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Summary Understanding the water‐use of plants is timely under increasing drought stress due to climate change. Despite the crucial role of roots in water uptake, relationships between water‐use and root traits are seldom considered. Combining a functional trait‐based approach with a water balance model, we tested whether root functional traits are related to spatial and temporal water‐use among 12 Mediterranean rangeland species grown in common garden monocultures. Soil water content was monitored for 10 months, and the dynamics of water uptake of each species was modelled at a daily time step. Root functional traits were measured at two soil depths (shallow and deep soil). Species with fast resource acquisition strategies in shallow soil, i.e. thin roots, maximised water uptake in a short period and consumed large amounts of water during periods of low water availability. Conversely, species with a more conservative root strategy, i.e. coarse roots, took up less water during the peak‐growing season, maintained water uptake over a longer period of time and consumed less water during periods of low water availability. Deep root traits are strongly related to species’ ability to take up water from deep soil. Deep roots with large diameters and low specific root length improve species’ ability to reach water from deep soil. Biomass investment in the deep soil layer was positively related to the amount of water consumed during periods of low water availability. Our results highlight that root functional traits influence a range of spatial and temporal water‐use among Mediterranean rangeland species. They account for the amount of water taken up during dry periods but not during the entire growing season. A lay summary is available for this article. Lay Summary

Comparative functional ecology seeks to understand why and how ecological systems and their components operate differently across environments. Although traditionally used in (semi)-natural situations, its concepts and methods could certainly apply to address key issues in the large variety of agricultural systems encountered across the world. In this review, we present major advances in comparative plant functional ecology that were made possible over the last two decades by the rapid development of a trait-based approach to plant functioning and prospects to apply it in agricultural situations. The strength of this approach is that it enables us to assess the interactions between organisms and their environment simultaneously on a large number of species, a prerequisite to address questions relative to species distribution, community assembly and ecosystem functioning. The trait concept will be first defined, before presenting a conceptual framework to understand the effects of environmental factors on plant community structure and ecosystem properties via plant traits. We will then argue that leading dimensions of variation among species can be captured by some selected traits and show that a combination of three easily measured traits—specific leaf area (the ratio of leaf area to leaf dry mass), plant height and seed mass—enables us to assess how different species use their resources, interact with neighbours and disperse in time and space. The use of traits to address central questions in community ecology will be reviewed next. It will be shown that traits allow us to (1) understand how plant species are sorted according to the nature of environmental gradients, (2) evaluate the relative importance of habitat filtering and limiting similarity in the process of community assembly and (3) quantify two main components of community functional structure, namely, community-weighted means of traits and community functional divergence. The relative impacts of these two components on ecosystem properties will then be discussed in the case of several components of primary productivity, litter decomposition, soil water content and carbon sequestration. There is strong support for the biomass ratio hypothesis, which states that the extent to which the traits of a species affect those ecosystem properties depends on the abundance of this species in the community. Assessing the role of functional divergence among species on ecosystem properties will require major methodological breakthroughs, both in terms of metrics and statistical procedures to be used. In agricultural situations, we show that trait-based approaches have been successfully developed to assess the impacts of management practices on (1) the agronomic value of grasslands and (2) the functional composition and structure of crop weed communities and how these could affect the functioning of the crop. Applications in forestry are still poorly developed, especially in temperate regions where the number of species in managed forest remains relatively low. The last decades of research have led to the constitution of large data sets of plant traits, which remain poorly compatible and accessible. Recent advances in the field of ecoinformatics suggest that major progress could be achieved in this area by using improved metadata standards and advancing trait domain ontologies. Finally, concluding remarks, unanswered questions and directions for research using the functional approach to biodiversity made possible by the use of traits will be discussed in the contexts of ecological and agronomical systems. The latter indeed cover a wide range of environmental conditions and biological diversity, and the prospect for reducing environmental impacts in highly productive, low-diversity systems will certainly imply improving our skills for the management of more diverse systems prone to a trait-based approach as reviewed here.

1. Understanding how environmental factors drive plant community assembly remains a major challenge in community ecology. The strength of different assembly processes along environmental gradients, such as environmental filtering and functional niche differentiation, can be quantified by analysing trait distributions in communities. While environmental filtering affects species occurrence among communities, functional divergence or convergence is strongly related to species abundances within communities, which few studies have taken into account. We examine the trait-mediated effect of these two processes along a stress-resource gradient. 2. We measured species abundances and the distributions of eight traits related to vegetative and regenerative phases in plant communities along a gradient of soil depth and resource availability in Mediterranean rangelands. We quantified environmental filtering, defined as a local restriction of trait range, and trait divergence, based on abundance-weighted trait variance, using a two-step approach with specifically designed null models. 3. Communities presented a clear functional response to the soil gradient, as evidenced by strong trends in community-weighted trait means. We detected environmental filtering of different traits at both ends of the gradient, suggesting that, contrary to widespread expectations, trait filtering may not necessarily be the result of abiotic filtering under harsh conditions but could likely also result from biotic interactions in productive habitats. 4. We found marked shifts in trait abundance distributions within communities along the gradient. Vegetative traits (e.g. leaf dry matter content) diverged on shallow soils, reflecting the coexistence of distinct water- and nutrient-use strategies in these constrained habitats and converged with increasing soil resource availability. By contrast, regenerative traits (e.g. seed mass) tended to diverge towards deeper soils, while plant reproductive heights diverged all along the gradient. 5. Synthesis: Our study highlights how the combination of abundance data with traits capturing different functional niches is critical to the detection of complex functional responses of plant communities to environmental gradients. We demonstrate that patterns of trait divergence and filtering are strongly contingent on both trait and environment such that there can be no expectation of a simple trend of increasing or decreasing functional divergence along a gradient of resource availability.

In its simplest definition, a trait is a surrogate of organismal performance, and this meaning of the term has been used by evolutionists for a long time. Over the last three decades, developments in community and ecosystem ecology have forced the concept of trait beyond these original boundaries, and trait-based approaches are now widely used in studies ranging from the level of organisms to that of ecosystems. Despite some attempts to fix the terminology, especially in plant ecology, there is currently a high degree of confusion in the use, not only of the term \"trait\" itself, but also in the underlying concepts it refers to. We therefore give an unambiguous definition of plant trait, with a particular emphasis on functional trait. A hierarchical perspective is proposed, extending the \"performance paradigm\" to plant ecology. \"Functional traits\" are defined as morpho-physio-phenological traits which impact fitness indirectly via their effects on growth, reproduction and survival, the three components of individual performance. We finally present an integrative framework explaining how changes in trait values due to environmental variations are translated into organismal performance, and how these changes may influence processes at higher organizational levels. We argue that this can be achieved by developing \"integration functions\" which can be grouped into functional response (community level) and effect (ecosystem level) algorithms.

The effects of increased atmospheric CO2concentrations on vegetative growth and competitive performance were evaluated, using five meta-analyses. Paying special attention to functional groups, we analysed responses at three integration levels: carbon economy parameters, vegetative biomass of isolated plants, and growth in competition. CO2effects on seed biomass and plant-to-plant variability were also studied. Underlying the growth stimulation is an increased unit leaf rate (ULR), especially for herbaceous dicots. This is mainly caused by an increase in the whole-plant rate of photosynthesis. The increased ULR is accompanied by a decrease in specific leaf area. The net result of these and other changes is that relative growth rate is only marginally stimulated. The biomass enhancement ratio (BER) of individually grown plants varies substantially across experiments/species, and size variability in the experimental populations is a vital factor in this. Fast-growing herbaceous C3 species respond more strongly than slow-growing C3 herbs or C4 plants. CAM species and woody plants show intermediate responses. When grown in competition, C4 species show lowest responses to elevated CO2at high nutrient conditions, whereas at low nutrient levels N2-fixing dicots respond relatively strongly. No systematic differences were found between slow- and fast-growing species. BER values obtained for isolated plants cannot be used to estimate BER of the same species grown in interspecific competition - the CO2response of monocultures may be a better predictor.

Summary The interactions between hydrological and ecological processes are key issues to improve our predictions of ecosystem responses to increasing droughts. However, predicting the dynamics and the impacts of vegetation water stress remains challenging because of complex ecohydrological feedbacks. The ecohydrological optimality approach proposes that functional adjustments within plant communities may buffer the increase in vegetation water stress despite local water shortage. This study aimed to test whether vegetation water stress may be invariant across contrasting plant communities, reflecting possible optimality processes. We addressed the following question: does a lower soil water storage capacity under the same climate generate greater vegetation water stress over time? We hypothesized that vegetation water stress would be buffered around a low and constant level through the adjustment of vegetation biomass productivity net primary productivity (NPP), evapotranspiration (ET) and/or water‐use efficiency (WUE) in relation with local soil water storage capacity. We monitored 12 native plant communities distributed along a gradient of soil water storage capacity (ranging from 20 mm to 120 mm) during five successive years. Net primary productivity, ET, WUE as well as soil water dynamics were assessed and modelled for each plant community throughout the 5 years of study. Vegetation water stress was determined for each plant community as the deviation of between actual ET and their maximum ETm rate achieved under non‐limiting conditions. We found that NPP and ET were together proportionally related to local soil water storage capacity across the 5 years of study while WUE did not differ between plant communities. Vegetation water stress was found quite similar for all plant communities whatever the soil water storage capacity. These results suggested that vegetation water stress was strongly buffered by the community‐level plant growth rates and total water use along the soil gradient, but not by WUE. Our results suggest that stressful environments rarely exist for plant communities. A dynamic scaling relationship between NPP and ET may underpin the control of vegetation water stress over seasonal and pluriannual time‐scales. Such results could contribute to better understanding processes associated with ecohydrological optimality and improve the predictions of vegetation dynamics under increasing droughts. A lay summary is available for this article. Lay Summary

Questions: How can one explicitly quantify, and separately measure, stress and disturbance gradients? How do these gradients affect functional composition in early successional plant communities and to what extent? Can we accurately predict trait composition from knowledge of these gradients? Location: Southern Quebec, Canada. Methods: Using eight environmental variables measured in 48 early successional plant communities, we estimated stress and disturbance gradients through structural equation modelling. We then measured 10 functional traits on the most abundant species of these 48 communities and calculated their community-level mean and variance weighted by the relative abundance of each species. Finally, we related these community-weighted means and variances to the estimated stress and disturbance gradients using general linear models or generalized additive models. Results: We obtained a well-fitting measurement model of the stress and disturbance gradients existing in our sites. Of the 10 studied traits, only average plant reproductive height was strongly correlated with the stress (r² = 0.464) and disturbance (r² = 0.543) gradients. Leaf traits were not significantly related to either the stress or disturbance gradients. Conclusions: The well-fitting measurement model of the stress and disturbance gradients, combined with the generally weak trait-environment linkages, suggests that community assembly in these early successional plant communities is driven primarily by stochastic processes linked to the history of arrival of propagules and not to trait-based environmental filtering.

Phenology, the study of seasonal timing of events in nature, plays a key role in the matching between organisms and their environment. Yet, it has been poorly integrated in trait-based descriptions of the plant phenotype. Here, we focus on three phases of reproductive phenology – time of flowering, time of seed dispersal and duration of seed maturation – to test how these traits relate to other recognized dimensions of plant functioning.Traits describing reproductive phenology, together with reproductive plant height, seed mass, area of a leaf, and traits involved in leaf economics, were compiled for 139 species growing under Mediterranean climate conditions.Across all species, flowering time was positively related to reproductive height, while the duration of seed maturation was related to leaf economics. Relationships differed among growth forms, however: flowering time and reproductive height were related both in annuals and in herbaceous perennials, whereas the duration of seed maturation was related to seed mass only in annuals; no correlations were found for woody species.Phenology relates to other dimensions of plant functioning in a complex manner, suggesting that it should be considered as an independent dimension in the context of plant strategies.

The phenology of organisms corresponds to the temporal match between the components of their life cycle and the seasonal distribution of resources and hazards. Flowering has been extensively studied to describe the reproductive phenology of plants, but in comparison, other key events for reproductive success such as the seed maturation period and the time of seed dispersal have been considerably less investigated.This study describes the temporal sequence of onset of flowering and seed dispersal, and the time necessary to mature seeds in 138 species growing in the strongly seasonal climate of Mediterranean southern France. Data for the three traits were compiled from several original surveys to characterize the reproductive phenology of 47 annual, 67 perennial herbaceous and 24 low stature woody species. The timing of these three phases was assessed in relation to local climatic conditions, and the degree to which they were coordinated was tested.All three phenological traits spanned a wide range of values from early spring to late summer. Annuals flowered slightly earlier than perennials but the largest difference between these groups was found for the seed maturation period, which was much shorter in annuals. This resulted in earlier dispersal dates in these species, which occurred before periods of high water deficit. Significant positive correlations were found between onset of flowering, onset of seed dispersal and seed maturation period. This suggests a consistent pattern of coordination between the different phases of reproductive phenology across species.Our results show that while the time slot for flowering appears restricted to periods with adequate temperature and water availability for most species, the seed maturation period and dispersal phase can occur during periods of substantial water deficit, at least for perennials. They also suggest that the different species can be arrayed along a fast-slow continuum based on reproductive events, from early flowering species with short seed maturation and early dispersal to late flowering species with long seed maturation and late dispersal. Whether this relates to the postulated fast-slow continuum identified for the functioning of vegetative organs is a promising avenue for future research.