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Uncertainties about controls on tree mortality make forest responses to land-use and climate change difficult to predict. We tracked biomass of tree functional groups in tropical forest inventories across Puerto Rico and the U.S. Virgin Islands, and with random forests we ranked 86 potential predictors of small tree survival (young or mature stems 2.5–12.6 cm diameter at breast height). Forests span dry to cloud forests, range in age, geology and past land use and experienced severe drought and storms. When excluding species as a predictor, top predictors are tree crown ratio and height, two to three species traits and stand to regional factors reflecting local disturbance and the system state (widespread recovery, drought, hurricanes). Native species, and species with denser wood, taller maximum height, or medium typical height survive longer, but short trees and species survive hurricanes better. Trees survive longer in older stands and with less disturbed canopies, harsher geoclimates (dry, edaphically dry, e.g., serpentine substrates, and highest-elevation cloud forest), or in intervals removed from hurricanes. Satellite image phenology and bands, even from past decades, are top predictors, being sensitive to vegetation type and disturbance. Covariation between stand-level species traits and geoclimate, disturbance and neighboring species types may explain why most neighbor variables, including introduced vs. native species, had low or no importance, despite univariate correlations with survival. As forests recovered from a hurricane in 1998 and earlier deforestation, small trees of introduced species, which on average have lighter wood, died at twice the rate of natives. After hurricanes in 2017, the total biomass of trees ≥12.7 cm dbh of the introduced species Spathodea campanulata spiked, suggesting that more frequent hurricanes might perpetuate this light-wooded species commonness. If hurricane recovery favors light-wooded species while drought favors others, climate change influences on forest composition and ecosystem services may depend on the frequency and severity of extreme climate events.

Global climate change has led to rising temperatures and to more frequent and intense climatic events, such as storms and droughts. Changes in climate and disturbance regimes can have non-additive effects on plant communities and result in complicated legacies we have yet to understand. This is especially true for tropical forests, which play a significant role in regulating global climate. We used understory vegetation data from the Tropical Responses to Altered Climate Experiment (TRACE) in Puerto Rico to evaluate how plant communities responded to climate warming and disturbance. The TRACE understory vegetation was exposed to a severe drought (2015), 2 years of experimental warming (4°C above ambient in half of the plots, 2016–2017 and 2018–2019), and two major hurricanes (Irma and María, September 2017). Woody seedlings and saplings were censused yearly from 2015 to 2019, with an additional census in 2015 after the drought ended. We evaluated disturbance-driven changes in species richness, diversity, and composition across ontogeny. We then used Bayesian predictive trait modeling to assess how species responded to disturbance and how this might influence the functional structure of the plant community. Our results show decreased seedling richness after hurricane disturbance, as well as increased sapling richness and diversity after warming. We found a shift in species composition through time for both seedlings and saplings, yet the individual effects of each disturbance were not significant. At both ontogenetic stages, we observed about twice as many species responding to experimental warming as those responding to drought and hurricanes. Predicted changes in functional structure point to disturbance-driven functional shifts toward a mixture of fast-growing and drought-tolerant species. Our findings demonstrate that the tropical forest understory community is more resistant to climatic stressors than expected, especially at the sapling stage. However, early signs of changes in species composition suggest that, in a warming climate with frequent droughts and hurricanes, plant communities might shift over time toward fast-growing or drought-tolerant species.

Abstract Tropical forests are the largest contributors to global emissions of carbon dioxide (CO2) to the atmosphere via soil respiration (Rs). As such, identifying the main controls on Rs in tropical forests is essential for accurately projecting the consequences of ongoing and future global environmental changes to the global C cycle. We measured hourly Rs in a secondary tropical moist forest in Puerto Rico over a 3‐year period to (a) quantify the magnitude of Rs and (b) identify the role of climatic, substrate, and nutrient controls on the seasonality of Rs. Across 3 years of measurements, mean Rs was 7.16 ± 0.02 μmol CO2 m‐2 s‐1 (or 2,710 g C m‐2 year‐1) and showed significant seasonal variation. Despite small month‐to‐month variation in temperature (~4°C), we found significant positive relationships between daily and monthly Rs with both air and soil temperature, highlighting the importance of temperature as a driver of Rs even in warm ecosystems, such as tropical forests. We also found a significant parabolic relationship between mean daily volumetric soil moisture and mean daily Rs, with an optimal moisture value of 0.34 m3 m‐3. Given the relatively consistent climate at this site, the large range in mean monthly Rs (~7 μmol CO2 m‐2 s‐1) was surprising and suggests that even small changes in climate can have large implications for ecosystem respiration. The strong positive relationship of Rs with temperature at monthly timescales particularly stands out, as moisture is usually considered a stronger control of Rs in tropical forests that already experience warm temperatures year‐round. Moreover, our results revealed the strong seasonality of Rs in tropical moist forests, which given its high magnitude, can represent a significant contribution to the seasonal patterns of atmospheric (CO2) globally.

Abstract The effects of climate change on tropical forests may have global consequences due to the forests’ high biodiversity and major role in the global carbon cycle. In this study, we document the effects of experimental warming on the abundance and composition of a tropical forest floor herbaceous plant community in the Luquillo Experimental Forest, Puerto Rico. This study was conducted within Tropical Responses to Altered Climate Experiment (TRACE) plots, which use infrared heaters under free‐air, open‐field conditions, to warm understory vegetation and soils + 4°C above nearby control plots. Hurricanes Irma and María damaged the heating infrastructure in the second year of warming, therefore, the study included one pretreatment year, one year of warming, and one year of hurricane response with no warming. We measured percent leaf cover of individual herbaceous species, fern population dynamics, and species richness and diversity within three warmed and three control plots. Results showed that one year of experimental warming did not significantly affect the cover of individual herbaceous species, fern population dynamics, species richness, or species diversity. In contrast, herbaceous cover increased from 20% to 70%, bare ground decreased from 70% to 6%, and species composition shifted pre to posthurricane. The negligible effects of warming may have been due to the short duration of the warming treatment or an understory that is somewhat resistant to higher temperatures. Our results suggest that climate extremes that are predicted to increase with climate change, such as hurricanes and droughts, may cause more abrupt changes in tropical forest understories than longer‐term sustained warming.

Despite the importance of tropical forests to global carbon balance, our understanding of how tropical plant physiology will respond to climate warming is limited. In addition, the contribution of tropical forest understories to global carbon cycling is predicted to increase with rising temperatures, however, in situ warming studies of tropical forest plants to date focus only on upper canopies. We present results of an in situ field-scale +4°C understory infrared warming experiment in Puerto Rico (Tropical Responses to Altered Climate Experiment; TRACE). We investigated gas exchange responses of two common understory shrubs, Psychotria brachiata and Piper glabrescens, after exposure to 4 and 8 months warming. We assessed physiological acclimation in two ways: (1) by comparing plot-level physiological responses in heated versus control treatments before and after warming, and (2) by examining physiological responses of individual plants to variation in environmental drivers across all plots, seasons, and treatments. P. brachiata has the capacity to up-regulate (i.e., acclimate) photosynthesis through broadened thermal niche and up-regulation of photosynthetic temperature optimum (Topt) with warmer temperatures. P. glabrescens, however, did not upregulate any photosynthetic parameter, but rather experienced declines in the rate of photosynthesis at the optimum temperature (Aopt), corresponding with lower stomatal conductance under warmer daily temperatures. Contrary to expectation, neither species showed strong evidence for respiratory acclimation. P. brachiata down-regulated basal respiration with warmer daily temperatures during the drier winter months only. P. glabrescens showed no evidence of respiratory acclimation. Unexpectedly, soil moisture, was the strongest environmental driver of daily physiological temperature responses, not vegetation temperature. Topt increased, while photosynthesis and basal respiration declined as soils dried, suggesting that drier conditions negatively affected carbon uptake for both species. Overall, P. brachiata, an early successional shrub, showed higher acclimation potential to daily temperature variations, potentially mitigating negative effects of chronic warming. The negative photosynthetic response to warming experienced by P. glabrescens, a mid-successional shrub, suggests that this species may not be able to as successfully tolerate future, warmer temperatures. These results highlight the importance of considering species when assessing climate change and relay the importance of soil moisture on plant function in large-scale warming experiments.

Abstract The response of tropical forests to global warming is one of the largest uncertainties in predicting the future carbon balance of Earth. To determine the likely effects of elevated temperatures on tropical forest understory plants and soils, as well as other ecosystems, an infrared (IR) heater system was developed to provide in situ warming for the Tropical Responses to Altered Climate Experiment (TRACE) in the Luquillo Experimental Forest in Puerto Rico. Three replicate heated 4‐m‐diameter plots were warmed to maintain a 4°C increase in understory vegetation compared to three unheated control plots, as sensed by IR thermometers. The equipment was larger than any used previously and was subjected to challenges different from those of many temperate ecosystem warming systems, including frequent power surges and outages, high humidity, heavy rains, hurricanes, saturated clayey soils, and steep slopes. The system was able to maintain the target 4.0°C increase in hourly average vegetation temperatures to within ± 0.1°C. The vegetation was heterogeneous and on a 21° slope, which decreased uniformity of the warming treatment on the plots; yet, the green leaves were fairly uniformly warmed, and there was little difference among 0–10 cm depth soil temperatures at the plot centers, edges, and midway between. Soil temperatures at the 40–50 cm depth increased about 3°C compared to the controls after a month of warming. As expected, the soil in the heated plots dried faster than that of the control plots, but the average soil moisture remained adequate for the plants. The TRACE heating system produced an adequately uniform warming precisely controlled down to at least 50‐cm soil depth, thereby creating a treatment that allows for assessing mechanistic responses of tropical plants and soil to warming, with applicability to other ecosystems. No physical obstacles to scaling the approach to taller vegetation (i.e., trees) and larger plots were observed.

Global climate models predict a future of increased severity of drought in many tropical forests. Soil microbes are central to the balance of these systems as sources or sinks of atmospheric carbon (C), yet how they respond metabolically to drought is not well understood. We simulated drought in the typically aseasonal Luquillo Experimental Forest, Puerto Rico, by intercepting precipitation falling through the forest canopy. This approach reduced soil moisture by 13 % and water potential by 0.14 MPa (from -0.2 to -0.34). Previous results from this experiment have demonstrated that the diversity and composition of these soil microbial communities are sensitive to even small changes in soil water. Here we show prolonged drought significantly alters the functional potential of the community and provokes a clear osmotic stress response, including the production of compatible solutes that increase intracellular C demand. Subsequently, a microbial population emerges with a greater capacity for extracellular enzyme production targeting macromolecular carbon. Significantly, some of these drought-induced functional shifts in the soil microbiota are attenuated by prior exposure to a short-term drought suggesting that acclimation may occur despite a lack of longer-term drought history.

Climate model projections for tropical regions show clear perturbation of precipitation patterns leading to increased frequency and severity of drought in some regions. Previous work has shown declining soil moisture to be a strong driver of changes in microbial trait distribution, however the feedback of any shift in functional potential on ecosystem properties related to carbon cycling are poorly understood. Here we show that drought-induced changes in microbial functional diversity and activity shape, and are in turn shaped by, the composition of dissolved and soil-associated carbon. We also demonstrate that a shift in microbial functional traits that favor the production of hygroscopic compounds alter the efflux of carbon dioxide following soil rewetting. Under drought the composition of the dissolved organic carbon pool changed in a manner consistent with a microbial metabolic response. We hypothesize that this microbial ecophysiological response to changing soil moisture elevates the intracellular carbon demand stimulating extracellular enzyme production, that prompts the observed decline in more complex carbon compounds (e.g., cellulose and lignin). Furthermore, a metabolic response to drought appeared to condition (biologically and physically) the soil, notably through the production of polysaccharides, particularly in experimental plots that had been pre-exposed to a short-term drought. This hysteretic response, in addition to an observed drought-related decline in phosphorus concentration, may have been responsible for a comparatively modest CO2 efflux following wet-up in drought plots relative to control plots.

Young secondary forests and plantations in the moist tropics often have rapid rates of biomass accumulation and thus sequester large amounts of carbon. Here, we compare results from mature forest and nearby 15-20 year old tree plantations in lowland Costa Rica to evaluate differences in allocation of carbon to aboveground production and root systems. We found that the tree plantations, which had fully developed, closed canopies, allocated more carbon belowground - to their root systems - than did mature forest. This increase in belowground carbon allocation correlated significantly with aboveground tree growth but not with canopy production (i.e., leaf fall or fine litter production). In contrast, there were no correlations between canopy production and either tree growth or belowground carbon allocation. Enhanced allocation of carbon to root systems can enhance plant nutrient uptake, providing nutrients beyond those required for the production of short-lived tissues such as leaves and fine roots, and thus enabling biomass accumulation. Our analyses support this deduction at our site, showing that enhanced allocation of carbon to root systems can be an important mechanism promoting biomass accumulation during forest growth in the moist tropics. Identifying factors that control when, where and for how long this occurs would help us to improve models of forest growth and nutrient cycling, and to ascertain the role that young forests play in mitigating increased atmospheric carbon dioxide.

Precipitation and temperature are important drivers of soil respiration. The role of moisture and temperature are generally explored at seasonal or inter-annual timescales; however, significant variability also occurs on hourly to daily time-scales. We used small (1.54 m(2)), throughfall exclusion shelters to evaluate the role soil moisture and temperature as temporal controls on soil CO2 efflux from a humid tropical forest in Puerto Rico. We measured hourly soil CO2 efflux, temperature and moisture in control and exclusion plots (n = 6) for 6-months. The variance of each time series was analyzed using orthonormal wavelet transformation and Haar-wavelet coherence. We found strong negative coherence between soil moisture and soil respiration in control plots corresponding to a two-day periodicity. Across all plots, there was a significant parabolic relationship between soil moisture and soil CO2 efflux with peak soil respiration occurring at volumetric soil moisture of approximately 0.375 m(3)/m(3). We additionally found a weak positive coherence between CO2 and temperature at longer time-scales and a significant positive relationship between soil temperature and CO2 efflux when the analysis was limited to the control plots. The coherence between CO2 and both temperature and soil moisture were reduced in exclusion plots. The reduced CO2 response to temperature in exclusion plots suggests that the positive effect of temperature on CO2 is constrained by soil moisture availability.