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Increasing aridity associated with climate change may lead to the crossing of critical ecosystem thresholds in drylands, compromising ecosystem services for millions of people. In this context, finding tools to detect at early stages the effects of increasing aridity on ecosystems is extremely urgent to avoid irreversible damage. Here, we assess shifts in plant community functional structure along a spatial aridity gradient in tropical dryland (Brazilian Caatinga), to select the most appropriate plant functional groups as ecological indicators likely useful to predict temporal ecosystem trajectories in response to aridity. We identified seven plant functional groups based on 13 functional traits associated with plant establishment, defense, regeneration, and dispersal, whose relative abundances changed, linearly and non-linearly, with increasing aridity, showing either increasing or decreasing trends. Of particular importance is the increase in abundance of plants with high chemical defense and Crassulacean Acid Metabolism (CAM) photosynthetic pathway, with increasing aridity. We propose the use of these functional groups as early warning indicators to detect aridity impacts on these dryland ecosystems and shifts in ecosystem functioning. This information can also be used in the elaboration of mitigation and ecological restoration measures to prevent and revert current and future climate change impacts on tropical dry forests.

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BACKGROUND AND AIMS: Trait-based frameworks assess plant survival strategies using different approaches. Some frameworks use functional traits to assign species to a priori defined ecological strategies. Others use functional traits as the central element of a species ecophysiological strategy. We compared these two approaches by asking: (1) what is the primary ecological strategy of three dominant co-occurring shrub species from inselbergs based on the CSR scheme, and (2) what main functional traits characterize the ecophysiological strategy of the species based on their use of carbon, water and light? METHODS: We conducted our study on a Colombian inselberg. In this extreme environment with multiple stressors (high temperatures and low resource availability), we expected all species to be stress tolerant (S in the CSR scheme) and have similar ecophysiological strategies. We measured 22 anatomical, morphological and physiological leaf traits. KEY RESULTS: The three species have convergent ecological strategies as measured by CSR (S, Acanthella sprucei; and S/CS, Mandevilla lancifolia and Tabebuia orinocensis) yet divergent resource-use strategies as measured by their functional traits. A. sprucei has the most conservative carbon use, risky water use and a shade-tolerant strategy. M. lancifolia has acquisitive carbon use, safe water use and a shade-tolerant strategy. T. orinocensis has intermediate carbon use, safe water use and a light-demanding strategy. Additionally, stomatal traits that are easy to measure are valuable to describe resource-use strategies because they are highly correlated with two physiological functions that are hard to measure: stomatal conductance and maximum photosynthesis per unit mass. CONCLUSIONS: The two approaches provide complementary information on species strategies. Plant species can co-occur in extreme environments, such as inselbergs, because they exhibit convergent primary ecological strategies but divergent ecophysiological strategies, allowing them to use limiting resources differently.

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The false Rhodes grasses [Leptochloa crinita (Lag.) P.M. Peterson and N.W. Snow and Leptochloa pluriflora (E. Fourn.) P.M. Peterson and N.W. Snow] are considered valuable native forage resources for arid and semiarid rangelands in Argentina and the United States. Effectively using plant materials as forage under aridity conditions requires understanding their resource allocation under those conditions. In the present study, plant functional traits were evaluated in six populations of each false Rhodes grass species from different geographic origin in a humid and an arid region. The evaluation was focused on seed weight, due to the key role of this trait in plant survival. The implication of seed weight in germination under osmotic stress and trade-off relationships between functional traits were also analysed. A fixed ontogenetic variation was found in both species, since populations maintained a stable seed weight across environments. The tolerance to osmotic stress at germination stage was more related to seed weight than to population origin or maternal environment of seeds; heavier-seeded populations produced heavier seedlings instead of a higher number of germinated seeds or higher germination rates. Some traits varied between environments but other traits exhibited a fixed response. Variation patterns among populations were similar within environments and in some cases even for populations from the same geographic origin, revealing a fixed ontogenetic variation; this phenomenon was clearer in L. crinita than in L. pluriflora. Moreover, several different trade-off strategies were detected in both species. These results reinforce the knowledge about the key role of seed weight in survival and performance of seedlings at initial growth stages under arid conditions; however, at advanced stages, other traits would have an important function in growth and development of false Rhodes grasses.

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The study of above- and below-ground organ plant coordination is crucial for understanding the biophysical constraints and trade-offs involved in species' performance under different environmental conditions. Environmental stress is expected to increase constraints on species trait combinations, resulting in stronger coordination among the organs involved in the acquisition and processing of the most limiting resource. To test this hypothesis, we compared the coordination of trait combinations in 94 tree seedling species from two tropical forest systems in Mexico: dry and moist. In general, we expected that the water limitation experienced by dry forest species would result in stronger leaf-stem-root coordination than light limitation experienced by moist forest species. Using multiple correlations analyses and tools derived from network theory, we found similar functional trait coordination between forests. However, the most important traits differed between the forest types. While in the dry forest the most central traits were all related to water storage (leaf and stem water content and root thickness), in the moist forest they were related to the capacity to store water in leaves (leaf water content), root efficiency to capture resources (specific root length), and stem toughness (wood density). Our findings indicate that there is a shift in the relative importance of mechanisms to face the most limiting resource in contrasting tropical forests.

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Microclimatic conditions change dramatically as forests age and impose strong filters on community assembly during succession. Light availability is the most limiting environmental factor in tropical wet forest succession; by contrast, water availability is predicted to strongly influence tropical dry forest (TDF) successional dynamics. While mechanisms underlying TDF successional trajectories are not well understood, observational studies have demonstrated that TDF communities transition from being dominated by species with conservative traits to species with acquisitive traits, the opposite of tropical wet forest. Determining how functional traits predict TDF tree species' responses to changing environmental conditions could elucidate mechanisms underlying tree performance during TDF succession. We implemented a 6-ha restoration experiment on a degraded Vertisol in Costa Rica to determine (1) how TDF tree species with different resource-use strategies performed along a successional gradient and (2) how ecophysiological functional traits correlated with tree performance in simulated successional stages. We used two management treatments to simulate distinct successional stages including: clearing all remnant vegetation (early-succession), or interplanting seedlings with no clearing (mid-succession). We crossed these two management treatments (cleared/interplanted) with two species mixes with different resource-use strategies (acquisitive/conservative) to examine their interaction. Overall seedling survival after 2 yr was low, 15.1-26.4% in the four resource-use-strategy × management-treatment combinations, and did not differ between the management treatments or resource-use-strategy groups. However, seedling growth rates were dramatically higher for all species in the cleared treatment (year 1, 69.1% higher; year 2, 143.3% higher) and defined resource-use strategies had some capacity to explain seedling performance. Overall, ecophysiological traits were better predictors of species' growth and survival than resource-use strategies defined by leaf and stem traits such as specific leaf area. Moreover, ecophysiological traits related to water use had a stronger influence on seedling performance in the cleared, early-successional treatment, indicating that the influence of microclimatic conditions on tree survival and growth shifts predictably during TDF succession. Our findings suggest that ecophysiological traits should be explicitly considered to understand shifts in TDF functional composition during succession and that using these traits to design species mixes could greatly improve TDF restoration outcomes.

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Although populations are phenotypically diverse, the majority of trait-based studies have focused on examining differences among species. The justification for this broadly applied approach is based on the assumption that differences among species are always greater than within species. This is likely true for local communities, but species are often broadly distributed across a wide range of environments and patterns of intraspecific variation might surpass differences among species. Therefore, an appropriate interpretation of the functional diversity requires an assessment of patterns of trait variation across different ecological scales. In this study, we examine and characterize patterns of leaf trait variation for species that are broadly distributed along an elevational gradient. We focus on seven leaf traits that represent a main axis of functional differentiation in plants reflecting the balance between photosynthetic efficiency, display, and stomatal conductance. We evaluated patterns of trait variance across ecological scales (elevation, species, populations, and individuals) and examined trait covariance at both within species and across species levels, along the elevation gradient. Our results show three key patterns: (1) intraspecific leaf trait variation for broadly distributed species is comparable to the interspecific trait variation, (2) the trait variance structure is highly variable across species, and (3) trait coordination between pairs of leaf traits is evident across species along the gradient, but not always within species. Combined, our results show that trait coordination and covariance are highly idiosyncratic across broadly distributed and co-occurring species, indicating that species may achieve similar functional roles even when exhibiting different phenotypes. This result challenges the traditional paradigm of functional ecology that assumes single trait values as optimal solutions for environments. In conclusion, patterns of trait variation both across and within species should be considered in future studies that assess trade-offs among traits over environmental gradients.

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Bark is a structure involved in multiple physiological functions, but which has been traditionally associated with protection against fire. Thus, little is known about how the morpho-anatomical variations of this structure are related to different ecological pressures, especially in tropical savanna species, which are commonly subjected to frequent fire and drought events. Here we evaluated how the structural and functional variations of bark are related to the processes of resilience and resistance to fire, as well as transport and storage of water in 31 native species from the Brazilian Cerrado. Because of their thick bark, none of the trees analyzed were top-killed after a severe fire event. The structural and functional variations of the bark were also associated with water storage and transport, functions related to properties of the inner bark. In fact, species with a thicker and less dense inner bark were the ones that had the highest water contents in the wood, bark, and leaves. Lower bark density was also related to higher stem hydraulic conductivity, carbon assimilation, and growth. Overall, we provide strong evidence that in addition to protection from fire, the relative investment in bark also reflects different strategies of water use and conservation among many Cerrado tree species.

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Tropical forests contribute significantly to the global carbon cycle, but little is known about the temperature response of photosynthetic carbon uptake in tropical species, and how this varies within and across forests. We determined in situ photosynthetic temperature-response curves for upper canopy leaves of 42 tree and liana species from two tropical forests in Panama with contrasting rainfall regimes. On the basis of seedling studies, we hypothesized that species with high photosynthetic capacity - light-demanding, fast-growing species - would have a higher temperature optimum of photosynthesis (TOpt ) than species with low photosynthetic capacity - shade-tolerant, slow-growing species - and that, therefore, TOpt would scale with the position of a species on the slow-fast continuum of plant functional traits. TOpt was remarkably similar across species, regardless of their photosynthetic capacity and other plant functional traits. Community-average TOpt was almost identical to mean maximum daytime temperature, which was higher in the dry forest. Photosynthesis above TOpt appeared to be more strongly limited by stomatal conductance in the dry forest than in the wet forest. The observation that all species in a community shared similar TOpt values suggests that photosynthetic performance is optimized under current temperature regimes. These results should facilitate the scaling up of photosynthesis in relation to temperature from leaf to stand level in species-rich tropical forests.

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BACKGROUND AND AIMS: Ecologists are increasingly using plant functional traits to predict community assembly, but few studies have linked functional traits to species' responses to fine-scale resource gradients. In this study, it was tested whether saplings of woody species partition fine-scale gradients in light availability based on their leaf mass per area (LMA) in three temperate rain forests and one Mediterranean forest in southern Chile. METHODS: LMA was measured under field conditions of all woody species contained in approx. 60 plots of 2 m2 in each site, and light availability, computed as the gap light index (GLI), was determined. For each site, species' pairwise differences in mean LMA (Δ LMA) and abundance-weighted mean GLI (Δ light response) of 2 m2 plots were calculated and it was tested whether they were positively related using Mantel tests, i.e. if species with different LMA values differed in their response to light availability. Additionally linear models were fitted to the relationship between plot-level mean LMA and GLI across plots for each site. KEY RESULTS: A positive and significant relationship was found between species' pairwise differences in mean LMA and differences in light response across species for all temperate rain forests, but not for the Mediterranean forest. The results also indicated a significant positive interspecific link between LMA and light availability for all forests. This is in contrast to what is traditionally reported and to expectations from the leaf economics spectrum. CONCLUSIONS: In environments subjected to light limitation, interspecific differences in a leaf trait (LMA) can explain the fine-scale partitioning of light availability gradients by woody plant species. This niche partitioning potentially facilitates species coexistence at the within-community level. The high frequency of evergreen shade-intolerant species in these forests may explain the positive correlation between light availability and LMA.

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Leaves as the main photosynthetic organ of plants must be well protected against various hazards to achieve their optimal lifespans. Yet, within-species variation and the material basis of leaf strength have been explored for very few species. Here, we present a large dataset of leaf fracture toughness from a species-rich humid tropical forest on Barro Colorado Island, Panama, reporting both among- and within-species variation in relation to light environment (sun-lit canopy versus shaded understorey) and ontogeny (seedlings versus adults). In this dataset encompassing 281 free-standing woody species and 428 species-light combinations, lamina fracture toughness varied ca 10 times. A central objective of our study was to identify generalizable patterns in the structural and material basis for interspecific variation in leaf lamina fracture toughness. The leaf lamina is a heterogeneous structure in which strong materials in cell walls, such as cellulose and lignin, contribute disproportionately to fracture toughness. We found significant increases in leaf fracture toughness from shade to sun and from seedling leaves to adult leaves. Both within and across species, leaf fracture toughness increased with total bulk density (dry biomass per unit volume) and cellulose mass concentration, but decreased with mass concentrations of lignin and hemicelluose. These bivariate relationships shift between light environments, but leaf cellulose density (cellulose mass per unit leaf volume) exhibits a common relationship with lamina fracture toughness between light environments and through ontogeny. Hence, leaf cellulose density is probably a universal predictor of leaf fracture toughness.

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The present study investigated the dynamic on a short-time scale in the vegetation in moist grassland of the Sete Cidades National Park, Piauí. Herb-subshrub layer samples was carried out in July 2007, 2009 and 2011. Changes in structural, floristic and functional traits in the community were assessed by species richness, diversity and similarity indices between those periods, as well as by hierarchical classification and ordination. Cluster and Principal Component Analyses identified functional groups according to 23 species trait state. To distinguish the contribution of space and time configuration in the community structure, we used the variance partition technique. The functional groups of chamaephytes and therophytes II were associated with wetter sites, while the groups of non-tussock hemicriptophytes I, tussock hemicriptophytes (FG4), and geophytes (FG5) were associated to the drier ones. We found a non-accelerated dynamics, at least on a short-time scale, represented by some descriptors in the community, such as the close similarity between the inventories and ordering of sampling transects in moist grassland. Therefore, besides considering the partition of the temporal niche as a mechanism for the co-existence of species, the heterogeneity of space dictated by environmental filters seems to determine the stability of the this grassland communities over time.

O presente estudo investigou a dinâmica em uma curta escala de tempo na vegetação de campo limpo úmido do Parque Nacional Sete Cidades Park, Piauí. Amostragens da camada herbáceo-subarbustiva foram realizadas em julho de 2007, 2009 e 2011. Mudanças estruturais, florísticas e de atributos funcionais da comunidade foram avaliadas por índices de riqueza de espécies, diversidade e similaridade entre esses períodos, bem como por classificação hierárquica e ordenação. Análise de Componentes Principais e de Cluster identificaram grupos funcionais de acordo com 23 atributos das espécies. Para distinguir a contribuição do espaço e do tempo nas análises, usamos técnicas de partição de variância. Os grupos funcionais das caméfitas e terófitas II foram associados com os locais mais úmidos, enquanto os grupos de hemicriptófitas I não entouceiradas, hemicriptófitos entouceiradas e das geófitas (GF5) foram associados aos mais secos. Nós encontramos uma dinâmica não-acelerada, pelo menos em uma escala de curta duração, representada por alguns descritores da comunidade, tais como a similaridade entre os inventários e a ordenação dos transectos no campo limpo úmido. Assim, além de considerar a partição do nicho temporal como um mecanismo para a coexistência de espécies, a heterogeneidade do espaço ditada por filtros ambientais parece determinar a estabilidade destas comunidades campestres ao longo do tempo.

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The present study investigated the dynamic on a short-time scale in the vegetation in moist grassland of the Sete Cidades National Park, Piauí. Herb-subshrub layer samples was carried out in July 2007, 2009 and 2011. Changes in structural, floristic and functional traits in the community were assessed by species richness, diversity and similarity indices between those periods, as well as by hierarchical classification and ordination. Cluster and Principal Component Analyses identified functional groups according to 23 species trait state. To distinguish the contribution of space and time configuration in the community structure, we used the variance partition technique. The functional groups of chamaephytes and therophytes II were associated with wetter sites, while the groups of non-tussock hemicriptophytes I, tussock hemicriptophytes (FG4), and geophytes (FG5) were associated to the drier ones. We found a non-accelerated dynamics, at least on a short-time scale, represented by some descriptors in the community, such as the close similarity between the inventories and ordering of sampling transects in moist grassland. Therefore, besides considering the partition of the temporal niche as a mechanism for the co-existence of species, the heterogeneity of space dictated by environmental filters seems to determine the stability of the this grassland communities over time.(AU)

O presente estudo investigou a dinâmica em uma curta escala de tempo na vegetação de campo limpo úmido do Parque Nacional Sete Cidades Park, Piauí. Amostragens da camada herbáceo-subarbustiva foram realizadas em julho de 2007, 2009 e 2011. Mudanças estruturais, florísticas e de atributos funcionais da comunidade foram avaliadas por índices de riqueza de espécies, diversidade e similaridade entre esses períodos, bem como por classificação hierárquica e ordenação. Análise de Componentes Principais e de Cluster identificaram grupos funcionais de acordo com 23 atributos das espécies. Para distinguir a contribuição do espaço e do tempo nas análises, usamos técnicas de partição de variância. Os grupos funcionais das caméfitas e terófitas II foram associados com os locais mais úmidos, enquanto os grupos de hemicriptófitas I não entouceiradas, hemicriptófitos entouceiradas e das geófitas (GF5) foram associados aos mais secos. Nós encontramos uma dinâmica não-acelerada, pelo menos em uma escala de curta duração, representada por alguns descritores da comunidade, tais como a similaridade entre os inventários e a ordenação dos transectos no campo limpo úmido. Assim, além de considerar a partição do nicho temporal como um mecanismo para a coexistência de espécies, a heterogeneidade do espaço ditada por filtros ambientais parece determinar a estabilidade destas comunidades campestres ao longo do tempo.(AU)

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