Phosphorus limitation of early growth differs between nitrogen-fixing and nonfixing dry tropical forest tree species.

Tropical forests are often characterized by low soil phosphorus (P) availability, suggesting that P limits plant performance. However, how seedlings from different functional types respond to soil P availability is poorly known but important for understanding and modeling forest dynamics under changing environmental conditions. We grew four nitrogen (N)-fixing Fabaceae and seven diverse non-N-fixing tropical dry forest tree species in a shade house under three P fertilization treatments and evaluated carbon (C) allocation responses, P demand, P-use, investment in P acquisition traits, and correlations among P acquisition traits. Nitrogen fixers grew larger with increasing P addition in contrast to non-N fixers, which showed fewer responses in C allocation and P use. Foliar P increased with P addition for both functional types, while P acquisition strategies did not vary among treatments but differed between functional types, with N fixers showing higher root phosphatase activity (RPA) than nonfixers. Growth responses suggest that N fixers are limited by P, but nonfixers may be limited by other resources. However, regardless of limitation, P acquisition traits such as mycorrhizal colonization and RPA were nonplastic across a steep P gradient. Differential limitation among plant functional types has implications for forest succession and earth system models.

Reduced ecosystem resilience quantifies fine-scale heterogeneity in tropical forest mortality responses to drought.

Sensitivity of forest mortality to drought in carbon-dense tropical forests remains fraught with uncertainty, while extreme droughts are predicted to be more frequent and intense. Here, the potential of temporal autocorrelation of high-frequency variability in Landsat Enhanced Vegetation Index (EVI), an indicator of ecosystem resilience, to predict spatial and temporal variations of forest biomass mortality is evaluated against in situ census observations for 64 site-year combinations in Costa Rican tropical dry forests during the 2015 ENSO drought. Temporal autocorrelation, within the optimal moving window of 24 months, demonstrated robust predictive power for in situ mortality (leave-one-out cross-validation R2  = 0.54), which allows for estimates of annual biomass mortality patterns at 30 m resolution. Subsequent spatial analysis showed substantial fine-scale heterogeneity of forest mortality patterns, largely driven by drought intensity and ecosystem properties related to plant water use such as forest deciduousness and topography. Highly deciduous forest patches demonstrated much lower mortality sensitivity to drought stress than less deciduous forest patches after elevation was controlled. Our results highlight the potential of high-resolution remote sensing to "fingerprint" forest mortality and the significant role of ecosystem heterogeneity in forest biomass resistance to drought.

A catastrophic tropical drought kills hydraulically vulnerable tree species.

Drought-related tree mortality is now a widespread phenomenon predicted to increase in magnitude with climate change. However, the patterns of which species and trees are most vulnerable to drought, and the underlying mechanisms have remained elusive, in part due to the lack of relevant data and difficulty of predicting the location of catastrophic drought years in advance. We used long-term demographic records and extensive databases of functional traits and distribution patterns to understand the responses of 20-53 species to an extreme drought in a seasonally dry tropical forest in Costa Rica, which occurred during the 2015 El Niño Southern Oscillation event. Overall, species-specific mortality rates during the drought ranged from 0% to 34%, and varied little as a function of tree size. By contrast, hydraulic safety margins correlated well with probability of mortality among species, while morphological or leaf economics spectrum traits did not. This firmly suggests hydraulic traits as targets for future research.

Resilience of seed production to a severe El Niño-induced drought across functional groups and dispersal types.

More frequent and severe El Niño Southern Oscillations (ENSO) are causing episodic periods of decreased rainfall. Although the effects of these ENSO-induced droughts on tree growth and mortality have been well studied, the impacts on other demographic rates such as reproduction are less well known. We use a four-year seed rain dataset encompassing the most severe ENSO-induced drought in more than 30 years to assess the resilience (i.e., resistance and recovery) of the seed composition and abundance of three forest types in a tropical dry forest. We found that forest types showed distinct differences in the timing, duration, and intensity of drought during the ENSO event, which likely mediated seed composition shifts and resilience. Drought-deciduous species were particularly sensitive to the drought with overall poor resilience of seed production, whereby seed abundance of this functional group failed to recover to predrought levels even two years after the drought. Liana and wind-dispersed species were able to maintain seed production both during and after drought, suggesting that ENSO events promote early successional species or species with a colonization strategy. Combined, these results suggest that ENSO-induced drought mediates the establishment of functional groups and dispersal types suited for early successional conditions with more open canopies and reduced competition among plants. The effects of the ENSO-induced drought on seed composition and abundance were still evident two years after the event suggesting the recovery of seed production requires multiple years that may lead to shifts in forest composition and structure in the long term, with potential consequences for higher trophic levels like frugivores.

Plant community responses to stand-level nutrient fertilization in a secondary tropical dry forest.

The size of the terrestrial carbon (C) sink is mediated by the availability of nutrients that limit plant growth. However, nutrient controls on primary productivity are poorly understood in the geographically extensive yet understudied tropical dry forest biome. To examine how nutrients influence above- and belowground biomass production in a secondary, seasonally dry tropical forest, we conducted a replicated, fully factorial nitrogen (N) and phosphorus (P) fertilization experiment at the stand scale in Guanacaste, Costa Rica. The production of leaves, wood, and fine roots was monitored through time; root colonization by mycorrhizal fungi and the abundance of N-fixing root nodules were also quantified. In this seasonal forest, interannual variation in rainfall had the largest influence on stand-level productivity, with lower biomass growth under drought. By contrast, aboveground productivity was generally not increased by nutrient addition, although fertilization enhanced growth of individual tree stems in a wet year. However, root growth increased markedly and consistently under P addition, significantly altering patterns of stand-level biomass allocation to above- vs. belowground compartments. Although nutrients did not stimulate total biomass production at the community scale, N-fixing legumes exhibited a twofold increase in woody growth in response to added P, accompanied by a dramatic increase in the abundance of root nodules. These data suggest that the relationship between nutrient availability and primary production in tropical dry forest is contingent on both water availability and plant functional diversity.