Evolutionary radiations of woody taxa within arid environments were made possible by multiple trait innovations including deep roots and embolism-resistant xylem, but little is known about how... Show moreEvolutionary radiations of woody taxa within arid environments were made possible by multiple trait innovations including deep roots and embolism-resistant xylem, but little is known about how these traits have coevolved across the phylogeny of woody plants or how they jointly influence the distribution of species. We synthesized global trait and vegetation plot datasets to examine how rooting depth and xylem vulnerability across 188 woody plant species interact with aridity, precipitation seasonality, and water table depth to influence species occurrence probabilities across all biomes. Xylem resistance to embolism and rooting depth are independent woody plant traits that do not exhibit an interspecific trade-off. Resistant xylem and deep roots increase occurrence probabilities in arid, seasonal climates over deep water tables. Resistant xylem and shallow roots increase occurrence probabilities in arid, nonseasonal climates over deep water tables. Vulnerable xylem and deep roots increase occurrence probabilities in arid, nonseasonal climates over shallow water tables. Lastly, vulnerable xylem and shallow roots increase occurrence probabilities in humid climates. Each combination of trait values optimizes occurrence probabilities in unique environmental conditions. Responses of deeply rooted vegetation may be buffered if evaporative demand changes faster than water table depth under climate change. Show less
As Earth's climate has varied strongly through geological time, studying the impacts of past climate change on biodiversity helps to understand the risks from future climate change. However, it... Show moreAs Earth's climate has varied strongly through geological time, studying the impacts of past climate change on biodiversity helps to understand the risks from future climate change. However, it remains unclear how paleoclimate shapes spatial variation in biodiversity. Here, we assessed the influence of Quaternary climate change on spatial dissimilarity in taxonomic, phylogenetic, and functional composition among neighboring 200-kilometer cells (beta-diversity) for angiosperm trees worldwide. We found that larger glacial-interglacial temperature change was strongly associated with lower spatial turnover (species replacements) and higher nestedness (richness changes) components of beta-diversity across all three biodiversity facets. Moreover, phylogenetic and functional turnover was lower and nestedness higher than random expectations based on taxonomic beta-diversity in regions that experienced large temperature change, reflecting phylogenetically and functionally selective processes in species replacement, extinction, and colonization during glacial-interglacial oscillations. Our results suggest that future human-driven climate change could cause local homogenization and reduction in taxonomic, phylogenetic, and functional diversity of angiosperm trees worldwide. Show less
Maynard, D.S.; Bialic-Murphy, L.; Zohner, C.M.; Averill, C.; Hoogen, J. van den; Ma, H.; ... ; Crowther, T.W. 2022
Due to massive energetic investments in woody support structures, trees are subject to unique physiological, mechanical, and ecological pressures not experienced by herbaceous plants. Despite a... Show moreDue to massive energetic investments in woody support structures, trees are subject to unique physiological, mechanical, and ecological pressures not experienced by herbaceous plants. Despite a wealth of studies exploring trait relationships across the entire plant kingdom, the dominant traits underpinning these unique aspects of tree form and function remain unclear. Here, by considering 18 functional traits, encompassing leaf, seed, bark, wood, crown, and root characteristics, we quantify the multidimensional relationships in tree trait expression. We find that nearly half of trait variation is captured by two axes: one reflecting leaf economics, the other reflecting tree size and competition for light. Yet these orthogonal axes reveal strong environmental convergence, exhibiting correlated responses to temperature, moisture, and elevation. By subsequently exploring multidimensional trait relationships, we show that the full dimensionality of trait space is captured by eight distinct clusters, each reflecting a unique aspect of tree form and function. Collectively, this work identifies a core set of traits needed to quantify global patterns in functional biodiversity, and it contributes to our fundamental understanding of the functioning of forests worldwide. Show less
Plant functional traits can predict community assembly and ecosystem functioning and are thus widely used in global models of vegetation dynamics and land–climate feedbacks. Still, we lack a global... Show morePlant functional traits can predict community assembly and ecosystem functioning and are thus widely used in global models of vegetation dynamics and land–climate feedbacks. Still, we lack a global understanding of how land and climate affect plant traits. A previous global analysis of six traits observed two main axes of variation: (1) size variation at the organ and plant level and (2) leaf economics balancing leaf persistence against plant growth potential. The orthogonality of these two axes suggests they are differently influenced by environmental drivers. We find that these axes persist in a global dataset of 17 traits across more than 20,000 species. We find a dominant joint effect of climate and soil on trait variation. Additional independent climate effects are also observed across most traits, whereas independent soil effects are almost exclusively observed for economics traits. Variation in size traits correlates well with a latitudinal gradient related to water or energy limitation. In contrast, varia- tion in economics traits is better explained by interactions of climate with soil fertility. These findings have the potential to improve our understanding of biodiversity patterns and our predictions of climate change impacts on biogeochemical cycles. Show less
The leaf economic spectrum is a widely studied axis of plant trait variability that defines a trade-off between leaf longevity and productivity. While this has been investigated at the global scale... Show moreThe leaf economic spectrum is a widely studied axis of plant trait variability that defines a trade-off between leaf longevity and productivity. While this has been investigated at the global scale, where it is robust, and at local scales, where deviations from it are common, it has received less attention at the intermediate scale of plant functional types (PFTs). We investigated whether global leaf economic relationships are also present within the scale of plant functional types (PFTs) commonly used by Earth System models, and the extent to which this global-PFT hierarchy can be used to constrain trait estimates. We developed a hierarchical multivariate Bayesian model that assumes separate means and covariance structures within and across PFTs and fit this model to seven leaf traits from the TRY database related to leaf longevity, morphology, biochemistry, and photosynthetic metabolism. Although patterns of trait covariation were generally consistent with the leaf economic spectrum, we found three approximate tiers to this consistency. Relationships among morphological and biochemical traits (specific leaf area [SLA], N, P) were the most robust within and across PFTs, suggesting that covariation in these traits is driven by universal leaf construction trade-offs and stoichiometry. Relationships among metabolic traits (dark respiration [R-d], maximum RuBisCo carboxylation rate [V-c,V-max], maximum electron transport rate [J(max)]) were slightly less consistent, reflecting in part their much sparser sampling (especially for high-latitude PFTs), but also pointing to more flexible plasticity in plant metabolistm. Finally, relationships involving leaf lifespan were the least consistent, indicating that leaf economic relationships related to leaf lifespan are dominated by across-PFT differences and that within-PFT variation in leaf lifespan is more complex and idiosyncratic. Across all traits, this covariance was an important source of information, as evidenced by the improved imputation accuracy and reduced predictive uncertainty in multivariate models compared to univariate models. Ultimately, our study reaffirms the value of studying not just individual traits but the multivariate trait space and the utility of hierarchical modeling for studying the scale dependence of trait relationships. Show less
Aim Plant trait databases often contain traits that are correlated, but for whom direct (undirected statistical dependency) and indirect (mediated by other traits) connections may be confounded.... Show moreAim Plant trait databases often contain traits that are correlated, but for whom direct (undirected statistical dependency) and indirect (mediated by other traits) connections may be confounded. The confounding of correlation and connection hinders our understanding of plant strategies, and how these vary among growth forms and climate zones. We identified the direct and indirect connections across plant traits relevant to competition, resource acquisition and reproductive strategies using a global database and explored whether connections within and between traits from different tissue types vary across climates and growth forms. Location Global. Major taxa studied Plants. Time period Present. Methods We used probabilistic graphical models and a database of 10 plant traits (leaf area, specific leaf area, mass- and area-based leaf nitrogen and phosphorous content, leaf life span, plant height, stem specific density and seed mass) with 16,281 records to describe direct and indirect connections across woody and non-woody plants across tropical, temperate, arid, cold and polar regions. Results Trait networks based on direct connections are sparser than those based on correlations. Land plants had high connectivity across traits within and between tissue types; leaf life span and stem specific density shared direct connections with all other traits. For both growth forms, two groups of traits form modules of more highly connected traits; one related to resource acquisition, the other to plant architecture and reproduction. Woody species had higher trait network modularity in polar compared to temperate and tropical climates, while non-woody species did not show significant differences in modularity across climate regions. Main conclusions Plant traits are highly connected both within and across tissue types, yet traits segregate into persistent modules of traits. Variation in the modularity of trait networks suggests that trait connectivity is shaped by prevailing environmental conditions and demonstrates that plants of different growth forms use alternative strategies to cope with local conditions. Show less
Mencuccini, M.; Rosa, T.; Rowland, L.; Choat, B.; Cornelissen, H.; Jansen, S.; ... ; Martinez-Vilalta, J. 2019
Biomass and area ratios between leaves, stems and roots regulate many physiological and ecological processes. The Huber value H-v (sapwood area/leaf area ratio) is central to plant water balance... Show moreBiomass and area ratios between leaves, stems and roots regulate many physiological and ecological processes. The Huber value H-v (sapwood area/leaf area ratio) is central to plant water balance and drought responses. However, its coordination with key plant functional traits is poorly understood, and prevents developing trait-based prediction models. Based on theoretical arguments, we hypothesise that global patterns in H-v of terminal woody branches can be predicted from variables related to plant trait spectra, that is plant hydraulics and size and leaf economics. Using a global compilation of 1135 species-averaged H-v, we show that H-v varies over three orders of magnitude. Higher H-v are seen in short small-leaved low-specific leaf area (SLA) shrubs with low K-s in arid relative to tall large-leaved high-SLA trees with high K-s in moist environments. All traits depend on climate but climatic correlations are stronger for explanatory traits than H-v. Negative isometry is found between H-v and K-s, suggesting a compensation to maintain hydraulic supply to leaves across species. This work identifies the major global drivers of branch sapwood/leaf area ratios. Our approach based on widely available traits facilitates the development of accurate models of above-ground biomass allocation and helps predict vegetation responses to drought. Show less
This paper introduces a modular processing chain to derive global high-resolution maps of leaf traits. In particular, we present global maps at 500 m resolution of specific leaf area, leaf dry... Show moreThis paper introduces a modular processing chain to derive global high-resolution maps of leaf traits. In particular, we present global maps at 500 m resolution of specific leaf area, leaf dry matter content, leaf nitrogen and phosphorus content per dry mass, and leaf nitrogen/phosphorus ratio. The processing chain exploits machine learning techniques along with optical remote sensing data (MODIS/Landsat) and climate data for gap filling and up-scaling of in-situ measured leaf traits. The chain first uses random forests regression with surrogates to fill gaps in the database (> 45% of missing entries) and maximizes the global representativeness of the trait dataset. Plant species are then aggregated to Plant Functional Types (PFTs). Next, the spatial abundance of PFTs at MODIS resolution (500 m) is calculated using Landsat data (30 m). Based on these PFT abundances, representative trait values are calculated for MODIS pixels with nearby trait data. Finally, different regression algorithms are applied to globally predict trait estimates from these MODIS pixels using remote sensing and climate data. The methods were compared in terms of precision, robustness and efficiency. The best model (random forests regression) shows good precision (normalized RMSE≤ 20%) and goodness of fit (averaged Pearson's correlation R = 0.78) in any considered trait. Along with the estimated global maps of leaf traits, we provide associated uncertainty estimates derived from the regression models. The process chain is modular, and can easily accommodate new traits, data streams (traits databases and remote sensing data), and methods. The machine learning techniques applied allow attribution of information gain to data input and thus provide the opportunity to understand trait-environment relationships at the plant and ecosystem scales. The new data products – the gap-filled trait matrix, a global map of PFT abundance per MODIS gridcells and the high-resolution global leaf trait maps – are complementary to existing large-scale observations of the land surface and we therefore anticipate substantial contributions to advances in quantifying, understanding and prediction of the Earth system. Show less
Aim The influence of soil properties on photosynthetic traits in higher plants is poorly quantified in comparison with that of climate.We address this situation by quantifying the unique and joint... Show moreAim The influence of soil properties on photosynthetic traits in higher plants is poorly quantified in comparison with that of climate.We address this situation by quantifying the unique and joint contributions to global leaf-trait variation from soils and climate. Location Terrestrial ecosystems world-wide. Methods Using a trait dataset comprising 1509 species from 288 sites, with climate and soil data derived from global datasets, we quantified the effects of 20 soil and 26 climate variables on light-saturated photosynthetic rate (Aarea), stomatal conductance (gs), leaf nitrogen and phosphorus (Narea and Parea) and specific leaf area (SLA) using mixed regression models and multivariate analyses. Results Soil variables were stronger predictors of leaf traits than climatic variables, except for SLA. On average, Narea, Parea and Aarea increased and SLA decreased with increasing soil pH and with increasing site aridity. gs declined and Parea increased with soil available P (Pavail). Narea was unrelated to total soil N. Joint effects of soil and climate dominated over their unique effects on Narea and Parea, while unique effects of soils dominated for Aarea and gs. Path analysis indicated that variation in Aarea reflected the combined independent influences of Narea and gs, the former promoted by high pH and aridity and the latter by low Pavail. Main conclusions Three environmental variables were key for explaining variation in leaf traits: soil pH and Pavail, and the climatic moisture index (the ratio ofprecipitation to potential evapotranspiration). Although the reliability of global soil datasets lags behind that of climate datasets, our results nonetheless provide compelling evidence that both can be jointly used in broad-scale analyses, and that effects uniquely attributable to soil properties are important determinants of leaf photosynthetic traits and rates. A significant future challenge is to better disentangle the covarying physiological, ecological Show less
Plant traits – the morphological, anatomical, physiological, biochemical and phenological characteristics of plants and their organs – determine how primary producers respond to environmental... Show morePlant traits – the morphological, anatomical, physiological, biochemical and phenological characteristics of plants and their organs – determine how primary producers respond to environmental factors, affect other trophic levels, influence ecosystem processes and services and provide a link from species richness to ecosystem functional diversity. Trait data thus represent the raw material for a wide range of research from evolutionary biology, community and functional ecology to biogeography. Here we present the global database initiative named TRY, which has united a wide range of the plant trait research community worldwide and gained an unprecedented buy-in of trait data: so far 93 trait databases have been contributed. The data repository currently contains almost three million trait entries for 69 000 out of the world’s 300 000 plant species, with a focus on 52 groups of traits characterizing the vegetative and regeneration stages of the plant life cycle, including growth, dispersal, establishment and persistence. A first data analysis shows that most plant traits are approximately log-normally distributed, with widely differing ranges of variation across traits. Most trait variation is between species (interspecific), but significant intraspecific variation is also documented, up to 40% of the overall variation. Plant functional types (PFTs), as commonly used in vegetation models, capture a substantial fraction of the observed variation – but for several traits most variation occurs within PFTs, up to 75% of the overall variation. In the context of vegetation models these traits would better be represented by state variables rather than fixed parameter values. The improved availability of plant trait data in the unified global database is expected to support a paradigm shift from species to trait-based ecology, offer new opportunities for synthetic plant trait research and enable a more realistic and empirically grounded representation of terrestrial vegetation in Earth system models. Show less