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Allometric equations are often used to estimate plant biomass allocation to different tissue types from easier-to-measure quantities. Biomass allocation, and thus allometric equations, often differs by species and sometimes varies with nutrient availability. We measured biomass components for five nitrogen-fixing tree species ( Robinia pseudoacacia , Gliricidia sepium , Casuarina equisetifolia , Acacia koa , Morella faya ) and three non-fixing tree species ( Betula nigra , Psidium cattleianum , Dodonaea viscosa ) grown in field sites in New York and Hawaii for 4–5 years and subjected to four fertilization treatments. We measured total aboveground, foliar, main stem, secondary stem, and twig biomass in all species, and belowground biomass in Robinia pseudoacacia and Betula nigra , along with basal diameter, height, and canopy dimensions. The individuals spanned a wide size range (<1–16 cm basal diameter; 0.24–8.8 m height). For each biomass component, aboveground biomass, belowground biomass, and total biomass, we determined the following four allometric equations: the most parsimonious (lowest AIC) overall, the most parsimonious without a fertilization effect, the most parsimonious without canopy dimensions, and an equation with basal diameter only. For some species, the most parsimonious overall equation included fertilization effects, but fertilization effects were inconsistent across fertilization treatments. We therefore concluded that fertilization does not clearly affect allometric relationships in these species, size classes, and growth conditions. Our best-fit allometric equations without fertilization effects had the following R 2 values: 0.91–0.99 for aboveground biomass (the range is across species), 0.95 for belowground biomass, 0.80–0.96 for foliar biomass, 0.94–0.99 for main stem biomass, 0.77–0.98 for secondary stem biomass, and 0.88–0.99 for twig biomass. Our equations can be used to estimate overall biomass and biomass of tissue components for these size classes in these species, and our results indicate that soil fertility does not need to be considered when using allometric relationships for these size classes in these species. 

Multidimensional trait frameworks are increasingly used to understand plant strategies for growth and survival. However, it is unclear if frameworks developed at a global level can be applied in local communities and how well these frameworks—based largely on plant morphological traits—align with plant physiology and response to stress.

We tested the ability of an integrated framework of plant form and function to characterise seedling trait variation and drought response among 22 grasses and forbs common in a semi‐arid grassland. We measured above‐ground and below‐ground traits, and survival to explore how drought response is linked to three trait dimensions (resource conservation, microbial collaboration, and plant size) associated with the framework as well as non‐morphological dimensions (e.g. physiological traits) that are under‐represented in global trait frameworks.

We found support for three globally‐recognised axes representing trade‐offs in strategies associated with tissue investment (leaf nitrogen, leaf mass per area, root tissue density), below‐ground resource uptake (root diameter, specific root length), and size (shoot mass). However, in contrast to global patterns, above‐ground and below‐ground resource conservation gradients were oppositely aligned: root tissue density was positively correlated with leaf N rather than leaf mass per area. This likely reflects different investment strategies of annual and perennial herbaceous species, as fast‐growing annual species invested in lower density roots and less nitrogen‐rich leaves to maximise plant‐level carbon assimilation. Species with longer drought survival minimised water loss through small above‐ground size and low leaf‐level transpiration rates, and drought survival was best predicted by a principal component axis representing plant size.

Contrary to our expectations, drought survival in seedlings did not align with the conservation or collaboration axes suggesting that seedlings with different functional strategies can achieve similar drought survival, as long as they minimise water loss. Our results also show that within local communities, expected trait relationships could be decoupled as some plant groups achieve similar performance through different trait combinations. The effectiveness of species mean trait values in predicting drought response highlights the value of trait‐based methods as a versatile tool for understanding ecological processes locally across various ecosystems.

Read the freePlain Language Summaryfor this article on the Journal blog.

 

Nitrogen (N)‐fixing trees are thought to break a basic rule of leaf economics: higher leaf N concentrations do not translate into higher rates of carbon assimilation. Understanding how leaf N affects photosynthesis and water use efficiency (WUE) in this ecologically important group is critical.

We grew six N‐fixing and four non‐fixing tree species for 4–5 years at four fertilization treatments in field experiments in temperate and tropical regions to assess how functional type (N fixer vs. non‐fixer) and N limitation affected leaf N and how leaf N affected light‐saturated photosynthesis (A_sat), stomatal conductance (g_sw) and WUE (WUE_iand δ¹³C).

A_sat, WUE_iand δ¹³C, but notg_sw, increased with higher leaf N. Surprisingly, N‐fixing and non‐fixing trees displayed similar scaling between leaf N and these physiological variables, and this finding was supported by reanalysis of a global dataset. N fixers generally had higher leaf N than non‐fixers, even when non‐fixers were not N‐limited at the leaf level. Leaf‐level N limitation did not alter the relationship ofA_sat,g_sw, WUE_iand δ¹³C with leaf N, although it did affect the photosynthetic N use efficiency. Higher WUE was associated with higher productivity, whereas higherA_satwas not.

Synthesis: The ecological success of N‐fixing trees depends on the effect of leaf N on carbon gain and water loss. Using a field fertilization experiment and reanalysis of a global dataset, we show that high leaf‐level photosynthesis and WUE in N fixers stems from their higher average leaf N, rather than a difference between N fixers and non‐fixers in the scaling of photosynthesis and WUE with leaf N. By clarifying the mechanism by which N fixers achieve and benefit from high WUE, our results further the understanding of global N fixer distributions.

 

Abstract The study of plant functional traits and variation among and within species can help illuminate functional coordination and trade-offs in key processes that allow plants to grow, reproduce and survive. We studied 20 leaf, above-ground stem, below-ground stem and fine-root traits of 17 Costus species from forests in Costa Rica and Panama to answer the following questions: (i) Do congeneric species show above-ground and below-ground trait coordination and trade-offs consistent with theory of resource acquisition and conservation? (ii) Is there correlated evolution among traits? (iii) Given the diversity of habitats over which Costus occurs, what is the relative contribution of site and species to trait variation? We performed a principal components analysis (PCA) to assess for the existence of a spectrum of trait variation and found that the first two PCs accounted for 21.4 % and 17.8 % of the total trait variation, respectively, with the first axis of variation being consistent with a continuum of resource-acquisitive and resource-conservative traits in water acquisition and use, and the second axis of variation being related to the leaf economics spectrum. Stomatal conductance was negatively related to both above-ground stem and rhizome specific density, and these relationships became stronger after accounting for evolutionary relatedness, indicating correlated evolution. Despite elevation and climatic differences among sites, high trait variation was ascribed to individuals rather than to sites. We conclude that Costus species present trait coordination and trade-offs that allow species to be categorized as having a resource-acquisitive or resource-conservative functional strategy, consistent with a whole-plant functional strategy with evident coordination and trade-offs between above-ground and below-ground function. Our results also show that herbaceous species and species with rhizomes tend to agree with trade-offs found in more species-rich comparisons. 

Light and soil nitrogen availability can be strong controls of plant nitrogen (N) fixation, but data on how understory N‐fixing plants respond to these drivers are limited despite their important role in ecosystem N cycling. Furthermore, ecosystem N cycling can be altered by the introduction of species with nutrient use patterns that differ from natives. We assessed how N fixation of two exotic, understory species responded to varying light and soil N environments.

We sampled leaf tissue fromMimosa pudicaL.,Desmodium triflorum(L.) DC., and a nonfixing reference plant (Axonopus) growing in control and two N fertilization treatments under either N‐fixing or non‐N‐fixing trees, which may alter local soil nutrient cycling, across a range of light conditions. We measured N fixation with¹⁵N isotope dilution, and ensured that N‐fixing neighbour trees were in fact fixing N. All understory plants were wild‐growing species not native to the study location.

DesmodiumandMimosaacquired 82.6% and 71.6% of their nitrogen from fixation (%N_dfa) in the control, compared to 66.8% and 58.1% in the +10 g N m⁻² year⁻¹treatment and 73.1% and 64.7% in the +15 g N m⁻² year⁻¹treatment. These subtle %N_dfadifferences across fertilization treatments were more apparent at low light availability and disappeared at high light availability. The amount of N fixed by neighbouring trees did not influence %N_dfain the understory species.

Synthesis. Our study shows some differences in N fixation across different nutrient environments at low light for two N‐fixing species, though the changes were small, and both species derived most of their N from fixation. These findings imply that introduced N‐fixing species could exacerbate ecosystem N enrichment, particularly under high soil N conditions.

 

Biodiversity losses are a major driver of global changes in ecosystem functioning. While most studies of the relationship between biodiversity and ecosystem functioning have examined randomized species losses, trait-based filtering associated with species-specific vulnerability to drivers of diversity loss can strongly influence how ecosystem functioning responds to declining biodiversity. Moreover, the responses of ecosystem functioning to diversity loss may be mediated by environmental variability interacting with the suite of traits remaining in depauperate communities. We do not yet understand how communities resulting from realistic diversity losses (filtered by response traits) influence ecosystem functioning (via effect traits of the remaining community), especially under variable environmental conditions. Here, we directly test how realistic and randomized plant diversity losses influence productivity and invasion resistance across multiple years in a California grassland. Compared with communities based on randomized diversity losses, communities resulting from realistic (drought-driven) species losses had higher invasion resistance under climatic conditions that matched the trait-based filtering they experienced. However, productivity declined more with realistic than with randomized species losses across all years, regardless of climatic conditions. Functional response traits aligned with effect traits for productivity but not for invasion resistance. Our findings illustrate that the effects of biodiversity losses depend not only on the identities of lost species but also on how the traits of remaining species interact with varying environmental conditions. Understanding the consequences of biodiversity change requires studies that evaluate trait-mediated effects of species losses and incorporate the increasingly variable climatic conditions that future communities are expected to experience.

 

Forests are a significant CO₂sink. However, CO₂sequestration in forests is radiatively offset by emissions of nitrous oxide (N₂O), a potent greenhouse gas, from forest soils. Reforestation, an important strategy for mitigating climate change, has focused on maximizing CO₂sequestration in plant biomass without integrating N₂O emissions from soils. Although nitrogen (N)‐fixing trees are often recommended for reforestation because of their rapid growth on N‐poor soil, they can stimulate significant N₂O emissions from soils. Here, we first used a field experiment to show that a N‐fixing tree (Robinia pseudoacacia) initially mitigated climate change more than a non‐fixing tree (Betula nigra). We then used our field data to parameterize a theoretical model to investigate these effects over time. Under lower N supply, N‐fixers continued to mitigate climate change more than non‐fixers by overcoming N limitation of plant growth. However, under higher N supply, N‐fixers ultimately mitigated climate change less than non‐fixers by enriching soil N and stimulating N₂O emissions from soils. These results have implications for reforestation, suggesting that N‐fixing trees are more effective at mitigating climate change at lower N supply, whereas non‐fixing trees are more effective at mitigating climate change at higher N supply.

 

Symbiotic nitrogen fixation (SNF) is a key ecological process whose impact depends on the strategy of SNF regulation—the degree to which rates of SNF change in response to limitation by N versus other resources. SNF that is obligate or exhibits incomplete downregulation can result in excess N fixation, whereas a facultative SNF strategy does not. We hypothesized that tree‐based SNF strategies differed by latitude (tropical vs. temperate) and symbiotic type (actinorhizal vs. rhizobial). Specifically, we expected tropical rhizobial symbioses to display strongly facultative SNF as an explanation of their success in low‐latitude forests. In this study we used¹⁵N isotope dilution field experiments in New York, Oregon, and Hawaii to determine SNF strategies in six N‐fixing tree symbioses. Nitrogen fertilization with +10 and +15 g N m⁻² year⁻¹for 4–5 years alleviated N limitation in all taxa, paving the way to determine SNF strategies. Contrary to our hypothesis, all six of the symbioses we studied sustained SNF even at high N.Robinia pseudoacacia(temperate rhizobial) fixed 91% of its N (%N_dfa) in controls, compared to 64% and 59% in the +10 and +15 g N m⁻² year⁻¹treatments. ForAlnus rubra(temperate actinorhizal), %N_dfawas 95%, 70%, and 60%. For the tropical species, %N_dfawas 86%, 80%, and 82% forGliricidia sepium(rhizobial); 79%, 69%, and 67% forCasuarina equisetifolia(actinorhizal); 91%, 42%, and 67% forAcacia koa(rhizobial); and 60%, 51%, and 19% forMorella faya(actinorhizal). Fertilization with phosphorus did not stimulate tree growth or SNF. These results suggest that the latitudinal abundance distribution of N‐fixing trees is not caused by a shift in SNF strategy. They also help explain the excess N in many forests where N fixers are common.

 

We commonly use trait variation to characterize plant function within and among species and understand how vegetation responds to the environment. Seedling emergence is an especially vulnerable window affecting population and community dynamics, yet trait‐based frameworks often bypass this earliest stage of plant life. Here we assess whether traits vary in ecologically meaningful ways when seedlings are just days old. How do shared evolutionary history and environmental conditions shape trait expression, and can traits explain which seedlings endure drought?

We measured seedling traits in the first 4 days of life for 16 annual plant species under two water treatments, exploring trait trade‐offs, species‐level plasticity and the ability of traits to predict duration of survival under drought.

Nearly half of traits showed the imprint of evolutionary history (i.e. significant phylogenetic signal), often reflecting differences between grasses and forbs, two groups separated by a deep evolutionary split. Water availability altered trait expression in most cases, though species‐level plastic responses also reflected evolutionary history.

On average, new seedlings exhibited substantial trait variation structured as multiple trade‐offs like those found in mature plants. Some species invested in thick roots and shoots, whereas others invested in more efficient tissues. Separately, some invested in tougher roots and others in deeper roots. We also observed trade‐offs related to growth rates (fast or slow) and biomass allocation (above‐ or below‐ground). Drought survival time was correlated most strongly with seed mass, root construction and allocation traits, and phylogeny (grasses vs. forbs).

Synthesis.Our results show that seed and seedling trait variation among annual species is substantial, and that a few attributes could capture major dimensions of ecological strategies during emergence. With seedling survival times ranging twofold among annuals (from 7.5 to 14.5 days), these strategies could mitigate recruitment responses to more frequent or longer dry spells. Multivariate trait and plasticity strategies should be further explored in studies designed to assess trait‐fitness linkages during recruitment.

A freePlain Language Summarycan be found within the Supporting Information of this article.