Aboveground carbon responses to experimental and natural hurricane impacts in a subtropical wet forest in Puerto Rico

Climate change and disturbance make it difficult to project long-term patterns of carbon sequestration in tropical forests, but large ecosystem experiments in these forests can inform predictions. The Canopy Trimming Experiment (CTE) manipulates two key components of hurricane disturbance, canopy openness and detritus deposition, in a tropical forest in Puerto Rico. We documented how the CTE and a real hurricane affected tree recruitment, biomass, and aboveground carbon storage over 15 years. In the CTE treatments, we trimmed branches, but we did not fell trees. We expected that during the 14-year period after initial canopy trimming, regrowth of branches and stems and stem recruitment stimulated by increased light and trimmed debris would help restore biomass and carbon loss due to trimming. Compared to control plots, in the trimmed plots recruitment of palms and dicot trees increased markedly after trimming, and stem diameters of standing trees increased. Data showed that recruitment of small trees adds little to aboveground carbon, compared to the amount in large trees. Nevertheless, this response restored pretreatment biomass and carbon in the experimental period. In particular, the experimental additions of trimmed debris on the forest floor seemed to stimulate increase in aboveground carbon. Toward the end of the experimental period, Hurricane Maria (Category 4 hurricane) trimmed and felled some trees but reduced aboveground carbon less in the plots (including untrimmed plots) than experimental trimming had. Thus, it appears that the amount of regrowth recorded after experimental trimming could also restore aboveground carbon in the forest after a severe hurricane in the same time span. However, Hurricane Maria, unlike the trimming treatments, felled large trees, and it may be that with predicted, more frequent severe hurricanes, that the continued loss of large trees would over the long term decrease aboveground carbon stored in this Puerto Rican forest and likewise in other tropical forests affected by cyclonic storms

Solar radiation and soil moisture drive tropical forest understory responses to experimental and natural hurricanes

Tropical forest understory regeneration occurs rapidly after disturbance with compositional trajectories that depend on species availability and environmental conditions. To predict future tropical forest regeneration dynamics, we need a deeper understanding of how pulse disturbance events, like hurricanes, interact with environmental variability to affect understory demography and composition. We examined fern and sapling mortality, recruitment, and community composition in relation to solar radiation and soil moisture using 17 years of forest dynamics data (2003–2019) from the Canopy Trimming Experiment in the Luquillo Experimental Forest, Puerto Rico. Solar radiation increased 150% and soil moisture increased 40% following canopy trimming of experimental plots relative to control plots. All plots were disturbed in 2017 by Hurricanes Irma and Maria, so experimentally trimmed plots presented the opportunity to study the effects of multiple hurricanes, while control plots isolated the effects of a single natural hurricane. Recruitment rates maximized at 0.14 individuals/plot/month for ferns and 0.20 stems/plot/month for saplings. Recruitment and mortality were distributed more evenly over the 17 years of monitoring in experimentally trimmed plots than in control plots; however, following Hurricane Maria demographic rates substantially increased in control plots only. In experimentally trimmed plots, the largest community compositional shifts occurred as a result of the trimming events, and compositional changes were greatest for control plots after Hurricane Maria in 2017. Pioneer tree and fern species increased in abundance in response to both simulated and natural hurricanes. Following Hurricane Maria, two dominant pioneer species, Cyathea arborea and Cecropia schreberiana, recruited abundantly, but only in control plots. In trimmed plots, increased solar radiation and soil moisture shifted understory species composition steadily toward pioneer and secondary-successional species, with soil moisture interacting strongly with canopy trimming. Thus, both solar radiation and soil moisture are environmental drivers affecting pioneer species recruitment following disturbance, which interact with canopy opening following hurricanes. Our results suggest that if hurricane disturbances increase in frequency and severity, as suggested by climate change predictions, the understory regeneration of late-successional species, such as Manilkara bidentata and Sloanea berteroana, which prefer deeper shade and slightly drier soil microsites, may become imperiled

Seven-year responses of trees to experimental hurricane effects in a tropical rainforest, Puerto Rico

We experimentally manipulated key components of severe hurricane disturbance, canopy openness and detritus deposition, to determine the independent and interactive effects of these components on tree recruitment, forest structure, and diversity in a wet tropical forest in the Luquillo Experimental Forest, Puerto Rico. Canopy openness was increased by trimming branches, and we manipulated debris by adding or subtracting the trimmed materials to the trimming treatments, in a 2 X 2 factorial design replicated in three blocks. Tree (stems \u3e 1 cm diameter at breast height) responses were measured during the 9-year study, which included at least 1 year of pre-manipulation monitoring. When the canopy was trimmed, stem densities increased \u3e2-fold and rates of recruitment increased \u3e25-fold. Deposition of canopy debris did not markedly affect stem densities but did have small yet significant effects on tree basal area. Basal area increased about 10% when debris was added to plots with intact canopies; the other treatments exhibited smaller or no increases in basal area over time. Much of the dynamics of stem densities were due to changes in the smallest size class (1–2.5 cm diameter), which responded with a pulse of recruitment in the canopy trimmed treatments, and a steady loss in plots with intact canopies. The decreases in stem densities in the plots with intact canopies is attributed to observed on-going forest thinning from the last natural severe hurricane disturbance in 1998. Given these repeated hurricane effects, our study enabled an experimental test of the Intermediate Disturbance Hypothesis (IDH), for which we predicted an increase in species diversity in canopy trimmed treatments and a loss of species in the treatments with intact canopies. Measured patterns of diversity gave partial support to the predictions of IDH, although raw species richness of sampled plots fit the predictions better than richness adjusted for differences in stem densities among treatments. Ordination of species responses in the community identified a guild of pioneer species responding to the trimmed treatments, but not the debris additions, amongst substantial background variation in species composition unrelated to the experimental treatments. These results are consistent with a growing consensus that, while trade-offs of resilience and resistance govern many species responses to hurricane disturbance, other environmental and historical factors are equally or more important in governing community dynamics in hurricane-disturbed forests

Solar radiation and soil moisture drive tropical forest understory responses to experimental and natural hurricanes

Abstract Tropical forest understory regeneration occurs rapidly after disturbance with compositional trajectories that depend on species availability and environmental conditions. To predict future tropical forest regeneration dynamics, we need a deeper understanding of how pulse disturbance events, like hurricanes, interact with environmental variability to affect understory demography and composition. We examined fern and sapling mortality, recruitment, and community composition in relation to solar radiation and soil moisture using 17 years of forest dynamics data (2003–2019) from the Canopy Trimming Experiment in the Luquillo Experimental Forest, Puerto Rico. Solar radiation increased 150% and soil moisture increased 40% following canopy trimming of experimental plots relative to control plots. All plots were disturbed in 2017 by Hurricanes Irma and Maria, so experimentally trimmed plots presented the opportunity to study the effects of multiple hurricanes, while control plots isolated the effects of a single natural hurricane. Recruitment rates maximized at 0.14 individuals/plot/month for ferns and 0.20 stems/plot/month for saplings. Recruitment and mortality were distributed more evenly over the 17 years of monitoring in experimentally trimmed plots than in control plots; however, following Hurricane Maria demographic rates substantially increased in control plots only. In experimentally trimmed plots, the largest community compositional shifts occurred as a result of the trimming events, and compositional changes were greatest for control plots after Hurricane Maria in 2017. Pioneer tree and fern species increased in abundance in response to both simulated and natural hurricanes. Following Hurricane Maria, two dominant pioneer species, Cyathea arborea and Cecropia schreberiana, recruited abundantly, but only in control plots. In trimmed plots, increased solar radiation and soil moisture shifted understory species composition steadily toward pioneer and secondary‐successional species, with soil moisture interacting strongly with canopy trimming. Thus, both solar radiation and soil moisture are environmental drivers affecting pioneer species recruitment following disturbance, which interact with canopy opening following hurricanes. Our results suggest that if hurricane disturbances increase in frequency and severity, as suggested by climate change predictions, the understory regeneration of late‐successional species, such as Manilkara bidentata and Sloanea berteroana, which prefer deeper shade and slightly drier soil microsites, may become imperiled