, stature, and diversity of tropical montane cloud forests (TMCF), systematic comparisons of soil resources within and between TMCF are lacking. This study combines published reports on TMCF soils with new data on the soils and forest structure of the Luquillo Mountains in Puerto Rico to assess the current state of knowledge regarding global and local-scale variation in TMCF soils. At the global scale, soils from 33 TMCF sites and over 150 pedons are reviewed. Compared to soils in humid lowland tropical forests, TMCF soils are relatively acidic, have higher organic matter content, and are relatively high in total nitrogen and extractable phosphorus. Across all sites, significant correlations also exist between mean annual precipitation and soil pH and base saturation, but not between any soil chemical factor and canopy height, site elevation, or air temperature. Although comparisons between TMCF are limited by inconsistent sampling protocols, analysis of available data does indicates that lower montane cloud forests (LMCF) have taller canopies, higher soil pH, lower soil nitrogen, and higher C/N ratios than upper montane cloud forests (UMCF). Within an UMCF in NE Puerto Rico, the abundance of soil nitrogen, carbon, and potassium accounted for 25% to 54% of the variation in canopy height. However, as much as 68% of the variation in stand height could be accounted for when site exposure, slope gradient, and the percent coverage of surface roots were also included in the analysis.

Little is known about ecosystem-level responses to multiple, climatic disturbance events. In the subtropical forests of Puerto Rico, the major natural disturbances are hurricanes and droughts. We tested the ecosystem-level effects of these disturbances in sites with different land use histories. From 1989 to 1992, data were collected to determine the effects of Hurricane Hugo and two droughts on litterfall inputs, fine-root biomass, and decomposition rates in three topographic locations (stream, riparian, upslope) within two watersheds. From 1994 to 1998, we added a third watershed and an experiment in which coarse-wood levels were manipulated to simulate hurricane inputs. Data were collected on tree and palm growth rates, litterfall inputs, fine-root biomass, and decomposition rates. From 1994 to 1998, four hurricanes and three droughts were recorded. Measured parameters had unique responses and recovery rates to hurricanes and droughts. Litterfall inputs returned to long-term mean rates within one month following droughts and small-to-moderate hurricanes but required five years to recover after an intense hurricane. In contrast, fine-root biomass recovered seven months after an intense hurricane but failed to recover after five years following a severe drought. Despite the dramatic effects of these weather events on some ecosystem parameters, we found that aboveground measures of tree and palm growth were more affected by preexisting site conditions (e.g., nitrogen availability due to past land use activities) than hurricanes or droughts. The addition of coarse woody debris increased tree and palm growth, fine-root biomass, and litter production; however, in the case of tree and palm growth, this effect was least measurable in the sites with the highest productivity. We found that decomposition rates were more controlled by litter quality than weather conditions. In conclusion, we found that certain ecosystem structures (e.g., canopy structure and fine-root biomass) generally recovered more slowly from disturbance events than certain ecosystem processes (e.g., plant growth rates, decomposition rates). We also found that past land use activities and disturbance legacies were important in determining the responses and recovery rates of the ecosystem to disturbance.

Determining the ubiquity of top-down control effects of predators on their prey and ecosystem processes is important for understanding community and ecosystem-level consequences that may result from predator loss. We conducted experiments at two spatial scales to investigate the effects of terrestrial frogs (Eleutherodactylus coqui) on aerial and litter invertebrates, plant growth and herbivory, and litter decomposition. At both scales, frogs reduced aerial invertebrates and leaf herbivory, but had no effect on litter invertebrates. At the smaller scale, frogs increased foliage production rates, measured as the number of new leaves and new leaf area produced, by 80% and decomposition rates by 20%. The influence of E. coqui on increasing primary productivity and decomposition rates at the smaller scale appeared to be a result of elimination and excretion rather than of controlling prey. While the results provide evidence for frogs controlling herbivorous prey at both scales, species effects on ecosystem processes were only detectable at the smaller scale. The results highlight the difficulties in conducting experiments at large spatial scales. The findings from this study imply that the loss of amphibians and other species of higher trophic levels may affect nutrient cycling rates in tropical forests.