Toward a coordinated understanding of hydro-biogeochemical root functions in tropical forests for application in vegetation models.

Tropical forest root characteristics and resource acquisition strategies are underrepresented in vegetation and global models, hampering the prediction of forest-climate feedbacks for these carbon-rich ecosystems. Lowland tropical forests often have globally unique combinations of high taxonomic and functional biodiversity, rainfall seasonality, and strongly weathered infertile soils, giving rise to distinct patterns in root traits and functions compared with higher latitude ecosystems. We provide a roadmap for integrating recent advances in our understanding of tropical forest belowground function into vegetation models, focusing on water and nutrient acquisition. We offer comparisons of recent advances in empirical and model understanding of root characteristics that represent important functional processes in tropical forests. We focus on: (1) fine-root strategies for soil resource exploration, (2) coupling and trade-offs in fine-root water vs nutrient acquisition, and (3) aboveground-belowground linkages in plant resource acquisition and use. We suggest avenues for representing these extremely diverse plant communities in computationally manageable and ecologically meaningful groups in models for linked aboveground-belowground hydro-nutrient functions. Tropical forests are undergoing warming, shifting rainfall regimes, and exacerbation of soil nutrient scarcity caused by elevated atmospheric CO2. The accurate model representation of tropical forest functions is crucial for understanding the interactions of this biome with the climate.

Las características de las raíces de los bosques tropicales y las estrategias de adquisición de recursos están subrepresentadas en modelos de vegetación, lo que dificulta la predicción del efecto de cambio de clima para estos ecosistemas ricos en carbono. Los bosques tropicales a menudo tienen combinaciones únicas a nivel mundial de alta biodiversidad taxonómica y funcional, estacionalidad de precipitación, y suelos infértiles, dando lugar a patrones distintos en los rasgos y funciones de las raíces en comparación con los ecosistemas de latitudes más altas. Integramos los avances recientes en nuestra comprensión de la función subterránea de los bosques tropicales en modelos de vegetación, centrándonos en la adquisición de agua y nutrientes. Ofrecemos comparaciones de avances recientes en la comprensión empírica y de modelos de las características de las raíces que representan procesos funcionales importantes en los bosques tropicales. Nos centramos en: (1) estrategias de raíces finas para adquisición de recursos del suelo, (2) acoplamiento y compensaciones entre adquisición del agua y de nutrientes, y (3) vínculos entre funciones sobre tierra y debajo del superficie en bosques tropicales. Sugerimos vías para representar estas comunidades de plantas extremadamente diversas en grupos computacionalmente manejables y ecológicamente significativos en modelos. Los bosques tropicales se están calentando, tienen cambios en los regímenes de lluvias, y tienen una exacerbación de la escasez de nutrientes del suelo causada por el elevado CO2 atmosférico. La representación precisa de las funciones de los bosques tropicales en modelos es crucial para comprender las interacciones de este bioma con el clima.

Exploring the role of biotic factors in regulating the spatial variability in land surface phenology across four temperate forest sites.

Land surface phenology (LSP), the characterization of plant phenology with satellite data, is essential for understanding the effects of climate change on ecosystem functions. Considerable LSP variation is observed within local landscapes, and the role of biotic factors in regulating such variation remains underexplored. In this study, we selected four National Ecological Observatory Network terrestrial sites with minor topographic relief to investigate how biotic factors regulate intra-site LSP variability. We utilized plant functional type (PFT) maps, functional traits, and LSP data to assess the explanatory power of biotic factors for the start and end of season (SOS and EOS) variability. Our results indicate that PFTs alone explain only 0.8-23.4% of intra-site SOS and EOS variation, whereas including functional traits significantly improves explanatory power, with cross-validation correlations ranging from 0.50 to 0.85. While functional traits exhibited diverse effects on SOS and EOS across different sites, traits related to competitive ability and productivity were important for explaining both SOS and EOS variation at these sites. These findings reveal that plants exhibit diverse phenological responses to comparable environmental conditions, and functional traits significantly contribute to intra-site LSP variability, highlighting the importance of intrinsic biotic properties in regulating plant phenology.

Soil fungal and bacterial communities reflect differently tundra vegetation state transitions and soil physico-chemical properties.

Strong disturbances may induce ecosystem transitions into new alternative states that sustain through plant-soil interactions, such as the transition of dwarf shrub-dominated into graminoid-dominated vegetation by herbivory in tundra. Little evidence exists on soil microbial communities in alternative states, and along the slow process of ecosystem return into the predisturbance state. We analysed vegetation, soil microbial communities and activities as well as soil physico-chemical properties in historical reindeer enclosures in northernmost Finland in the following plot types: control heaths in the surrounding tundra; graminoid-dominated; 'shifting'; and recovered dwarf shrub-dominated vegetation inside enclosures. Soil fungal communities followed changes in vegetation, whereas bacterial communities were more affected by soil physico-chemical properties. Graminoid plots were characterized by moulds, pathotrophs and dark septate endophytes. Ericoid mycorrhizal and saprotrophic fungi were typical for control and recovered plots. Soil microbial communities inside the enclosures showed historical contingency, as their spatial variation was high in recovered plots despite the vegetation being more homogeneous. Self-maintaining feedback loops between plant functional types, soil microbial communities, and carbon and nutrient mineralization act effectively to stabilize alternative vegetation states, but once predisturbance vegetation reestablishes itself, soil microbial communities and physico-chemical properties return back towards their predisturbance state.

Thermography captures the differential sensitivity of dryland functional types to changes in rainfall event timing and magnitude.

Drylands of the southwestern United States are rapidly warming, and rainfall is becoming less frequent and more intense, with major yet poorly understood implications for ecosystem structure and function. Thermography-based estimates of plant temperature can be integrated with air temperature to infer changes in plant physiology and response to climate change. However, very few studies have evaluated plant temperature dynamics at high spatiotemporal resolution in rainfall pulse-driven dryland ecosystems. We address this gap by incorporating high-frequency thermal imaging into a field-based precipitation manipulation experiment in a semi-arid grassland to investigate the impacts of rainfall temporal repackaging. All other factors held constant, we found that fewer/larger precipitation events led to cooler plant temperatures (1.4°C) compared to that of many/smaller precipitation events. Perennials, in particular, were 2.5°C cooler than annuals under the fewest/largest treatment. We show these patterns were driven by: increased and consistent soil moisture availability in the deeper soil layers in the fewest/largest treatment; and deeper roots of perennials providing access to deeper plant available water. Our findings highlight the potential for high spatiotemporal resolution thermography to quantify the differential sensitivity of plant functional groups to soil water availability. Detecting these sensitivities is vital to understanding the ecohydrological implications of hydroclimate change.

Global net biome CO2 exchange predicted comparably well using parameter-environment relationships and plant functional types.

Accurate estimation and forecasts of net biome CO2 exchange (NBE) are vital for understanding the role of terrestrial ecosystems in a changing climate. Prior efforts to improve NBE predictions have predominantly focused on increasing models' structural realism (and thus complexity), but parametric error and uncertainty are also key determinants of model skill. Here, we investigate how different parameterization assumptions propagate into NBE prediction errors across the globe, pitting the traditional plant functional type (PFT)-based approach against a novel top-down, machine learning-based "environmental filtering" (EF) approach. To do so, we simulate these contrasting methods for parameter assignment within a flexible model-data fusion framework of the terrestrial carbon cycle (CARDAMOM) at a global scale. In the PFT-based approach, model parameters from a small number of select locations are applied uniformly within regions sharing similar land cover characteristics. In the EF-based approach, a pixel's parameters are predicted based on underlying relationships with climate, soil, and canopy properties. To isolate the role of parametric from structural uncertainty in our analysis, we benchmark the resulting PFT-based and EF-based NBE predictions with estimates from CARDAMOM's Bayesian optimization approach (whereby "true" parameters consistent with a suite of data constraints are retrieved on a pixel-by-pixel basis). When considering the mean absolute error of NBE predictions across time, we find that the EF-based approach matches or outperforms the PFT-based approach at 55% of pixels-a narrow majority. However, NBE estimates from the EF-based approach are susceptible to compensation between errors in component flux predictions and predicted parameters can align poorly with the assumed "true" values. Overall, though, the EF-based approach is comparable to conventional approaches and merits further investigation to better understand and resolve these limitations. This work provides insight into the relationship between terrestrial biosphere model performance and parametric uncertainty, informing efforts to improve model parameterization via PFT-free and trait-based approaches.

Water level drawdown makes boreal peatland vegetation more responsive to weather conditions.

Climate warming and projected increase in summer droughts puts northern peatlands under pressure by subjecting them to a combination of gradual drying and extreme weather events. The combined effect of those on peatland functions is poorly known. Here, we studied the impact of long-term water level drawdown (WLD) and contrasting weather conditions on leaf phenology and biomass production of ground level vegetation in boreal peatlands. Data were collected during two contrasting growing seasons from a WLD experiment including a rich and a poor fen and an ombrotrophic bog. Results showed that WLD had a strong effect on both leaf area development and biomass production, and these responses differed between peatland types. In the poor fen and the bog, WLD increased plant growth, while in the rich fen, WLD reduced the growth of ground level vegetation. Plant groups differed in their response, as WLD reduced the growth of graminoids, while shrubs and tree seedlings benefited from it. In addition, the vegetation adjusted to the lower WTs, was more responsive to short-term climatic variations. The warmer summer resulted in a greater maximum and earlier peaking of leaf area index, and greater biomass production by vascular plants and Sphagnum mosses at WLD sites. In particular, graminoids benefitted from the warmer conditions. The change towards greater production in the WLD sites in general and during the warmer weather in particular, was related to the observed transition in plant functional type composition towards arboreal vegetation.

Consistent diurnal pattern of leaf respiration in the light among contrasting species and climates.

Leaf daytime respiration (leaf respiration in the light, RL ) is often assumed to constitute a fixed fraction of leaf dark respiration (RD ) (i.e. a fixed light inhibition of respiration (RD )) and vary diurnally due to temperature fluctuations. These assumptions were tested by measuring RL , RD and the light inhibition of RD in the field at a constant temperature using the Kok method. Measurements were conducted diurnally on 21 different species: 13 deciduous, four evergreen and four herbaceous from humid continental and humid subtropical climates. RL and RD showed significant diurnal variations and the diurnal pattern differed in trajectory and magnitude between climates, but not between plant functional types (PFTs). The light inhibition of RD varied diurnally and differed between climates and in trajectory between PFTs. The results highlight the entrainment of leaf daytime respiration to the diurnal cycle and that time of day should be accounted for in studies seeking to examine the environmental and biological drivers of leaf daytime respiration.

Cross-scale regulation of seasonal microclimate by vegetation and snow in the Arctic tundra.

Climate warming is inducing widespread vegetation changes in Arctic tundra ecosystems, with the potential to alter carbon and nutrient dynamics between vegetation and soils. Yet, we lack a detailed understanding of how variation in vegetation and topography influences fine-scale temperatures ("microclimate") that mediate these dynamics, and at what resolution vegetation needs to be sampled to capture these effects. We monitored microclimate at 90 plots across a tundra landscape in western Greenland. Our stratified random study design covered gradients of topography and vegetation, while nested plots (0.8-100 m2 ) enabled comparison across different sampling resolutions. We used Bayesian mixed-effect models to quantify the direct influence of plot-level topography, moisture and vegetation on soil, near-surface and canopy-level temperatures (-6, 2, and 15 cm). During the growing season, colder soils were predicted by shrub cover (-0.24°C per 10% increase), bryophyte cover (-0.35°C per 10% increase), and vegetation height (-0.17°C per 1 cm increase). The same three factors also predicted the magnitude of differences between soil and above-ground temperatures, indicating warmer soils at low cover/height, but colder soils under closed/taller canopies. These findings were consistent across plot sizes, suggesting that spatial predictions of microclimate may be possible at the operational scales of satellite products. During winter, snow cover (+0.75°C per 10 snow-covered days) was the key predictor of soil microclimate. Topography and moisture explained little variation in the measured temperatures. Our results not only underline the close connection of vegetation and snow with microclimate in the Arctic tundra but also point to the need for more studies disentangling their complex interplay across tundra environments and seasons. Future shifts in vegetation cover and height will likely mediate the impact of atmospheric warming on the tundra soil environment, with potential implications for below-ground organisms and ecosystem functioning.

Representing plant diversity in land models: An evolutionary approach to make "Functional Types" more functional.

Plants are critical mediators of terrestrial mass and energy fluxes, and their structural and functional traits have profound impacts on local and global climate, biogeochemistry, biodiversity, and hydrology. Yet, Earth System Models (ESMs), our most powerful tools for predicting the effects of humans on the coupled biosphere-atmosphere system, simplify the incredible diversity of land plants into a handful of coarse categories of "Plant Functional Types" (PFTs) that often fail to capture ecological dynamics such as biome distributions. The inclusion of more realistic functional diversity is a recognized goal for ESMs, yet there is currently no consistent, widely accepted way to add diversity to models, that is, to determine what new PFTs to add and with what data to constrain their parameters. We review approaches to representing plant diversity in ESMs and draw on recent ecological and evolutionary findings to present an evolution-based functional type approach for further disaggregating functional diversity. Specifically, the prevalence of niche conservatism, or the tendency of closely related taxa to retain similar ecological and functional attributes through evolutionary time, reveals that evolutionary relatedness is a powerful framework for summarizing functional similarities and differences among plant types. We advocate that Plant Functional Types based on dominant evolutionary lineages ("Lineage Functional Types") will provide an ecologically defensible, tractable, and scalable framework for representing plant diversity in next-generation ESMs, with the potential to improve parameterization, process representation, and model benchmarking. We highlight how the importance of evolutionary history for plant function can unify the work of disparate fields to improve predictive modeling of the Earth system.

Cover crop functional types differentially alter the content and composition of soil organic carbon in particulate and mineral-associated fractions.

Cover crops (CCs) can increase soil organic carbon (SOC) sequestration by providing additional OC residues, recruiting beneficial soil microbiota, and improving soil aggregation and structure. The various CC species that belong to distinct plant functional types (PFTs) may differentially impact SOC formation and stabilization. Biogeochemical theory suggests that selection of PFTs with distinct litter quality (C:N ratio) should influence the pathways and magnitude of SOC sequestration. Yet, we lack knowledge on the effect of CCs from different PFTs on the quantity and composition of physiochemical pools of SOC. We sampled soils under monocultures of three CC PFTs (legume [crimson clover]; grass [triticale]; and brassica [canola]) and a mixture of these three species, from a long-term CC experiment in Pennsylvania, USA. We measured C content in bulk soil and C content and composition in contrasting physical fractions: particulate organic matter, POM; and mineral-associated organic matter, MAOM. The bulk SOC content was higher in all CC treatments compared to the fallow. Compared to the legume, monocultures of grass and brassica with lower litter quality (wider C:N) had higher proportion of plant-derived C in POM, indicating selective preservation of complex structural plant compounds. In contrast, soils under legumes had greater accumulation of microbial-derived C in MAOM. Our results for the first time, revealed that the mixture contributed to a higher concentration of plant-derived compounds in POM relative to the legume, and a greater accumulation of microbial-derived C in MAOM compared to monocultures of grass and brassica. Mixtures with all three PFTs can thus increase the short- and long-term SOC persistence balancing the contrasting effects on the chemistries in POM and MAOM imposed by monoculture CC PFTs. Thus, despite different cumulative C inputs in CC treatments from different PFTs, the total SOC stocks did not vary between CC PFTs, rather PFTs impacted whether C accumulated in POM or MAOM fractions. This highlights that CCs of different PFTs may shift the dominant SOC formation pathways (POM vs. MAOM), subsequently impacting short- and long-term SOC stabilization and stocks. Our work provides a strong applied field test of biogeochemical theory linking litter quality to pathways of C accrual in soil.

Drivers of phenological changes in southern Europe.

The life cycle of plants is largely determined by climate, which renders phenological responses to climate change a highly suitable bioindicator of climate change. Yet, it remains unclear, which are the key drivers of phenological patterns at certain life stages. Furthermore, the varying responses of species belonging to different plant functional types are not fully understood. In this study, the role of temperature and precipitation as environmental drivers of phenological changes in southern Europe is assessed. The trends of the phenophases leaf unfolding, flowering, fruiting, and senescence are quantified, and the corresponding main environmental drivers are identified. A clear trend towards an earlier onset of leaf unfolding, flowering, and fruiting is detected, while there is no clear pattern for senescence. In general, the advancement of leaf unfolding, flowering and fruiting is smaller for deciduous broadleaf trees in comparison to deciduous shrubs and crops. Many broadleaf trees are photoperiod-sensitive; therefore, their comparatively small phenological advancements are likely the effect of photoperiod counterbalancing the impact of increasing temperatures. While temperature is identified as the main driver of phenological changes, precipitation also plays a crucial role in determining the onset of leaf unfolding and flowering. Phenological phases advance under dry conditions, which can be linked to the lack of transpirational cooling leading to rising temperatures, which subsequently accelerate plant growth.

Fire does not transform shrublands of Echinospartum horridum (Vahl) Rothm. into grasslands in the Pyrenees: Development of community structure and nutritive value after single prescribed burns.

Prescribed fire has been extensively used in recent years to control woody encroachment into mountain and other grassland-dominated landscapes. In the Aragon Pyrenees, prescribed burns have been mainly used to remove the native thorny shrub Echinospartum horridum (Vahl) Rothm., whose populations are spreading to the detriment of grasslands. To study the effectiveness of the burning of E. horridum to preserve grasslands for livestock grazing, the vegetation of six sites burned 0.5, 2, 3, 6, 15 and 35 years ago was sampled and compared with that of nearby unburned shrubland (control) and grassland (reference). In addition, the nutritional quality of E. horridum was examined and compared to that of the reference grassland to evaluate to what extent shrub growth can be controlled by herbivores after burning. Initial shrub cover recovered as early as 15 years after fire, with E. horridum being dominant. Plant diversity was greatest at intermediate number of years after fire. Initial floristic composition and life-form spectrum were restored 15-35 years after burning. Echinospartum horridum exhibited early lignification that restricts its availability as a palatable forage for the first two years after burning and makes it unlikely to be consumed thereafter, highlighting the difficulty in controlling the expansion of this species by livestock herbivory. The analysis of the nutrient levels suggested an increased shortage of limiting elements, such as phosphorus or sulfur, in the mid-term after burning due to substantial nutrient losses and exports during and after the burn. Our results question the suitability and sustainability of a single prescribed burn as management tool alone to control the expansion of E. horridum and call for caution in its application for fighting shrub encroachment in the Central Pyrenees.

Global patterns of biomass allocation in woody species with different tolerances of shade and drought: evidence for multiple strategies.

The optimal partitioning theory predicts that plants of a given species acclimate to different environments by allocating a larger proportion of biomass to the organs acquiring the most limiting resource. Are similar patterns found across species adapted to environments with contrasting levels of abiotic stress? We tested the optimal partitioning theory by analysing how fractional biomass allocation to leaves, stems and roots differed between woody species with different tolerances of shade and drought in plants of different age and size (seedlings to mature trees) using a global dataset including 604 species. No overarching biomass allocation patterns at different tolerance values across species were found. Biomass allocation varied among functional types as a result of phenological (deciduous vs evergreen broad-leaved species) and broad phylogenetical (angiosperms vs gymnosperms) differences. Furthermore, the direction of biomass allocation responses between tolerant and intolerant species was often opposite to that predicted by the optimal partitioning theory. We conclude that plant functional type is the major determinant of biomass allocation in woody species. We propose that interactions between plant functional type, ontogeny and species-specific stress tolerance adaptations allow woody species with different shade and drought tolerances to display multiple biomass partitioning strategies.

Wetland Heterogeneity Determines Methane Emissions: A Pan-Arctic Synthesis.

Methane (CH4) emissions from pan-Arctic wetlands provide a potential positive feedback to global warming. However, the differences in CH4 emissions across wetland types in these regions have not been well understood. We synthesized approximately 9000 static chamber CH4 measurements during the growing season from 83 sites across pan-Arctic regions. We highlighted spatial variations of CH4 emissions corresponding to environmental heterogeneity across wetland types. CH4 emission is the highest in fens, followed by marshes, bogs, and the lowest in swamps. This gradient is controlled by the water table, soil temperature, and dominant plant functional types and their interactions. The water table position for maximum CH4 emission is below, close to, and above the ground surface in bogs, marshes/fens, and swamps, respectively. The temperature sensitivity (Q10) of CH4 emissions varied among different wetland types, ranging from the lowest in swamps to the highest in fens. The interactive impact of temperature and the water table positions on CH4 emissions are regulated with dominant plant functional types. CH4 emissions from wetlands dominated by vascular plants rely more on species composition than that dominated by non-vascular plants. Wetlands with greater abundance of graminoids (e.g., fens) have higher CH4 emissions than tree-dominated wetlands (e.g., swamps). This synthesis emphasizes the role of wetland heterogeneity in determining the strength of CH4 emissions.

Divergent drivers of leaf trait variation within species, among species, and among functional groups.

Understanding variation in leaf functional traits-including rates of photosynthesis and respiration and concentrations of nitrogen and phosphorus-is a fundamental challenge in plant ecophysiology. When expressed per unit leaf area, these traits typically increase with leaf mass per area (LMA) within species but are roughly independent of LMA across the global flora. LMA is determined by mass components with different biological functions, including photosynthetic mass that largely determines metabolic rates and contains most nitrogen and phosphorus, and structural mass that affects toughness and leaf lifespan (LL). A possible explanation for the contrasting trait relationships is that most LMA variation within species is associated with variation in photosynthetic mass, whereas most LMA variation across the global flora is associated with variation in structural mass. This hypothesis leads to the predictions that (i) gas exchange rates and nutrient concentrations per unit leaf area should increase strongly with LMA across species assemblages with low LL variance but should increase weakly with LMA across species assemblages with high LL variance and that (ii) controlling for LL variation should increase the strength of the above LMA relationships. We present analyses of intra- and interspecific trait variation from three tropical forest sites and interspecific analyses within functional groups in a global dataset that are consistent with the above predictions. Our analysis suggests that the qualitatively different trait relationships exhibited by different leaf assemblages can be understood by considering the degree to which photosynthetic and structural mass components contribute to LMA variation in a given assemblage.

Lineage-based functional types: characterising functional diversity to enhance the representation of ecological behaviour in Land Surface Models.

Process-based vegetation models attempt to represent the wide range of trait variation in biomes by grouping ecologically similar species into plant functional types (PFTs). This approach has been successful in representing many aspects of plant physiology and biophysics but struggles to capture biogeographic history and ecological dynamics that determine biome boundaries and plant distributions. Grass-dominated ecosystems are broadly distributed across all vegetated continents and harbour large functional diversity, yet most Land Surface Models (LSMs) summarise grasses into two generic PFTs based primarily on differences between temperate C3 grasses and (sub)tropical C4 grasses. Incorporation of species-level trait variation is an active area of research to enhance the ecological realism of PFTs, which form the basis for vegetation processes and dynamics in LSMs. Using reported measurements, we developed grass functional trait values (physiological, structural, biochemical, anatomical, phenological, and disturbance-related) of dominant lineages to improve LSM representations. Our method is fundamentally different from previous efforts, as it uses phylogenetic relatedness to create lineage-based functional types (LFTs), situated between species-level trait data and PFT-level abstractions, thus providing a realistic representation of functional diversity and opening the door to the development of new vegetation models.

Biome: evolution of a crucial ecological and biogeographical concept.

A biome is a key community ecological and biogeographical concept and, as such, has profited from the overall progress of community ecology, punctuated by two major innovations: shifting the focus from pure pattern description to understanding functionality, and changing the approach from observational to explanatory and, most importantly, from descriptive to predictive. The functional focus enabled development of mechanistic and function-focused predictive and retrodictive modelling; it also shaped the current understanding of the concept of a biome as a dynamic biological entity having many aspects, with deep roots in the evolutionary past, and which is undergoing change. The evolution of the biome concept was punctuated by three synthetic steps: the first synthesis formulated a solid body of theory explaining the ecological and biogeographical meaning of zonality and collated our knowledge on drivers of vegetation patterns at large spatial scales; the second translated this knowledge into effective mechanistic modelling tools, developing further the link between ecosystem functionality and biogeography; and the third (still in progress) is seeking common ground between large-scale ecological and biogeographic phenomena, using macroecology and macroevolutionary research tools.

Traditional plant functional groups explain variation in economic but not size-related traits across the tundra biome.

AIM: Plant functional groups are widely used in community ecology and earth system modelling to describe trait variation within and across plant communities. However, this approach rests on the assumption that functional groups explain a large proportion of trait variation among species. We test whether four commonly used plant functional groups represent variation in six ecologically important plant traits. LOCATION: Tundra biome. TIME PERIOD: Data collected between 1964 and 2016. MAJOR TAXA STUDIED: 295 tundra vascular plant species. METHODS: We compiled a database of six plant traits (plant height, leaf area, specific leaf area, leaf dry matter content, leaf nitrogen, seed mass) for tundra species. We examined the variation in species-level trait expression explained by four traditional functional groups (evergreen shrubs, deciduous shrubs, graminoids, forbs), and whether variation explained was dependent upon the traits included in analysis. We further compared the explanatory power and species composition of functional groups to alternative classifications generated using post hoc clustering of species-level traits. RESULTS: Traditional functional groups explained significant differences in trait expression, particularly amongst traits associated with resource economics, which were consistent across sites and at the biome scale. However, functional groups explained 19% of overall trait variation and poorly represented differences in traits associated with plant size. Post hoc classification of species did not correspond well with traditional functional groups, and explained twice as much variation in species-level trait expression. MAIN CONCLUSIONS: Traditional functional groups only coarsely represent variation in well-measured traits within tundra plant communities, and better explain resource economic traits than size-related traits. We recommend caution when using functional group approaches to predict tundra vegetation change, or ecosystem functions relating to plant size, such as albedo or carbon storage. We argue that alternative classifications or direct use of specific plant traits could provide new insights for ecological prediction and modelling.

The role of adaptive strategies in plant naturalization.

Determining the factors associated with the naturalization of alien species is a central theme in ecology. Here, we tested the usefulness of a metric for quantifying Grime's seminal concept of adaptive strategies - competitors, stress-tolerators and ruderals (CSR) - to explain plant naturalizations worldwide. Using a global dataset of 3004 vascular plant species, and accounting for phylogenetic relatedness and species' native biomes, we assessed the associations between calculated C-, S- and R-scores and naturalization success for species exhibiting different life forms. Across different plant life forms, C-scores were positively and S-scores negatively associated with both the probability of naturalization and the number of regions where the species has naturalized. R-scores had positive effects on the probability of naturalization. These effects of the scores were, however, weak to absent for tree species. Our findings demonstrate the utility of CSR-score calculation to broadly represent, and potentially explain, the naturalization success of plant species.

Quantitative mesophyll parameters rather than whole-leaf traits predict response of C3 steppe plants to aridity.

We studied the impact of aridity on leaf and mesophyll traits in dominant and very abundant plant species of Eurasian steppe plant communities. We covered a 500-km latitudinal gradient across three vegetation zones in the Volga region of southern European Russia. Whole-leaf traits, volumetric fractions of leaf tissues, quantitative parameters of photosynthetic cells and chloroplasts, and chlorophyll, carbon (C) and nitrogen (N) contents were analyzed and related to plant functional type (PFT), type of mesophyll anatomy, phylogeny and climate aridity. The proportions of prevailing PFTs in the communities, such as C3 monocots, C3 dicots with dorsiventral and isopalisade anatomy and C4 dicots, changed with increasing aridity which influenced the whole-leaf parameters and tissue composition in the leaf. Leaf mass per unit area and leaf thickness slightly increased along the aridity gradient, but the most significant changes were observed in the mesophyll. Mesophyll cell surface area, chloroplast number and chloroplast surface area per unit leaf area were higher in C3 plants growing in the desert steppe compared with those of the forest steppe, while chlorophyll content per single chloroplast and per unit N content as well chlorophyll a/b ratio decreased. Our results identify a suite of mesophyll traits as a typical 'syndrome' of increasingly drought-adapted steppe plants.