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Molly A. Cavaleri

Publications and source records attributed to Molly A. Cavaleri.

16 recordsLinked to original sources

Lowland tropical forests remain a methane sink under warming and long-term hurricane disturbance recovery

Methane (CH 4 ) is a potent greenhouse gas, and tropical forests account for roughly one–third of global atmospheric CH 4 uptake by soils. Projected warming and more frequent hurricanes in these ecosystems may alter soil CH 4 sink strength, as warmer and wetter soils enhance methanogenesis activity. We measured soil CH 4 and CO 2 efflux during the calendar summer months of 2023 and 2024 alongside continuous records of soil moisture, soil and air temperature, and precipitation in an in–situ warming experiment (TRACE) located in a lowland tropical forest in Puerto Rico, six to seven years after Hurricanes Irma and Maria (2017). The realized warming (∼1.95°C) enhanced soil respiration only in summer 2023 ( p < 0.05), but net soil CH 4 uptake was invariant in both campaigns ( p > 0.05). Instead, sampling day and between–plot variability explained soil CH 4 dynamics much more than treatment contrasts. Importantly, CH 4 uptake was consistently coupled to CO 2 efflux, suggesting tight linkages between methanotrophic and heterotrophic activities. Between treatments, CH 4 and CO 2 responses to soil temperature variation were less sensitive in warmed plots, which may suggest weak metabolic upregulation under elevated temperatures. Together, these findings indicate that lowland tropical soils remain CH 4 sink even under warming and years after hurricane disturbance, with CH 4 dynamics driven more by spatial and temporal variability than experimental warming. Long–term, high–resolution monitoring integrating soil biogeochemistry and microbial processes will be critical to determine whether the observed net CH 4 uptake signal represents a sustainable or transient response under continued warming and disturbance.

Puerto Rico, Sabana Field Research Station

Experimental warming alters free-living nitrogen fixation in a humid tropical forest

Microbial nitrogen (N) fixation accounts for c. 97% of natural N inputs to terrestrial ecosystems. These microbes can be free-living in the soil and leaf litter (asymbiotic) or in symbiosis with plants. Warming is expected to increase N-fixation rates because warmer temperatures favor the growth and activity of N-fixing microbes. We investigated the effects of warming on asymbiotic components of N fixation at a field warming experiment in Puerto Rico. We analyzed the function and composition of bacterial communities from surface soil and leaf litter samples. Warming significantly increased asymbiotic N-fixation rates in soil by 55% (to 0.002 kg ha −1 yr −1 ) and by 525% in leaf litter (to 14.518 kg ha −1 yr −1 ) . This increase in N fixation was associated with changes in the N-fixing bacterial community composition and soil nutrients. Our findings suggest that warming increases the natural N inputs from the atmosphere into this tropical forest due to changes in microbial function and composition, especially in the leaf litter. Given the importance of leaf litter in nutrient cycling, future research should investigate other aspects of N cycles in the leaf litter under warming conditions.

Luquillo Experimental Forest, northeastern Puerto

Warming induces unexpectedly high soil respiration in a wet tropical forest

Tropical forests are a dominant regulator of the global carbon cycle, exchanging more carbon dioxide with the atmosphere than any other terrestrial biome. Climate models predict unprecedented climatic warming in tropical regions in the coming decades; however, in situ field warming studies are severely lacking in tropical forests. Here we present results from an in situ warming experiment in Puerto Rico, where soil respiration responses to + 4 o C warming were assessed half-hourly for a year. Soil respiration rates were 42-204% higher in warmed relative to ambient plots, representing some of the highest soil respiration rates reported for any terrestrial ecosystem. While respiration rates were significantly higher in the warmed plots, the temperature sensitivity (Q 10 ) was 71.7% lower, pointing to a mechanistic shift. Even with reduced Q 10 , if observed soil respiration rates persist in a warmer world, the feedback to future climate could be considerably greater than previously predicted or observed.

Puerto Rico

Respiratory acclimation of tropical forest roots in response to in situ experimental warming and hurricane disturbance

Climate projections predict higher temperatures and more frequent hurricanes in the tropics. Tropical plants subjected to these stresses may respond by acclimating their physiology. We investigated tropical forest root respiration in response to in situ experimental warming and hurricane disturbance in eastern Puerto Rico. We measured mass-normalized root specific respiration, root biomass, and root traits at the Tropical Responses to Altered Climate Experiment (TRACE), where understory vegetation is warmed + 4 °C above ambient. Our measurements span 5 years, including before and after two major hurricanes, to quantify root contributions to ecosystem carbon fluxes. Experimental warming did not affect root specific respiration at a standard temperature of 25° (RSR 25 , mean = 3.89 nmol CO 2 g −1 s −1 ) or the temperature sensitivity of root respiration (Q 10 , mean = 1.75), but did result in decreased fine-root biomass, thereby decreasing area-based estimations of ecosystem-level root respiration in warmed plots by ~ 35%. RSR 25 of newer roots, which increased with increasing root nitrogen, showed greater rates 6 months after the hurricanes, but subsequently decreased after 12 months. Root specific respiration did not acclimate to higher temperatures, based on lack of adjustments in either Q 10 or RSR 25 in the warmed plots; however, decreased root biomass indicates the root contribution to soil carbon dioxide efflux was overall lower with warming. Lower root biomass may also limit nutrient and water uptake, having potential negative effects on carbon assimilation. Our results show that warming and hurricane disturbance have strong potential to affect tropical forest roots, as well as ecosystem carbon fluxes.

Ecosystems

Understory plant communities show resistance to drought, hurricanes, and experimental warming in a wet tropical forest

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.

Bosque experimental de Luquillo, Luquillo Experime

Experimental warming and its legacy effects on root dynamics following two hurricane disturbances in a wet tropical forest

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 m 2 . Following Hurricanes Irma and María 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.

Puerto Rico

Experimental warming across a tropical forest canopy height gradient reveals minimal photosynthetic and respiratory acclimation

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 ( Q 10 ) was down-regulated in heated leaves. The optimum temperatures for photosynthesis ( T opt ) decreased 3–5°C from understory to the highest canopy position, perhaps due to upper canopy stomatal conductance limitations. Guarea upper canopy T opt was similar to the mean daytime temperatures, while Ocotea canopy leaves often operated above T opt . With minimal acclimation to warmer temperatures in the upper canopy, further warming could put these forests at risk of reduced CO 2 uptake, which could weaken the overall carbon sink strength of this tropical forest.

Plant, Cell, and Environment

Only sun-lit leaves of the uppermost canopy exceed both air temperature and photosynthetic thermal optima in a wet tropical forest

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 (T opt = 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.

Agricultural and Forest Meteorology

Tropical understory herbaceous community responds more strongly to hurricane disturbance than to experimental warming

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.

Puerto Rico

Altered climate leads to positive density‐dependent feedbacks in a tropical wet forest

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.

Puerto Rico

Soil biogeochemical responses of a tropical forest to warming and hurricane disturbance

Tropical forests represent <15% of Earths terrestrial surface yet support >50% of the planets 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 an occurrence of 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 Mara 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, 43, and 46% 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.

Puerto Rico

On the shoulders of giants: Continuing the legacy of large-scale ecosystem manipulation experiments in Puerto Rico

There is a long history of experimental research in the Luquillo Experimental Forest in Puerto Rico. These experiments have addressed questions about biotic thresholds, assessed why communities vary along natural gradients, and have explored forest responses to a range of both anthropogenic and non-anthropogenic disturbances. Combined, these studies cover many of the major disturbances that affect tropical forests around the world and span a wide range of topics, including the effects of forest thinning, ionizing radiation, hurricane disturbance, nitrogen deposition, drought, and global warming. These invaluable studies have greatly enhanced our understanding of tropical forest function under different disturbance regimes and informed the development of management strategies. Here we summarize the major field experiments that have occurred within the Luquillo Experimental Forest. Taken together, results from the major experiments conducted in the Luquillo Experimental Forest demonstrate a high resilience of Puerto Rico’s tropical forests to a variety of stressors.

Luquillo Experimental Forest

Soil warming effects on tropical forests with highly weathered soils

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.

Book chapter

Temperate and tropical forest canopies are already functioning beyond their thermal thresholds for photosynthesis

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.

Forests

Infrared heater system for warming tropical forest understory plants and soils

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.

Puerto Rico

Tropical forest carbon balance in a warmer world: a critical review spanning microbial- to ecosystem-scale processes

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.

Biological Reviews