Abstract. Foliar nitrogen to phosphorus (N:P) ratios are widely used to indicate soil nutrient availability and limitation, but the foliar ratios of woody plants have proven more complicated to interpret than ratios from whole biomass of herbaceous species. This may be related to tissues in woody species acting as nutrient reservoirs during active growth, allowing maintenance of optimal N:P ratios in recently produced, fully expanded leaves (i.e., ''new'' leaves, the most commonly sampled tissue). Here we address the hypothesis that N:P ratios of newly expanded leaves are less sensitive indicators of soil nutrient availability than are other tissue types in woody plants. Seedlings of five naturally established tree species were harvested from plots receiving two years of fertilizer treatments in a lowland tropical forest in the Republic of Panama. Nutrient concentrations were determined in new leaves, old leaves, stems, and roots. For stems and roots, N:P ratios increased after N addition and decreased after P addition, and trends were consistent across all five species. Older leaves also showed strong responses to N and P addition, and trends were consistent for four of five species. In comparison, overall N:P ratio responses in new leaves were more variable across species. These results indicate that the N:P ratios of stems, roots, and older leaves are more responsive indicators of soil nutrient availability than are those of new leaves. Testing the generality of this result could improve the use of tissue nutrient ratios as indices of soil nutrient availability in woody plants.

Quantifying nutrient limitation of primary productivity is a fundamental task of terrestrial ecosystem ecology, but in a high carbon dioxide environment it is even more critical that we understand potential nutrient constraints on plant growth. Ecologists often manipulate nutrients with fertilizer to assess nutrient limitation, yet for a variety of reasons, nutrient fertilization experiments are either impractical or incapable of resolving ecosystem responses to some global changes. The challenges of conducting large, in situ fertilization experiments are magnified in forests, especially the high-diversity forests common throughout the lowland tropics. A number of methods, including fertilization experiments, could be seen as tools in a toolbox that ecologists may use to attempt to assess nutrient limitation, but there has been no compilation or synthetic discussion of those methods in the literature. Here, we group these methods into one of three categories (indicators of soil nutrient supply, organismal indicators of nutrient limitation, and lab-based experiments and nutrient depletions), and discuss some of the strengths and limitations of each. Next, using a case study, we compare nutrient limitation assessed using these methods to results obtained using large-scale fertilizations across the Hawaiian Archipelago. We then explore the application of these methods in high-diversity tropical forests. In the end, we suggest that, although no single method is likely to predict nutrient limitation in all ecosystems and at all scales, by simultaneously utilizing a number of the methods we describe, investigators may begin to understand nutrient limitation in complex and diverse ecosystems such as tropical forests. In combination, these methods represent our best hope for understanding nutrient constraints on the global carbon cycle, especially in tropical forest ecosystems.

Leaching is a mechanism for the release of nutrients from litter or senesced leaves that can drive interactions among plants, microbes, and soil. Although leaching is well established in conceptual models of litter decomposition, potential nutrient solubility of mineral elements from recently senesced litter has seldom been quantified. Using a standardized extraction (1:50 litter-to-water ratio and four-hour extraction) and recently senesced leaf litter of 41 tropical tree and liana species, we investigated how solubility varies among elements, and whether the solubility of elements could be predicted by litter traits (e.g., lignin, total element concentrations). In addition, we investigated nutrient forms (i.e., inorganic and organic) and ratios in leachate. Water-soluble elements per unit litter mass were strongly predicted by total initial litter element concentrations for potassium (K; r2 = 0.79), sodium (Na; r2 = 0.51) and phosphorus (P; r2 = 0.66), while a significant but weaker positive relationship was found for nitrogen (N; r2 = 0.36). There was no significant relationship for carbon (C) or calcium (Ca). Element-specific solubility varied markedly. On average 100% of total K, 35% of total P, 28% of total Na, 5% of total N, 4% of total Ca, and 3% of total C were soluble. For soluble P, 90% was inorganic orthophosphate. The high solubility of K, Na, and P as inorganic orthophosphate suggests that these nutrients can become rapidly available to litter microbes with no metabolic cost. Few common predictors of decomposition rates were correlated with element solubility, although soluble C (milligrams per gram of litter) was negatively related to lignin content (r2 = 0.19; P < 0.004). Solubility of elements was linked within a species: when a species ranked high in the soluble fraction of one element, it also ranked high in the solubility of other elements. Overall nutrient-specific patterns of solubility from recently senesced litter emphasize that litter elements cannot be treated equally in our conceptual and empirical models of decomposition. The relatively high potential solubility of P as orthophosphate from fresh litter advances our understanding of ecological stoichiometric ratios and nutrient bioavailability in tropical forests.

BackgroundLocal plant-soil associations are commonly studied at the species-level, while associations at the level of nodes within a phylogeny have been less well explored. Understanding associations within a phylogenetic context, however, can improve our ability to make predictions across systems and can advance our understanding of the role of evolutionary history in structuring communities.Methodology/Principal FindingsHere we quantified evolutionary signal in plant-soil associations using a DNA sequence-based community phylogeny and several soil variables (e.g., extractable phosphorus, aluminum and manganese, pH, and slope as a proxy for soil water). We used published plant distributional data from the 50-ha plot on Barro Colorado Island (BCI), Republic of Panamá. Our results suggest some groups of closely related species do share similar soil associations. Most notably, the node shared by Myrtaceae and Vochysiaceae was associated with high levels of aluminum, a potentially toxic element. The node shared by Apocynaceae was associated with high extractable phosphorus, a nutrient that could be limiting on a taxon specific level. The node shared by the large group of Laurales and Magnoliales was associated with both low extractable phosphorus and with steeper slope. Despite significant node-specific associations, this study detected little to no phylogeny-wide signal. We consider the majority of the ‘traits’ (i.e., soil variables) evaluated to fall within the category of ecological traits. We suggest that, given this category of traits, phylogeny-wide signal might not be expected while node-specific signals can still indicate phylogenetic structure with respect to the variable of interest.ConclusionsWithin the BCI forest dynamics plot, distributions of some plant taxa are associated with local-scale differences in soil variables when evaluated at individual nodes within the phylogenetic tree, but they are not detectable by phylogeny-wide signal. Trends highlighted in this analysis suggest how plant-soil associations may drive plant distributions and diversity at the local-scale.

To investigate causes of tree species distributions across soil resources in northern Michigan, we conducted a seedling transplant experiment with five species showing different site affinities: Acer saccharum Marsh. (sugar maple), Prunus serotina Ehrh. (black cherry), and Fraxinus americana L. (white ash), which are associated with high-fertility mesic moraine; Quercus rubra L. (red oak), associated with intermediate sites; and Quercus velutina Lam. (black oak), associated with low-fertility droughty outwash sites. Seedlings were planted in plots stratified across variation in light and soil nutrient and water availability. After one growing season, under 14%–27% canopy openness, species tended to trade off between high survival on outwash versus high relative growth rate of root + stem mass (RGRrs) on moraine. The high survivorship of black and red oak on outwash was associated with greater root and whole-plant mass in comparison with sugar maple, white ash, and black cherry. High RGRrs on high-fertility moraine for these latter species was associated with high fine root area per unit whole-plant mass and plasticity to increase specific root area in response to increased soil resources. We did not detect a similar survival–growth trade-off for seedlings grown at lower light (3%–10%) on intermediate versus high-fertility sites. Overall, these results suggest that species distributions across soil resource gradients can in part be explained by a trade-off between tolerance of low soil resources versus competitive ability (i.e., growth) under high soil resources.

In strongly weathered soils, leaf litter not only returns phosphorus (P) to the soil environment, it may also modify soil properties and soil solution chemistry, with the potential to decrease phosphate sorption and increase plant available P. Using a radioactive phosphate tracer ( 32 P) and 1 h laboratory incubations we investigated the effect of litter inputs on phosphate sorption over two time scales: (1) long-term field litter manipulations (litter addition, control and litter removal) and (2) pulses of litter leachate (i.e. water extracts of leaf litter) from five species. Leachate pulse effects were compared to a simulated throughfall, which served as a control solution. Soil receiving long-term doubling of leaf litter maintained five-fold more phosphate in solution than the litter removal soil. In addition to the quantity of phosphate sorbed, the field litter addition treatment decreased the strength of phosphate sorption, as evaluated through extraction of sorbed 32 P using a weakly acidic ammonium fluoride solution (Bray 1). In litter removal soil, leachate pulses significantly reduced phosphate sorption in comparison to the throughfall control for all five species evaluated. However, the ability of leachate pulses to reduce phosphate sorption decreased when soil had received field litter inputs. Across soils the effect of leachate pulses on phosphate sorption increased with net sorption of dissolved organic C, with the exception of leachate from one species that had a higher index of aromatic C concentration. These results demonstrate that litter inputs, as both long-term inputs and short-term leachate pulses, can decrease the quantity and strength of phosphate sorption, which may increase the biological availability of this key nutrient.

Studies of biodiversity and terrestrial ecosystem functioning have concentrated almost exclusively on temperate grasslands. To broaden the reach of biodiversity‐functioning research, five fast growing species, comprising three eudicot trees and two congeneric palms (none symbiotic with nitrogen‐fixing microorganisms), were grown for 13 yr in a replacement‐series mixture and monocultures on a fertile soil in a high‐rainfall area of lowland Costa Rica. The mixture accrued more biomass and had greater net productivity than the average, but not the most productive, monoculture. Relative Land Output (a measure of comparative yield) increased steeply. The combined evidence points to an increase in resource partitioning or facilitation among species over time. Spatial partitioning of aboveground space (for light capture) and soil (possibly for retrieval of deep nitrogen), and facilitation of phosphorus availability by one species, are mechanisms that may account for the inferred complementarity. Extending the generalized findings on biodiversity–productivity relationships from well‐studied grasslands to tropical forests is warranted. Mixtures of fast growing trees can out‐perform the average of their component monocultures, whether the metric is biomass accrual or productivity. The modular growth of long‐lived structure enables arborescent species to retain crown space previously captured and may lead to increased spatial partitioning and facilitation of resources over time.

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