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.

Tropical dry forest trees and lianas differ in leaf economic spectrum traits but have overlapping water-use strategies.

Tree species in tropical dry forests employ a wide range of strategies to cope with seasonal drought, including regulation of hydraulic function. However, it is uncertain if co-occurring lianas also possess a diversity of strategies. For a taxonomically diverse group of 14 tree and 7 liana species, we measured morphological and hydraulic functional traits during an unusual drought and under non-drought conditions to determine (i) if trees have different water-use strategies than lianas and (ii) if relationships among these traits can be used to better understand how tree and liana species regulate diurnal leaf water potential (Ψdiurnal). In this Costa Rican tropical dry forest, lianas and trees had overlapping water-use strategies, but differed in many leaf economic spectrum traits. Specifically, we found that both lianas and trees employed a diversity of Ψdiurnal regulation strategies, which did not differ statistically. However, lianas and trees did significantly differ in terms of certain traits including leaf area, specific leaf area, petiole length, wood vessel diameter and xylem vessel density. All liana and tree species we measured fell along a continuum of isohydric (partial) to anisohydric (strict or extreme) Ψdiurnal regulation strategies, and leaf area, petiole length, stomatal conductance and wood vessel diameter correlated with these strategies. These findings contribute to a trait-based understanding of how plants regulate Ψdiurnal under both drought stress and sufficient water availability, and underscore that lianas and trees employ a similarly wide range of Ψdiurnal regulation strategies, despite having vastly different growth forms.

Scale-dependent variation in nitrogen cycling and soil fungal communities along gradients of forest composition and age in regenerating tropical dry forests.

Rates of ecosystem nitrogen (N) cycling may be mediated by the presence of ectomycorrhizal fungi, which compete directly with free-living microbes for N. In the regenerating tropical dry forests of Central America, the distribution of ectomycorrhizal trees is affected by succession and soil parent material, both of which may exert independent influence over soil N fluxes. In order to quantify these interacting controls, we used a scale-explicit sampling strategy to examine soil N cycling at scales ranging from the microsite to ecosystem level. We measured fungal community composition, total and inorganic N pools, gross proteolytic rate, net N mineralization and microbial extracellular enzyme activity at multiple locations within 18 permanent plots that span dramatic gradients of soil N concentration, stand age and forest composition. The ratio of inorganic to organic N cycling was correlated with variation in fungal community structure, consistent with a strong influence of ectomycorrhiza on ecosystem-scale N cycling. However, on average, > 61% of the variation in soil biogeochemistry occurred within plots, and the effects of forest composition were mediated by this local-scale heterogeneity in total soil N concentrations. These cross-scale interactions demonstrate the importance of a spatially explicit approach towards an understanding of controls on element cycling.

Nitrogen, phosphorus, and cation use efficiency in stands of regenerating tropical dry forest.

Plants on infertile soils exhibit physiological and morphological traits that support conservative internal nutrient cycling. However, potential trade-offs among use efficiencies for N, P, and cations are not well explored in species-rich habitats where multiple elements may limit plant production. We examined uptake efficiency and use efficiency of N, P, K, Ca, Mg, Al, and Na in plots of regenerating tropical dry forests spanning a gradient of soil fertility. Our aim was to determine whether plant responses to multiple elements are correlated, or whether there are trade-offs among exploitation strategies across stands varying in community composition, soil quality, and successional stage. For all elements, both uptake efficiency and use efficiency decreased as availability of the corresponding element increased. Plant responses to N, Na, and Al were uncoupled from uptake and use efficiencies for P and essential base cations, which were tightly correlated. N and P use efficiencies were associated with shifts in plant species composition along the soil fertility gradient, and there was also a trend towards increasing N use efficiency with stand age. N uptake efficiency was positively correlated with the abundance of tree species that associate with ectomycorrhizal fungi. Taken together, our results suggest that successional processes and local species composition interact to regulate plant responses to availability of multiple resources. Successional tropical dry forests appear to employ different strategies to maximize response to N vs. P and K.

Short-term precipitation exclusion alters microbial responses to soil moisture in a wet tropical forest.

Many wet tropical forests, which contain a quarter of global terrestrial biomass carbon stocks, will experience changes in precipitation regime over the next century. Soil microbial responses to altered rainfall are likely to be an important feedback on ecosystem carbon cycling, but the ecological mechanisms underpinning these responses are poorly understood. We examined how reduced rainfall affected soil microbial abundance, activity, and community composition using a 6-month precipitation exclusion experiment at La Selva Biological Station, Costa Rica. Thereafter, we addressed the persistent effects of field moisture treatments by exposing soils to a controlled soil moisture gradient in the lab for 4 weeks. In the field, compositional and functional responses to reduced rainfall were dependent on initial conditions, consistent with a large degree of spatial heterogeneity in tropical forests. However, the precipitation manipulation significantly altered microbial functional responses to soil moisture. Communities with prior drought exposure exhibited higher respiration rates per unit microbial biomass under all conditions and respired significantly more CO2 than control soils at low soil moisture. These functional patterns suggest that changes in microbial physiology may drive positive feedbacks to rising atmospheric CO2 concentrations if wet tropical forests experience longer or more intense dry seasons in the future.