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Vargas G., German

Publications and source records attributed to Vargas G., German.

Aridity and forest age mediate landscape scale patterns of tropical forest resistance to cyclonic storms

Abstract Cyclonic storms, or hurricanes, are expected to intensify as ocean heat energy rises due to climate change. Ecological theory suggests that tropical forest resistance to hurricanes should increase with forest age and wood density. However, most data on hurricane effects on tropical forests come from a limited number of well‐studied long‐term monitoring sites, restricting our capacity to evaluate the resistance of tropical forests to hurricanes across broad environmental gradients. In this study, we assessed whether forest age and aridity mediate the effects of hurricanes Irma and Maria in Puerto Rico, Vieques and Culebra islands. We leveraged functional trait data for 410 tree species, remotely sensed measurements of canopy height and cover, along with data on forest stand characteristics of 180 of 338 forest monitoring plots, each covering an area of 0.067 ha. The plots represent a broad mean annual precipitation (MAP) gradient from 701 to 4598 mm and a complex mosaic of forest age from 5 to around 85 years since deforestation. Hurricanes resulted in a 25% increase in basal area mortality rates, a 45% decrease in canopy height and a 21% reduction in canopy cover. These effects intensified with forest age, even after considering proximity to the hurricane path. The links between forest age and hurricane disturbances were likely due the prevalence of tall canopies. Tall forest canopies were strongly linked with low community‐weighted wood density (WD). These characteristics were on average more common in moist and wet forests (MAP >1250 mm). Conversely, dry forests were dominated by short species with high wood density (WD > 0.6 g cm −3 ) and did not show significant increases in basal area mortality rates after the hurricanes. Synthesis . Our findings show that selection towards drought‐tolerant traits across aridity gradients, such as short stature and dense wood, enhances resistance to hurricanes. However, forest age modulated responses to hurricanes, with older forests being less resistant across the islands. This evidence highlights the importance of considering the intricate links between ecological succession and plant function when forecasting tropical forests’ responses to increasingly strong hurricanes.

Vargas G., German↗

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

Summary 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.

59 BASIC BIOLOGICAL SCIENCES↗

Leaf habit affects the distribution of drought sensitivity but not water transport efficiency in the tropics

Abstract Considering the global intensification of aridity in tropical biomes due to climate change, we need to understand what shapes the distribution of drought sensitivity in tropical plants. We conducted a pantropical data synthesis representing 1117 species to test whether xylem‐specific hydraulic conductivity (K S ), water potential at leaf turgor loss (Ψ TLP ) and water potential at 50% loss of K S (Ψ P50 ) varied along climate gradients. The Ψ TLP and Ψ P50 increased with climatic moisture only for evergreen species, but K S did not. Species with high Ψ TLP and Ψ P50 values were associated with both dry and wet environments. However, drought‐deciduous species showed high Ψ TLP and Ψ P50 values regardless of water availability, whereas evergreen species only in wet environments. All three traits showed a weak phylogenetic signal and a short half‐life. These results suggest strong environmental controls on trait variance, which in turn is modulated by leaf habit along climatic moisture gradients in the tropics.

54 ENVIRONMENTAL SCIENCES↗

Increasing Liana Abundance and Associated Reductions in Tree Growth in Secondary Seasonally Dry Tropical Forest

Lianas are thought to be increasing and altering tree growth and ecosystem productivity in tropical forests, but less research has focused on secondary or seasonally dry tropical forest. We report on an 11-year study of tree growth and liana presence from Guanacaste, Costa Rica, where we measured the diameter growth and liana presence on more than 1,700 trees in regenerating forest of different ages. We find that the proportion of trees without lianas is decreasing and the number of trees with lianas occupying more than 10% of tree’s crowns is increasing. We also find that lianas are affecting the diameter growth of trees. The 11-year average relative growth rates of trees with lianas in more than 10% of the tree’s crown are lower than the relative growth of trees with no lianas or lianas in less than 10% of their crown. Year-to-year, tree relative growth rate is related to annual precipitation and tree diameter. However, trees that were heavily infested with lianas (i.e., with lianas in more than 50% of their crowns) had lower relative growth and a weaker precipitation-growth relationship. This work underscores the value of long-term longitudinal data in secondary forest and adds critical data on dry forest liana abundance change.

54 ENVIRONMENTAL SCIENCES↗

Intra-annual variation in microclimatic conditions in relation to vegetation type and structure in two tropical dry forests undergoing secondary succession

Microclimate acts as a strong filter on species performance in restored and regenerating forests, particularly in seasonally dry tropical forests (SDTF). Yet few studies have measured microclimate patterns across succession in SDTF. Furthermore, although dynamic vegetation models simulate microclimate, evaluation of these simulated variables with field observations has been relatively uncommon. Here, we investigated the seasonal patterns of soil temperature and soil water in naturally regenerated and planted successional vegetation in SDTF in Costa Rica and Puerto Rico, using complementary approaches of intensive field observations and simulation modeling with the Ecosystem Demography model. We found that plots representing later successional stages were wetter on average, but only during the dry season. During the wet season, mean soil water did not differ across vegetation types, but open, early successional vegetation experienced more frequent extreme wet and dry conditions than older forest and plantations. Soil temperature tended to decline with forest structure, and later successional vegetation also experienced less extreme daily temperature fluctuations. Basal area and leaf area index were the best predictors of differences in soil water and temperature across plots. Model simulations were consistent with observations of wet season soil temperature and soil water, but the model failed to reproduce dry season soil moisture dynamics, suggesting that further work is needed to reduce model biases in microclimate variables. Collectively, our results imply that common assumptions about how microclimates influence successional processes in SDTF should be revisited.

54 ENVIRONMENTAL SCIENCES↗

Two Co-occurring Liana Species Strongly Differ in Their Hydraulic Traits in a Water-Limited Neotropical Forest

Lianas are a key growth form in tropical forests. They are believed to be strong competitors for water, thanks to their presumed efficient vascular systems. However, despite being a large polyphyletic group, they are currently often considered as a functionally homogeneous entity. In this study, we challenged this assumption by estimating the variability in hydraulic traits of two common, co-occurring liana species in a water-limited environment, namely, a seasonally dry tropical forest in Costa Rica. We measured vulnerability to embolism at the leaf and branch levels using two different methods (optical and acoustic vulnerability) and found that both species had very different hydraulic properties. Compared to reported P 50 values in literature, we found two extreme P 50 values: a low value for Bignonia diversifolia (−4.30 ± 0.54 MPa at the leaf level; −7.42 ± 0.54 MPa at the branch level) and a high value for Cissus microcarpa (−1.07 ± 0.14 at the leaf level; −1.20 ± 0.05 MPa at the branch level). Furthermore, B. diversifolia had a higher apparent modulus of elasticity in the radial direction (556.6 ± 401.0 MPa) and a variable midday water potential. On the other hand, C. microcarpa had a low apparent modulus of elasticity in the radial direction (37.8 ± 26.3 MPa) and a high branch water content, which enabled the species to keep its water potential stable during the dehydration experiments and during a drought period in the field. This mechanism may enable this species to coexist with species that are more resistant to drought-induced embolisms such as B. diversifolia . Although only two species were studied, considerable overlap was found between the range of hydraulic properties of trees growing in the same location and trees and lianas growing in two forests in Panama. These findings demonstrate that lianas cannot be considered as a homogeneous group and call for further research into the intra-growth form diversity of liana properties.

Coppieters, Kasper↗

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, in this study, 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 R 2 = 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.

59 BASIC BIOLOGICAL SCIENCES↗

Above-ground net primary productivity in regenerating seasonally dry tropical forest: Contributions of rainfall, forest age and soil

Identifying factors controlling forest productivity is critical to understanding forest-climate change feedbacks, modelling vegetation dynamics and carbon finance schemes. However, little research has focused on productivity in regenerating tropical forests which are expanding in their fraction of global area have an order of magnitude larger carbon uptake rates relative to older forest. Here We examined above-ground net primary productivity (ANPP) and its components (wood production and litterfall) over 10 years in forest plots that vary in successional age, soil characteristics and species composition using band dendrometers and litterfall traps in regenerating seasonally dry tropical forests in northwestern Costa Rica. We show that the components of ANPP are differentially driven by age and annual rainfall and that local soil variation is important. Total ANPP was explained by a combination of age, annual rainfall and soil variation. Wood production comprised 35% of ANPP on average across sites and years, and was explained by annual rainfall but not forest age. Conversely, litterfall increased with forest age and soil fertility yet was not affected by annual rainfall. In this region, edaphic variability is highly correlated with plant community composition. Thus, variation in ecosystem processes explained by soil may also be partially explained by species composition. These results suggest that future changes in annual rainfall can alter the secondary forest carbon sink, but this effect will be buffered by the litterfall flux which varies little among years. In determining the long-term strength of the secondary forest carbon sink, both rainfall and forest age will be critical variables to track. We also conclude that detailed understanding of local site variation in soils and plant community may be required to accurately predict the impact of changing rainfall on forest carbon uptake. Synthesis. We show that in seasonally dry tropical forest, annual rainfall has a positive relationship with the growth of above-ground woody tissues of trees and that droughts lead to significant reductions in above-ground productivity. These results provide evidence for climate change—carbon cycle feedbacks in the seasonal tropics and highlight the value of longitudinal data on forest regeneration.

54 ENVIRONMENTAL SCIENCES↗

Beyond leaf habit: generalities in plant function across 97 tropical dry forest tree species

Leaf habit has been hypothesized to define a linkage between the slow-fast plant economic spectrum and the drought resistance-avoidance trade-off in tropical forests (‘slow-safe vs fast-risky’). However, variation in hydraulic traits as a function of leaf habit has rarely been explored for a large number of species. We sampled leaf and branch functional traits of 97 tropical dry forest tree species from four sites to investigate whether patterns of trait variation varied consistently in relation to leaf habit along the ‘slow-safe vs fast-risky’ trade-off. Leaf habit explained from 0% to 43.69% of individual trait variation. We found that evergreen and semi-deciduous species differed in their location along the multivariate trait ordination when compared to deciduous species. While deciduous species showed consistent trait values, evergreen species trait values varied as a function of the site. Last, trait values varied in relation to the proportion of deciduous species in the plant community. Here We found that leaf habit describes the strategies that define drought avoidance and plant economics in tropical trees. However, leaf habit alone does not explain patterns of trait variation, which suggests quantifying site-specific or species-specific uncertainty in trait variation as the way forward.

59 BASIC BIOLOGICAL SCIENCES↗

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. In this work, 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.

59 BASIC BIOLOGICAL SCIENCES↗