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Tropical forest canopies cycle vast amounts of carbon, yet we still have a limited understanding of how these critical ecosystems will respond to climate warming. We implemented in situ leaf-level + 3°C experimental warming from the understory to the upper canopy of two Puerto Rican tropical tree species, Guarea guidonia and Ocotea sintenisii. After approximately 1 month of continuous warming, we assessed adjustments in photosynthesis, chlorophyll fluorescence, stomatal conductance, leaf traits and foliar respiration. Warming did not alter net photosynthetic temperature response for either species; however, the optimum temperature of Ocotea understory leaf photosynthetic electron transport shifted upward. There was no Ocotea respiratory treatment effect, while Guarea respiratory temperature sensitivity (Q10 ) was down-regulated in heated leaves. The optimum temperatures for photosynthesis (Topt ) decreased 3-5°C from understory to the highest canopy position, perhaps due to upper canopy stomatal conductance limitations. Guarea upper canopy Topt was similar to the mean daytime temperatures, while Ocotea canopy leaves often operated above Topt . With minimal acclimation to warmer temperatures in the upper canopy, further warming could put these forests at risk of reduced CO2 uptake, which could weaken the overall carbon sink strength of this tropical forest.

Tropical forests are the largest contributors to global emissions of carbon dioxide (CO2) to the atmosphere via soil respiration (R s). As such, identifying the main controls on R s 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 R s in a secondary tropical moist forest in Puerto Rico over a 3‐year period to (a) quantify the magnitude of R s and (b) identify the role of climatic, substrate, and nutrient controls on the seasonality of R s. Across 3 years of measurements, mean R s 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 R s with both air and soil temperature, highlighting the importance of temperature as a driver of R s even in warm ecosystems, such as tropical forests. We also found a significant parabolic relationship between mean daily volumetric soil moisture and mean daily R s, 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 R s (~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 R s with temperature at monthly timescales particularly stands out, as moisture is usually considered a stronger control of R s in tropical forests that already experience warm temperatures year‐round. Moreover, our results revealed the strong seasonality of R s in tropical moist forests, which given its high magnitude, can represent a significant contribution to the seasonal patterns of atmospheric (CO2) globally., We measured hourly soil respiration in a tropical moist forest in Puerto Rico over a 3‐year period to (a) quantify the magnitude of R s and (b) identify the role of climatic, substrate, and nutrient controls on the seasonality of R s. Across 3 years of measurements, mean R s was 7.08 ± 0.02 μmol CO2 m‐2 s‐1 (2,728 g C m‐2 year‐1) and showed significant seasonal variation at daily and monthly timescales. Despite small month‐to‐month variation in temperature (~4 C), seasonal patterns of daily and monthly R s were strongly correlated with air temperature (R 2 ranged from 0.49 to 0.70 depending on the timescale), highlighting the importance of temperature as a driver of R s even in warm ecosystems, such as tropical forests.

Habitability has been generally defined as the capability of an environment to support life. Ecologists have been using Habitat Suitability Models (HSMs) for more than four decades to study the habitability of Earth from local to global scales. Astrobiologists have been proposing different habitability models for some time, with little integration and consistency among them, being different in function to those used by ecologists. Habitability models are not only used to determine if environments are habitable or not, but they also are used to characterize what key factors are responsible for the gradual transition from low to high habitability states. Here we review and compare some of the different models used by ecologists and astrobiologists and suggest how they could be integrated into new habitability standards. Such standards will help to improve the comparison and characterization of potentially habitable environments, prioritize target selections, and study correlations between habitability and biosignatures. Habitability models are the foundation of planetary habitability science and the synergy between ecologists and astrobiologists is necessary to expand our understanding of the habitability of Earth, the Solar System, and extrasolar planets., Published in Astrobiology, 21(8). arXiv admin note: substantial text overlap with arXiv:2007.05491

Tropical forests are expected to experience unprecedented warming and increases in hurricane disturbances in the coming decades; yet, our understanding of how these productive systems, especially their belowground component, will respond to the combined effects of varied environmental changes remains empirically limited. Here we evaluated the responses of root dynamics (production, mortality, and biomass) to soil and understory warming (+4°C) and after two consecutive tropical hurricanes in our in situ warming experiment in a tropical forest of Puerto Rico: Tropical Responses to Altered Climate Experiment (TRACE). We collected minirhizotron images from three warmed plots and three control plots of 12 m2 . Following Hurricanes Irma and Maria in September 2017, the infrared heater warming treatment was suspended for repairs, which allowed us to explore potential legacy effects of prior warming on forest recovery. We found that warming significantly reduced root production and root biomass over time. Following hurricane disturbance, both root biomass and production increased substantially across all plots; the root biomass increased 2.8-fold in controls but only 1.6-fold in previously warmed plots. This pattern held true for both herbaceous and woody roots, suggesting that the consistent antecedent warming conditions reduced root capacity to recover following hurricane disturbance. Root production and mortality were both related to soil ammonium nitrogen and microbial biomass nitrogen before and after the hurricanes. This experiment has provided an unprecedented look at the complex interactive effects of disturbance and climate change on the root component of a tropical forested ecosystem. A decrease in root production in a warmer world and slower root recovery after a major hurricane disturbance, as observed here, are likely to have longer-term consequences for tropical forest responses to future global change.

Globally, soils store two to three times as much carbon as currently resides in the atmosphere, and it is critical to understand how soil greenhouse gas (GHG) emissions and uptake will respond to ongoing climate change. In particular, the soil‐to‐atmosphere CO2 flux, commonly though imprecisely termed soil respiration (R S), is one of the largest carbon fluxes in the Earth system. An increasing number of high‐frequency R S measurements (typically, from an automated system with hourly sampling) have been made over the last two decades; an increasing number of methane measurements are being made with such systems as well. Such high frequency data are an invaluable resource for understanding GHG fluxes, but lack a central database or repository. Here we describe the lightweight, open‐source COSORE (COntinuous SOil REspiration) database and software, that focuses on automated, continuous and long‐term GHG flux datasets, and is intended to serve as a community resource for earth sciences, climate change syntheses and model evaluation. Contributed datasets are mapped to a single, consistent standard, with metadata on contributors, geographic location, measurement conditions and ancillary data. The design emphasizes the importance of reproducibility, scientific transparency and open access to data. While being oriented towards continuously measured R S, the database design accommodates other soil‐atmosphere measurements (e.g. ecosystem respiration, chamber‐measured net ecosystem exchange, methane fluxes) as well as experimental treatments (heterotrophic only, etc.). We give brief examples of the types of analyses possible using this new community resource and describe its accompanying R software package., Here we describe the lightweight, open source COSORE (COntinuous SOil REspiration) database and software. COSORE focuses on automated, continuous and long‐term greenhouse gas flux datasets, and is intended to serve as a community resource for earth sciences, climate change syntheses and model evaluation.

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 four and eight 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 an 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 wetter 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.

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.

Methane (CH4) is a potent greenhouse gas that is both produced and consumed in soils by microbially mediated processes sensitive to soil redox. We evaluated the classical conceptual model of peatland CH4 dynamics—in which the water table position determines the vertical distribution of methanogenesis and methanotrophy—versus an emerging model in which methanogenesis and methanotrophy can both occur throughout the soil profile due to spatially heterogeneous redox and anaerobic CH4 oxidation. We simultaneously measured gross CH4 production and oxidation in situ across a microtopographical gradient in a drained temperate peatland and ex situ along the soil profile, giving us novel insight into the component fluxes of landscape-level net CH4 fluxes. Net CH4 fluxes varied among landforms (p

Tropical forests represent 50% of the planet's species and play a disproportionately large role in determining climate due to the vast amounts of carbon they store and exchange with the atmosphere. Currently, disturbance patterns in tropical ecosystems are changing due to factors such as increased land use pressure and altered patterns in hurricanes. At the same time, these regions are expected to experience unprecedented warming before 2100. Despite the importance of these ecosystems for forecasting the global consequences of multiple stressors, our understanding of how projected changes in climate and disturbance will affect the biogeochemical cycling of tropical forests remains in its infancy. Until now, no studies to our knowledge have evaluated forest recovery following hurricane disturbance within the context of concurrent climatic change. Here, we present soil biogeochemical results from a tropical forest field warming experiment in Puerto Rico where, a year after experimental warming began, Hurricanes Irma and Maria greatly altered the forest, allowing a unique opportunity to explore the interacting effects of hurricane disturbance and warming. We tracked post-hurricane forest recovery for a year without warming to assess legacy effects of prior warming on the disturbance response, and then reinitiated warming treatments to further evaluate interactions between forest recovery and warmer temperatures. The data showed that warming affected multiple aspects of soil biogeochemical cycling even in the first year of treatment, with particularly large positive effects on soil microbial biomass pools (e.g., increases of 54%, 33%, and 38% relative to the control plots were observed for microbial biomass carbon, nitrogen, and phosphorus, respectively after 6 months of warming). We also observed significant effects of the hurricanes on soil biogeochemical cycling, as well as interactive controls of warming and disturbance. Taken together, our results showed dynamic soil responses that suggest the future of biogeochemical cycling in this tropical wet forest will be strongly shaped by the directional effects of warming and the episodic effects of hurricanes.

Climate change is predicted to result in warmer and drier Neotropical forests relative to current conditions. Negative density-dependent feedbacks, mediated by natural enemies, are key to maintaining the high diversity of tree species found in the tropics, yet we have little understanding of how projected changes in climate are likely to affect these critical controls. Over 3 years, we evaluated the effects of a natural drought and in situ experimental warming on density-dependent feedbacks on seedling demography in a wet tropical forest in Puerto Rico. In the +4°C warming treatment, we found that seedling survival increased with increasing density of the same species (conspecific). These positive density-dependent feedbacks were not associated with a decrease in aboveground natural enemy pressure. If positive density-dependent feedbacks are not transient, the diversity of tropical wet forests, which may rely on negative density dependence to drive diversity, could decline in a future warmer, drier world.

International audience; Transpiration in humid tropical forests modulates the global water cycle and is a key driver of climate regulation. Yet, our understanding of how tropical trees regulate sap flux in response to climate variability remains elusive. With a progressively warming climate, atmospheric evaporative demand [i.e., vapor pressure deficit (VPD)] will be increasingly important for plant functioning, becoming the major control of plant water use in the twenty-first century. Using measurements in 34 tree species at seven sites across a precipitation gradient in the neotropics, we determined how the maximum sap flux velocity (vmax) and the VPD threshold at which vmax is reached (VPDmax) vary with precipitation regime [mean annual precipitation (MAP); seasonal drought intensity (PDRY)] and two functional traits related to foliar and wood economics spectra [leaf mass per area (LMA); wood specific gravity (WSG)]. We show that, even though vmax is highly variable within sites, it follows a negative trend in response to increasing MAP and PDRY across sites. LMA and WSG exerted little effect on vmax and VPDmax, suggesting that these widely used functional traits provide limited explanatory power of dynamic plant responses to environmental variation within hyper-diverse forests. This study demonstrates that long-term precipitation plays an important role in the sap flux response of humid tropical forests to VPD. Our findings suggest that under higher evaporative demand, trees growing in wetter environments in humid tropical regions may be subjected to reduced water exchange with the atmosphere relative to trees growing in drier climates.

Tropical forests have evolved under relatively narrow temperature regimes, and therefore may be more susceptible to climatic change than forests in higher latitudes. Recent evidence shows that lowland tropical forest canopies may already be exceeding thermal maxima for photosynthesis. Height can strongly influence both the microclimate and physiology of forest canopy foliage, yet vertical trends in canopy micrometeorology are rarely examined in tropical forests. To improve our understanding of how climatological and micrometeorological conditions affect tropical tree function, we assessed vertical gradients of photosynthetic photon flux density, vapor pressure deficit, air temperature, leaf temperature, and the difference between leaf and air temperature (ΔT) in a Puerto Rican tropical wet forest. Both air temperature and vapor pressure deficit increased linearly with height. Leaf temperature, however, did not significantly differ across the shaded foliage from 0-16 m, while the uppermost layer (20 m) was up to 4°C hotter than the rest of the foliage and up to 5°C hotter than air temperature at the highest radiation intensity. As a result, leaf temperatures in the shaded middle canopy and understory showed nearly poikilothermic behavior (i.e., leaf temperatures = air temperature), while the uppermost canopy strata showed megathermic behavior (i.e., leaf temperatures greater than air temperature), revealing different thermoregulation strategies for sun-lit versus shaded foliage. In addition, the uppermost canopy was the only stratum to exceed mean photosynthetic temperature optima for this site (Topt = 30.2 ± 1.1°C). Because the upper canopy plays a disproportionately large role in whole-forest photosynthesis, continued warming could potentially weaken the tropics’ carbon sink capacity. However, the shaded leaves may be able increase carbon uptake with further warming because they appear to be able to maintain temperatures below photosynthetic optima, possibly with the help of radiation shielding provided by the uppermost canopy layer.

The Luquillo Experimental Forest (LEF) has a long history of research on tropical forestry, ecology, and conservation, dating as far back as the early 19th Century. Scientific surveys conducted by early explorers of Puerto Rico, followed by United States institutions contributed early understanding of biogeography, species endemism, and tropical soil characteristics. Research in the second half of the 1900s established the LEF as an exemplar of forest management and restoration research in the tropics. Research conducted as part of a radiation experiment funded by the Atomic Energy Commission in the 1960s on forest metabolism established the field of ecosystem ecology in the tropics. Subsequent research has built on these early advances to develop new theories on ecosystem response to disturbance regimes and the role of the biota in ecosystem resilience. Recent and current research in the LEF has advanced understanding of resilience to hurricane disturbances, human land use, gamma irradiation, landslides, drought, and warming, showing that even following the most severe disturbances (e.g., landslides, agriculture) forests reestablish within 60 years. Work in the LEF has reversed the paradigm that tropical ecosystems are fragile, but instead exhibit remarkable resilience to many forms of disturbance present at multiple spatial and temporal scales. Current research is already advancing understanding of how climate change and attendant effects on the disturbance regime might affect the composition, structure, and function of tropical forest ecosystems.

The tropics are a region encircling the equator, delineated to the north by the Tropic of Cancer (23°26′14.0″N) and to the south by the Tropic of Capricorn (23°26′14.0″S). While we often think of the tropics as consistently warm and wet throughout the year, in reality, the tropics maintain a myriad of climates. Of the 116 Holdridge life zones (a global bioclimatic classification scheme), the tropics contain more life zones than the sum of all the planet's other geographic regions combined ( Holdridge, 1967 ). In addition to high climatic diversity, the tropics support a wide range of parent materials, landforms, geomorphic characteristics, and soil ages, and maintain all 12 soil types of the USDA soil taxonomy system ( Palm et al., 2007 ; Porder et al., 2007 ; Quesada et al., 2010 ; Richter and Babbar, 1991 ; Sanchez, 1977 ; Soil Survey Staff, 2006 ; Townsend et al., 2008 ). Accordingly, there is no single representative tropical ecosystem. Given the diversity of tropical biomes, this chapter will focus specifically on tropical forested ecosystems and their responses to warming because of their global importance, potential sensitivity to change, and the fact that an improved understanding of how these ecosystems may respond to warmer climate conditions is of significant importance to ecology and society. Furthermore, while generally considering all tropical forest types, emphasis in this chapter is on the humid tropics for which we have most data.

For more accurate projections of both the global carbon (C) cycle and the changing climate, a critical current need is to improve the representation of tropical forests in Earth system models. Tropical forests exchange more C, energy, and water with the atmosphere than any other class of land ecosystems. Further, tropical-forest C cycling is likely responding to the rapid global warming, intensifying water stress, and increasing atmospheric CO2 levels. Projections of the future C balance of the tropics vary widely among global models. A current effort of the modeling community, the ILAMB (International Land Model Benchmarking) project, is to compile robust observations that can be used to improve the accuracy and realism of the land models for all major biomes. Our goal with this paper is to identify field observations of tropical-forest ecosystem C stocks and fluxes, and of their long-term trends and climatic and CO2 sensitivities, that can serve this effort. We propose criteria for reference-level field data from this biome and present a set of documented examples from old-growth lowland tropical forests. We offer these as a starting point towards the goal of a regularly updated consensus set of benchmark field observations of C cycling in tropical forests.

Primary productivity in tropical forests is often considered limited by phosphorus (P) availability. Microbial activity is a key regulator of available P through organic matter decomposition (supply) as well as microbial immobilization (depletion). Environmental conditions, such as soil moisture and temperature can fluctuate significantly on hourly to daily time-scales in tropical forested ecosystems. Given the ability of microbes to respond rapidly to changing environmental conditions we would expect concomitant changes in the available soil P pool. Despite the potential for soil P availability to vary on short time-scales, research that investigates hourly to daily changes in the available soil P pool in tropical forests is extremely rare. We quantified diurnal fluctuations in labile soil P and the importance of biotic and abiotic factors in driving these patterns in a wet tropical forest in Puerto Rico. Hourly measurements of Bray-extractable P were made from sunrise to sunset on five separate days along with measurements of soil temperature, moisture, pH, soil respiration, and solar radiation. While we found no significant diurnal variation in labile P, it did, however, vary significantly across the five sample days (2.8–3.8 µg/g). The day-to-day variation in labile P was positively related to soil moisture (R2 = 0.42, p = 0.009). These findings illustrate the potential for rapid change in the available P pool in response to variable soil moisture status as well as the importance of considering soil moisture conditions when estimating P availability in the humid tropics.

Tropical tree species have evolved under very narrow temperature ranges compared to temperate forest species. Studies suggest that tropical trees may be more vulnerable to continued warming compared to temperate species, as tropical trees have shown declines in growth and photosynthesis at elevated temperatures. However, regional and global vegetation models lack the data needed to accurately represent such physiological responses to increased temperatures, especially for tropical forests. To address this need, we compared instantaneous photosynthetic temperature responses of mature canopy foliage, leaf temperatures, and air temperatures across vertical canopy gradients in three forest types: tropical wet, tropical moist, and temperate deciduous. Temperatures at which maximum photosynthesis occurred were greater in the tropical forests canopies than the temperate canopy (30 ± 0.3 °C vs. 27 ± 0.4 °C). However, contrary to expectations that tropical species would be functioning closer to threshold temperatures, photosynthetic temperature optima was exceeded by maximum daily leaf temperatures, resulting in sub-optimal rates of carbon assimilation for much of the day, especially in upper canopy foliage (>10 m). If trees are unable to thermally acclimate to projected elevated temperatures, these forests may shift from net carbon sinks to sources, with potentially dire implications to climate feedbacks and forest community composition.

For more accurate projections of both the global carbon (C) cycle and the changing climate, a critical current need is to improve the representation of tropical forests in Earth system models. Tropical forests exchange more C, energy, and water with the atmosphere than any other class of land ecosystems. Further, tropical-forest C cycling is likely responding to the rapid global warming, intensifying water-stress, and increasing atmospheric CO2 levels. Projections of the future C balance of the tropics vary widely among global models. A current effort of the modeling community, ILAMB (the International Land Model Benchmarking Project), is to compile robust observations that can be used to improve the accuracy and realism of the land models for all major biomes. Our goal with this paper is to identify field observations of tropical-forest ecosystem C stocks and fluxes that can support this effort. We propose criteria for reference-level field data from this biome and present a set of documented examples from old-growth lowland tropical forests. We offer these as a starting point towards the goal of a regularly updated consensus set of benchmark field observations of C-cycling in tropical forests.

The removal of nutrients from senescing tissues, nutrient resorption, is a key strategy for conserving nutrients in plants. However, our understanding of what drives patterns of nutrient resorption in tropical trees is limited. We examined the effects of nutrient sources (stand-level and tree-level soil fertility) and sinks (reproductive effort) on nitrogen (N) and phosphorus (P) resorption. We evaluated resorption efficiency (percentage of original nutrients removed during senescence) and resorption proficiency (indicated by senesced-leaf nutrient concentrations) in a symbiotic N-fixing tree species, Pentaclethra macroloba, common to tropical forests in Costa Rica. Although tree-level soil P alone did not drive patterns in nutrient resorption, P efficiency and proficiency declined with increasing tree-level soil P when reproductive status was also considered. Nutrient resorption declined with increasing tree-level soil P in trees that were actively fruiting or that experienced high seedfall the year prio...

© 2016 Bouskill, Wood, Baran, Ye, Bowen, Lim, Zhou, Van Nostrand, Nico, Northen, Silver and Brodie. 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.

© 2016 Bouskill, Wood, Baran, Hao, Ye, Bowen, Lim, Nico, Holman, Gilbert, Silver, Northen and Brodie. 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 CO2efflux following wet-up in drought plots relative to control plots.

Many tropical rain forest regions are at risk of increased future drought. The net effects of drought on forest ecosystem functioning will be substantial if important ecological thresholds are passed. However, understanding and predicting these effects is challenging using observational studies alone. Field-based rainfall exclusion (canopy throughfall exclusion; TFE) experiments can offer mechanistic insight into the response to extended or severe drought and can be used to help improve model-based simulations, which are currently inadequate. Only eight TFE experiments have been reported for tropical rain forests. We examine them, synthesizing key results and focusing on two processes that have shown threshold behavior in response to drought: (1) tree mortality and (2) the efflux of carbon dioxdie from soil, soil respiration. We show that: (a) where tested using large-scale field experiments, tropical rain forest tree mortality is resistant to long-term soil moisture deficit up to a threshold of 50% of the water that is extractable by vegetation from the soil, but high mortality occurs beyond this value, with evidence from one site of increased autotrophic respiration, and (b) soil respiration reaches its peak value in response to soil moisture at significantly higher soil moisture content for clay-rich soils than for clay-poor soils. This first synthesis of tropical TFE experiments offers the hypothesis that low soil moisture–related thresholds for key stress responses in soil and vegetation may prove to be widely applicable across tropical rain forests despite the diversity of these forests.

Global climate models project a decrease in the magnitude of precipitation in tropical regions. Changes in rainfall patterns have important implications for the moisture content and redox status of tropical soils, yet little is known about how these changes may affect microbial community structure. Specifically, does exposure to prior stress confer increased resistance to subsequent perturbation? Here we reduced the quantity of precipitation throughfall to tropical forest soils in the Luquillo Mountains, Puerto Rico. Treatments included newly established throughfall exclusion plots (de novo excluded), plots undergoing reduction for a second time (pre-excluded) and ambient control plots. Ten months of throughfall exclusion led to a small but statistically significant decline in soil water potential and bacterial populations clearly adapted to increased osmotic stress. Although the water potential decline was small and microbial biomass did not change, phylogenetic diversity in the de novo-excluded plots decreased by ∼40% compared with the control plots, yet pre-excluded plots showed no significant change. On the other hand, the relative abundances of bacterial taxa in both the de novo-excluded and pre-excluded plots changed significantly with throughfall exclusion compared with control plots. Changes in bacterial community structure could be explained by changes in soil pore water chemistry and suggested changes in soil redox. Soluble iron declined in treatment plots and was correlated with decreased soluble phosphorus concentrations, which may have significant implications for microbial productivity in these P-limited systems.

The influence of environmental gradients on the foliar nutrient economy of forests has been well documented; however, we have little understanding of what drives variability among individuals within a single forest stand, especially tropical forests. We evaluated inter- and intra-specific variation in nutrient resorption, foliar nutrient concentrations and physical leaf traits of nine canopy tree species within a 1-ha secondary tropical rain forest in northeastern Costa Rica. Both nitrogen (N) and phosphorus (P) resorption efficiency (RE) and proficiency of the nine tree species varied significantly among species, but not within. Both N and P RE were significantly negatively related to leaf specific strength. Green leaf N and P concentrations were strongly negatively related to leaf mass per area, and senesced leaf nutrient concentrations were significantly positively related to green leaf nutrient concentrations. This study reveals a strong influence of physical leaf traits on foliar nutrient and resorption traits of co-occurring species in a secondary wet tropical forest stand.

The productivity of many tropical wet forests is generally limited by bioavailable phosphorus (P). Microbial activity is a key regulator of P availability in that it determines both the supply of P through organic matter decomposition and the depletion of bioavailable P through microbial uptake. Both microbial uptake and mineralization occur rapidly, and their net effect on P availability varies with soil moisture, temperature, and soil organic matter quantity and quality. Exploring the mechanisms driving P availability at fine temporal scales can provide insight into the coupling of carbon, water, and nutrient cycles, and ultimately, the response of tropical forests to climate change. Despite the recognized importance of P cycling to the dynamics of wet tropical forests and their potential sensitivity to short-term fluctuations in bioavailable P, the diurnal pattern of P remains poorly understood. This study quantifies diurnal fluctuations in labile soil P and evaluates the importance of biotic and abiotic factors in driving these patterns. To this end, measurements of labile P were made every other hour in a Costa Rican wet tropical forest oxisol. Spatial and temporal variation in Bray-extractable P were investigated in relation to ecosystem carbon flux, soil CO2 efflux, soil moisture, soil temperature, solar radiation, and sap-flow velocity. Spatially averaged bi-hourly (every two hours) labile P ranged from 0.88 to 2.48 microg/g across days. The amplitude in labile P throughout the day was 0.61-0.82 microg/g (41-54% of mean P concentrations) and was characterized by a bimodal pattern with a decrease at midday. Labile P increased with soil CO2 efflux and soil temperature and declined with increasing sap flow and solar radiation. Together, soil CO2 efflux, soil temperature, and sap flow explained 86% of variation in labile P.

Litter-induced pulses of nutrient availability could play an important role in the productivity and nutrient cycling of forested ecosystems, especially tropical forests. Tropical forests experience such pulses as a result of wet-dry seasonality and during major climatic events, such as strong El Ninos. We hypothesized that (1) an increase in the quantity and quality of litter inputs would stimulate leaf litter production, woody growth, and leaf litter nutrient cycling, and (2) the timing and magnitude of this response would be influenced by soil fertility and forest age. To test these hypotheses in a Costa Rican wet tropical forest, we established a large-scale litter manipulation experiment in two secondary forest sites and four old-growth forest sites of differing soil fertility. In replicated plots at each site, leaves and twigs (,2 cm diameter) were removed from a 400-m 2 area and added to an adjacent 100-m 2 area. This transfer was the equivalent of adding 5-25 kg/ha of organic P to the forest floor. We analyzed leaf litter mass, (N) and (P), and N and P inputs for addition, removal, and control plots over a two-year period. We also evaluated basal area increment of trees in removal and addition plots. There was no response of forest productivity or nutrient cycling to litter removal; however, litter addition significantly increased leaf litter production and N and P inputs 4-5 months following litter application. Litter production increased as much as 92%, and P and N inputs as much as 85% and 156%, respectively. In contrast, litter manipulation had no significant effect on woody growth. The increase in leaf litter production and N and P inputs were significantly positively related to the total P that was applied in litter form. Neither litter treatment nor forest type influenced the temporal pattern of any of the variables measured. Thus, environmental factors such as rainfall drive temporal variability in litter and nutrient inputs, while nutrient release from decomposing litter influences the magnitude. Seasonal or annual variation in leaf litter mass, such as occurs in strong El Nino events, could positively affect leaf litter nutrient cycling and forest productivity, indicating an ability of tropical trees to rapidly respond to increased nutrient availability.

We experimentally manipulated forest floor litter to investigate the influence of litter quality and quantity on soil properties over the short-term (weeks to months) in a wet tropical forest in NE Costa Rica. The study included old growth forest on high fertility soils, old growth forest on low fertility soils, and secondary forest on intermediate fertility soils. Forest floor litter was removed from a 16 m2 area and added to an adjacent 4 m2 area in March 2003, resulting in a one to four-fold increase in the annual litter input to the forest floor. We created three addition, three removal and three control plots per forest type. We measured treatment effects on variation in soil moisture, temperature, pH, and Bray-1 P (plant available) over a 5-month period that captured the dry-wet season transition. Litter manipulation had no effect on any of the soil properties measured during the 5-month study period. Significant variability through time and a similar temporal pattern across the three forest stands suggest that climatic variability is driving short-term patterns in these soil properties rather than seasonal inputs of litter. In general, soils were warmer, drier and more basic with higher available P during dry season months. Even in wet tropical forests, small variability in climate can play an important role in soil dynamics over periods of weeks to months. Although litter manipulation did not influence soil properties over the 5-month study period, a longer lag may exist between the timing of litter inputs and the influence of that litter on soil properties, especially plant available P.

The stoichiometry of resources may explain bottom-up regulation of higher trophic levels. We tested the effects of soil and litter nutrient stoichiometry on the invertebrate litter fauna of a Costa Rican tropical rain forest. Animal densities were estimated from 15 sites across a phosphorus gradient. The density of the invertebrate litter fauna varied considerably, and was strongly tied to soil and litter phosphorus concentrations. An increase in phosphorus concentrations corresponded with an equally proportionate increase in animal densities. Natural variation in nutrient levels can thus serve as a predictor of density in a highly diverse tropical animal community.

By assessing current leaf litter nutrient dynamics, we may be able to predict responses of nutrient cycling in tropical ecosystems to future environmental change. The goal of this study was to assess whether nutrient cycling is related to seasonal variation in rainfall in a wet tropical forest. We examined leaf litter of an old-growth tropical rain forest in N.E. Costa Rica over a 4-year period to explore seasonal and inter-annual changes in leaf litter nutrient concentrations, and to evaluate potential short- and long-term drivers of variation in litter nutrient concentration, particularly that of phosphorus (P) and nitrogen (N). We also examined the temporal dynamics of calcium, potassium, and magnesium in the leaf litter. Leaf litter [P] and %N changed significantly with time, both seasonally and inter-annually. Seasonal changes in leaf litter [P] were strongly positively correlated with rainfall from the previous 2 weeks; cations, however, were inversely related to this measure of current rainfall, while %N was not related to rainfall. We propose that the positive relationship between current rainfall and leaf litter [P] is due to a response by the vegetation to an increase in nutrient availability and uptake. In contrast, given the negative relationship between current rainfall and cation concentrations, leaching from live leaf tissue is a more likely driver of short-term changes in cations. Should global climate change include altered rainfall patterns in this biome, one class of ecosystem-level responses could be significant changes in P and cation cycling.

Although tropical forests account for only a fraction of the planet's terrestrial surface, they exchange more carbon dioxide with the atmosphere than any other biome on Earth, and thus play a disproportionate role in the global climate. In the next 20 years, the tropics will experience unprecedented warming, yet there is exceedingly high uncertainty about their potential responses to this imminent climatic change. Here, we prioritize research approaches given both funding and logistical constraints in order to resolve major uncertainties about how tropical forests function and also to improve predictive capacity of earth system models. We investigate overall model uncertainty of tropical latitudes and explore the scientific benefits and inevitable trade-offs inherent in large-scale manipulative field experiments. With a Coupled Model Intercomparison Project Phase 5 analysis, we found that model variability in projected net ecosystem production was nearly 3 times greater in the tropics than for any other latitude. Through a review of the most current literature, we concluded that manipulative warming experiments are vital to accurately predict future tropical forest carbon balance, and we further recommend the establishment of a network of comparable studies spanning gradients of precipitation, edaphic qualities, plant types, and/or land use change. We provide arguments for long-term, single-factor warming experiments that incorporate warming of the most biogeochemically active ecosystem components (i.e. leaves, roots, soil microbes). Hypothesis testing of underlying mechanisms should be a priority, along with improving model parameterization and constraints. No single tropical forest is representative of all tropical forests; therefore logistical feasibility should be the most important consideration for locating large-scale manipulative experiments. Above all, we advocate for multi-faceted research programs, and we offer arguments for what we consider the most powerful and urgent way forward in order to improve our understanding of tropical forest responses to climate change.

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.

[1] Soil moisture is a key driver of biogeochemical processes in terrestrial ecosystems, strongly affecting carbon (C) and nutrient availability as well as trace gas production and consumption in soils. Models predict increasing drought frequency in tropical forest ecosystems, which could feed back on future climate change directly via effects on trace gasdynamics and indirectly through changes in nutrient availability. We used throughfall exclusion shelters to determine effects of short-term (3 month) drought on trace gas fluxes and nutrient availability in humid tropical forests in Puerto Rico. Exclusion and control plots were replicated within and across three topographic zones (ridge, slope, valley) to account for spatial heterogeneity typical of these ecosystems. Throughfall exclusion reduced soil moisture in all sites and lowered exchangeable phosphorus (P) on ridges and slopes. Drought decreased soil carbon dioxide (CO2) emissions by 30% in ridge sites and 28% in slope sites, and increased net methane (CH4) consumption by 480% in valley sites. Both valley and ridge sites became net nitrous oxide (N2O) sinks in response to soil drying. Emissions of CO2 and N2O, as well as CH4 consumption were positively related to exchangeable P and the nitrate:ammonium ratio. These findings suggest that drought has the potential to decrease net trace gas emissions from humid tropical forest soils. The differential response of trace gas emissions and nutrients from different topographic zones to drought underscores the complexity of biogeochemical cycling in these ecosystems and the importance of considering spatial heterogeneity when estimating whole system responses.

Tropical forests play a major role in regulating global carbon (C) fluxes and stocks, and even small changes to C cycling in this productive biome could dramatically affect atmospheric carbon dioxide (CO(2) ) concentrations. Temperature is expected to increase over all land surfaces in the future, yet we have a surprisingly poor understanding of how tropical forests will respond to this significant climatic change. Here we present a contemporary synthesis of the existing data and what they suggest about how tropical forests will respond to increasing temperatures. Our goals were to: (i) determine whether there is enough evidence to support the conclusion that increased temperature will affect tropical forest C balance; (ii) if there is sufficient evidence, determine what direction this effect will take; and, (iii) establish what steps should to be taken to resolve the uncertainties surrounding tropical forest responses to increasing temperatures. We approach these questions from a mass-balance perspective and therefore focus primarily on the effects of temperature on inputs and outputs of C, spanning microbial- to ecosystem-scale responses. We found that, while there is the strong potential for temperature to affect processes related to C cycling and storage in tropical forests, a notable lack of data combined with the physical, biological and chemical diversity of the forests themselves make it difficult to resolve this issue with certainty. We suggest a variety of experimental approaches that could help elucidate how tropical forests will respond to warming, including large-scale in situ manipulation experiments, longer term field experiments, the incorporation of a range of scales in the investigation of warming effects (both spatial and temporal), as well as the inclusion of a diversity of tropical forest sites. Finally, we highlight areas of tropical forest research where notably few data are available, including temperature effects on: nutrient cycling, heterotrophic versus autotrophic respiration, thermal acclimation versus substrate limitation of plant and microbial communities, below-ground C allocation, species composition (plant and microbial), and the hydraulic architecture of roots. Whether or not tropical forests will become a source or a sink of C in a warmer world remains highly uncertain. Given the importance of these ecosystems to the global C budget, resolving this uncertainty is a primary research priority.

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.