We excluded throughfall from humid tropical forests in Puerto Rico for a period of three months to determine how drought affects greenhouse gas emissions from tropical forest soils. We established five 1.24 m2 throughfall exclusion and five control plots of equal size in three sites located on ridges, slopes, and an upland valley dominated by palms (total of 30 plots). We measured weekly changes in carbon dioxide (CO2) and bi-weekly changes in nitrous oxide (N2O) and methane (CH4) in response to manipulation. We additionally measured the effects of throughfall exclusion on soil temperature and moisture, nutrient availability, and pH. Rainout shelters significantly reduced throughfall by 22 to 32 % and decreased soil moisture by 16 to 36% (top 10 cm). Rates of CO2 emissions decreased significantly in the ridge and slope sites (30%, 28%, respectively), but not the palm during the experimental drought. In contrast, the palm site became a significantly stronger sink for CH4 in response to drying (480% decline relative to controls), while CH4 fluxes in the ridge and slope sites did not respond to drought. Both the palm and ridge site became a sink for N2O in response to drought and the slope site followed a similar trend. Soil pH and available P decreased significantly in response to soil drying; however, available N was not affected. Variability in the response of greenhouse gas emissions to drought among the three sites highlights the complexity of biogeochemical cycling in tropical forested ecosystems, as well as the need for research that incorporates the high degree of spatial heterogeneity in experimental designs. Our results show that humid tropical forests are sensitive to climate change and that short-term declines in rainfall could result in a negative feedback to climate change via lowered greenhouse gas emissions and increased greenhouse gas consumption by soils.

The effects of historical land use on tropical forest must be examined to understand present forest characteristics and to plan conservation strategies. We compared the effects of past land use, topography, soil type, and other environmental variables on tree species composition in a subtropical wet forest in the Luquillo Mountains, Puerto Rico. The study involved stems > 10 cm diameter measured at 130 cm above the ground, within the 16-ha Luquillo Forest Dynamics Plot (LFDP), and represents the forest at the time Hurricane Hugo struck in 1989. Topography in the plot is rugged, and soils are variable. Historical documents and local residents described past land uses such as clear-felling and selective logging followed by farming, fruit and coffee production, and timber stand improvement in the forest area that now includes the LFDP. These uses ceased 40-60 yr before the study, but their impacts could be differentiated by percent canopy cover seen in aerial photographs from 1936. Using these photographs, we defined four historic cover classes within the LFDP. These ranged from cover class 1, the least tree-covered area in 1936, to cover class 4, with the least intensive historic land use (selective logging and timber stand improvement). In 1989, cover class 1 had the lowest stem density and proportion of large stems, whereas cover class 4 had the highest basal area, species richness, and number of rare and endemic species. Ordination of tree species composition (89 species, 13 167 stems) produced arrays that primarily corresponded to the four cover classes (i.e., historic land uses). The ordination arrays corresponded secondarily to soil characteristics and topography. Natural disturbances (hurricanes, landslides, and local treefalls) affected tree composition, but these effects did not correlate with the major patterns of species distributions on the plot. Thus, it appears that forest development and natural disturbance have not masked the effects of historical land use in this tropical forest, and that past land use was the major influence on the patterns of tree composition in the plot in 1989. The least disturbed stand harbors more rare and endemic species, and such stands should be protected.

leaffall periodicity has been related to rainfall regime and dry season length. In weakly seasonal forests, where there is no marked dry season, other climatic factors could trigger leaf shed. In this study, we aimed to determine if other climatic variables (wind speed, solar radiation, photosynthetic photon flux density [PPFD], day length, temperature, and relative humidity) could be better correlated with patterns of litter and leaffall in a weakly seasonal subtropical wet forest in Puerto Rico. Leaffall patterns were correlated mainly with solar radiation, PPFD, day length, and temperature; and secondarily with rainfall. Two main peaks of leaffall were observed: April–June and August–September, coinciding with the periods of major solar radiation at this latitude. Community leaffall patterns were the result of overlapping peaks of individual species. Of the 32 species analyzed, 21 showed phenological patterns, either unimodal (16 species), bimodal (three species), or multimodal (two species). Lianas also presented leaffall seasonality, suggesting that they are subject to the same constraints and triggering factors affecting trees. In addition to solar radiation as a main determinant of leaffall timing in tropical forests, our findings highlight the importance of interannual variation and asynchrony, suggesting that leaffall is the result of a complex interaction between environmental and physiological factors.

1. Excretion of nitrogen (N) and phosphorus (P) is a direct and potentially important role for aquatic consumers in nutrient cycling that has recently garnered increased attention. The ecosystem-level significance of excreted nutrients depends on a suite of abiotic and biotic factors, however, and few studies have coupled measurements of excretion with consideration of its likely importance for whole-system nutrient fluxes. 2. We measured rates and ratios of N and P excretion by shrimps (Xiphocaris elongata and Atya spp.) in two tropical streams that differed strongly in shrimp biomass because a waterfall excluded predatory fish from one site. We also made measurements of shrimp and basal resource carbon (C), N and P content and estimated shrimp densities and ecosystem-level N and P excretion and uptake. Finally, we used a 3-year record of discharge and NH4-N concentration in the high-biomass stream to estimate temporal variation in the distance required for excretion to turn over the ambient NH4-N pool. 3. Per cent C, N, and P body content of Xiphocaris was significantly higher than that of Atya. Only per cent P body content showed significant negative relationships with body mass. C:N of Atya increased significantly with body mass and was higher than that of Xiphocaris. N : P of Xiphocaris was significantly higher than that of Atya. 4. Excretion rates ranged from 0.16–3.80 lmol NH4-N shrimp)1 h)1, 0.23–5.76 lmol total dissolved nitrogen (TDN) shrimp)1 h)1 and 0.002–0.186 lmol total dissolved phosphorus (TDP) shrimp)1 h)1. Body size was generally a strong predictor of excretion rates in both taxa, differing between Xiphocaris and Atya for TDP but not NH4-N and TDN. Excretion rates showed statistically significant but weak relationships with body content stoichiometry. 5. Large between-stream differences in shrimp biomass drove differences in total excretion by the two shrimp communities (22.3 versus 0.20 lmol NH4-N m)2 h)1, 37.5 versus 0.26 lmol TDN m)2 h)1 and 1.1 versus 0.015 lmol TDP m)2 h)1), equivalent to 21% and 0.5% of NH4-N uptake and 5% and <0.1% of P uptake measured in the high- and lowbiomass stream, respectively. Distances required for excretion to turn over the ambient NH4-N pool varied more than a hundredfold over the 3-year record in the high-shrimp stream, driven by variability in discharge and NH4-N concentration. 6. Our results underscore the importance of both biotic and abiotic factors in controlling consumer excretion and its significance for nutrient cycling in aquatic ecosystems. Differences in community-level excretion rates were related to spatial patterns in shrimp biomass dictated by geomorphology and the presence of predators. Abiotic factors also had important effects through temporal patterns in discharge and nutrient concentrations. Future excretion studies that focus on nutrient cycling should consider both biotic and abiotic factors in assessing the significance of consumer excretion in aquatic ecosystems.