Intraspecific variation in fine-root traits is larger than in aboveground traits in European herbaceous species regardless of drought.

Differences within species (Intraspecific trait variation - ITV) contribute substantially to overall trait variability and environmental harshness can reduce among-species variation. While aboveground traits have received considerable attention, knowledge about ITV in fine-root traits and how it differs from ITV in aboveground traits remains limited. This study examined the partitioning of trait variation aboveground and fine-root traits in 52 European herbaceous species and how such proportions change in response to drought, offering valuable insights for accurate functional species characterization and inter-species comparisons. We studied seven morphological aboveground and fine-root traits under drought and well-watered conditions in a greenhouse experiment. Linear mixed effect models and permutational multivariate analysis of variance (PERMANOVA) were employed to decompose trait variation, ensuring the robustness of our results. We also calculated variance partitioning for the combination of aboveground traits and the combination of fine-root traits, as well as pairs of analogous leaf and fine-root traits (i.e., traits that fulfill similar functions) for each treatment (control and drought). Among-species trait differences explained a greater proportion of overall variance than within-species variation, except for root dry matter content (RDMC). Height and leaf area stood out, with species' identity accounting for 87-90% of total trait variation. Drought had no significant effect on the proportions of variation in any of the traits. However, the combination of fine-root traits exhibited higher intraspecific variability (44-44%) than aboveground traits (19-21%) under both drought and control. Analogous root traits also showed higher ITV (51-50%) than analogous leaf traits (27-31%). Our findings highlight substantial within-species variation and the nuanced responses of fine-root traits, particularly RDMC, suggesting root traits' flexibility to soil heterogeneity that fosters less differentiation among species. Among-species trait differences, especially aboveground, may underscore distinct strategies and competitive abilities for resource acquisition and utilization. This study contributes to elucidate the mechanisms underlying the multifunctionality of the above- and belowground plants compartments.

The great escape: patterns of enemy release are not explained by time, space or climate.

When a plant is introduced to a new ecosystem it may escape from some of its coevolved herbivores. Reduced herbivore damage, and the ability of introduced plants to allocate resources from defence to growth and reproduction can increase the success of introduced species. This mechanism is known as enemy release and is known to occur in some species and situations, but not in others. Understanding the conditions under which enemy release is most likely to occur is important, as this will help us to identify which species and habitats may be most at risk of invasion. We compared in situ measurements of herbivory on 16 plant species at 12 locations within their native European and introduced Australian ranges to quantify their level of enemy release and understand the relationship between enemy release and time, space and climate. Overall, plants experienced approximately seven times more herbivore damage in their native range than in their introduced range. We found no evidence that enemy release was related to time since introduction, introduced range size, temperature, precipitation, humidity or elevation. From here, we can explore whether traits, such as leaf defences or phylogenetic relatedness to neighbouring plants, are stronger indicators of enemy release across species.

Environmental heterogeneity modulates the effect of plant diversity on the spatial variability of grassland biomass.

Plant productivity varies due to environmental heterogeneity, and theory suggests that plant diversity can reduce this variation. While there is strong evidence of diversity effects on temporal variability of productivity, whether this mechanism extends to variability across space remains elusive. Here we determine the relationship between plant diversity and spatial variability of productivity in 83 grasslands, and quantify the effect of experimentally increased spatial heterogeneity in environmental conditions on this relationship. We found that communities with higher plant species richness (alpha and gamma diversity) have lower spatial variability of productivity as reduced abundance of some species can be compensated for by increased abundance of other species. In contrast, high species dissimilarity among local communities (beta diversity) is positively associated with spatial variability of productivity, suggesting that changes in species composition can scale up to affect productivity. Experimentally increased spatial environmental heterogeneity weakens the effect of plant alpha and gamma diversity, and reveals that beta diversity can simultaneously decrease and increase spatial variability of productivity. Our findings unveil the generality of the diversity-stability theory across space, and suggest that reduced local diversity and biotic homogenization can affect the spatial reliability of key ecosystem functions.

Fine-root traits in the global spectrum of plant form and function.

Plant traits determine how individual plants cope with heterogeneous environments. Despite large variability in individual traits, trait coordination and trade-offs1,2 result in some trait combinations being much more widespread than others, as revealed in the global spectrum of plant form and function (GSPFF3) and the root economics space (RES4) for aboveground and fine-root traits, respectively. Here we combine the traits that define both functional spaces. Our analysis confirms the major trends of the GSPFF and shows that the RES captures additional information. The four dimensions needed to explain the non-redundant information in the dataset can be summarized in an aboveground and a fine-root plane, corresponding to the GSPFF and the RES, respectively. Both planes display high levels of species aggregation, but the differentiation among growth forms, families and biomes is lower on the fine-root plane, which does not include any size-related trait, than on the aboveground plane. As a result, many species with similar fine-root syndromes display contrasting aboveground traits. This highlights the importance of including belowground organs to the GSPFF when exploring the interplay between different natural selection pressures and whole-plant trait integration.

Extinction of threatened vertebrates will lead to idiosyncratic changes in functional diversity across the world.

Although species with larger body size and slow pace of life have a higher risk of extinction at a global scale, it is unclear whether this global trend will be consistent across biogeographic realms. Here we measure the functional diversity of terrestrial and freshwater vertebrates in the six terrestrial biogeographic realms and predict their future changes through scenarios mimicking a gradient of extinction risk of threatened species. We show vastly different effects of extinctions on functional diversity between taxonomic groups and realms, ranging from almost no decline to deep functional losses. The Indo-Malay and Palearctic realms are particularly inclined to experience a drastic loss of functional diversity reaching 29 and 31%, respectively. Birds, mammals, and reptiles regionally display a consistent functional diversity loss, while the projected losses of amphibians and freshwater fishes differ across realms. More efficient global conservation policies should consider marked regional losses of functional diversity across the world.

Erosion of global functional diversity across the tree of life.

Although one-quarter of plant and vertebrate species are threatened with extinction, little is known about the potential effect of extinctions on the global diversity of ecological strategies. Using trait and phylogenetic information for more than 75,000 species of vascular plants, mammals, birds, reptiles, amphibians, and freshwater fish, we characterized the global functional spectra of each of these groups. Mapping extinction risk within these spectra showed that larger species with slower pace of life are universally threatened. Simulated extinction scenarios exposed extensive internal reorganizations in the global functional spectra, which were larger than expected by chance for all groups, and particularly severe for mammals and amphibians. Considering the disproportionate importance of the largest species for ecological processes, our results emphasize the importance of actions to prevent the extinction of the megabiota.

Seeds tend to disperse further in the tropics.

Despite the importance of seed dispersal in a plant's life cycle, global patterns in seed dispersal distance have seldom been studied. This paper presents the first geographically and taxonomically broad quantification of the latitudinal gradient in seed dispersal distance. Although there is substantial variation in the seed dispersal distances of different species at a given latitude, seeds disperse on average more than an order of magnitude further at the equator than towards the poles. This pattern is partially explained by plant life-history traits that simultaneously associate with seed dispersal distance and latitude, including dispersal mode and plant height. The extended seed shadow of tropical plants could increase the distance between conspecific individuals. This could facilitate species coexistence and contribute to the maintenance of high plant diversity in tropical communities. The latitudinal gradient in dispersal distance also has implications for species' persistence in the face of habitat fragmentation and climate change.

Can dispersal investment explain why tall plant species achieve longer dispersal distances than short plant species?

Tall plant species disperse further distances than do short species, within and across dispersal syndromes, yet the driver underpinning this relationship is unclear. The ability of taller plants to invest more in dispersal structures may explain the positive relationship between plant height and dispersal distance. Here, we quantify the cross-species relationships between presence of dispersal structures, dispersal investment plant height and dispersal distance. Plant height, dispersal syndrome and dispersal investment data were collated for 1613 species from the literature, with dispersal distance data collated for 114 species. We find that species with high dispersal investment disperse further than do species with low dispersal investment. Tall species have a greater probability of having dispersal structures on their seeds compared with short species. For species with dispersal structures on their seeds, plant height is very weakly related to dispersal investment. Our results provide the first global confirmation of the dispersal investment-distance hypothesis, and show dispersal investment can be used for predicting species dispersal distances. However, our results and those of previous studies indicate plant height is still the best proxy for estimating species dispersal distances due to it being such a readily available plant trait.

Species pools, community completeness and invasion: disentangling diversity effects on the establishment of native and alien species.

Invasion should decline with species richness, yet the relationship is inconsistent. Species richness, however, is a product of species pool size and biotic filtering. Invasion may increase with richness if large species pools represent weaker environmental filters. Measuring species pool size and the proportion realised locally (completeness) may clarify diversity-invasion relationships by separating environmental and biotic effects, especially if species' life-history stage and origin are accounted for. To test these relationships, we added seeds and transplants of 15 native and alien species into 29 grasslands. Species pool size and completeness explained more variation in invasion than richness alone. Although results varied between native and alien species, seed establishment and biotic resistance to transplants increased with species pool size, whereas transplant growth and biotic resistance to seeds increased with completeness. Consequently, species pools and completeness represent multiple independent processes affecting invasion; accounting for these processes improves our understanding of invasion.

Functional diversity enhances the resistance of ecosystem multifunctionality to aridity in Mediterranean drylands.

We used a functional trait-based approach to assess the impacts of aridity and shrub encroachment on the functional structure of Mediterranean dryland communities (functional diversity (FD) and community-weighted mean trait values (CWM)), and to evaluate how these functional attributes ultimately affect multifunctionality (i.e. the provision of several ecosystem functions simultaneously). Shrub encroachment (the increase in the abundance/cover of shrubs) is a major land cover change that is taking place in grasslands worldwide. Studies conducted on drylands have reported positive or negative impacts of shrub encroachment depending on the functions and the traits of the sprouting or nonsprouting shrub species considered. FD and CWM were equally important as drivers of multifunctionality responses to both aridity and shrub encroachment. Size traits (e.g. vegetative height or lateral spread) and leaf traits (e.g. specific leaf area and leaf dry matter content) captured the effect of shrub encroachment on multifunctionality with a relative high accuracy (r(2)  = 0.63). FD also improved the resistance of multifunctionality along the aridity gradient studied. Maintaining and enhancing FD in plant communities may help to buffer negative effects of ongoing global environmental change on dryland multifunctionality.

Predicting species' maximum dispersal distances from simple plant traits.

Many studies have shown plant species' dispersal distances to be strongly related to life-history traits, but how well different traits can predict dispersal distances is not yet known. We used cross-validation techniques and a global data set (576 plant species) to measure the predictive power of simple plant traits to estimate species' maximum dispersal distances. Including dispersal syndrome (wind, animal, ant, ballistic, and no special syndrome), growth form (tree, shrub, herb), seed mass, seed release height, and terminal velocity in different combinations as explanatory variables we constructed models to explain variation in measured maximum dispersal distances and evaluated their power to predict maximum dispersal distances. Predictions are more accurate, but also limited to a particular set of species, if data on more specific traits, such as terminal velocity, are available. The best model (R2 = 0.60) included dispersal syndrome, growth form, and terminal velocity as fixed effects. Reasonable predictions of maximum dispersal distance (R2 = 0.53) are also possible when using only the simplest and most commonly measured traits; dispersal syndrome and growth form together with species taxonomy data. We provide a function (dispeRsal) to be run in the software package R. This enables researchers to estimate maximum dispersal distances with confidence intervals for plant species using measured traits as predictors. Easily obtainable trait data, such as dispersal syndrome (inferred from seed morphology) and growth form, enable predictions to be made for a large number of species.

A negative heterogeneity-diversity relationship found in experimental grassland communities.

Recent meta-analyses and simulation studies have suggested that the relationship between soil resource heterogeneity and plant diversity (heterogeneity-diversity relationship; HDR) may be negative when heterogeneity occurs at small spatial scales. To explore different mechanisms that can explain a negative HDR, we conducted a mesocosm experiment combining a gradient of soil nutrient availability (low, medium, high) and scale of heterogeneity (homogeneous, large-scale heterogeneous, small-scale heterogeneous). The two heterogeneous treatments were created using chessboard combinations of low and high fertility patches, and had the same overall fertility as the homogeneous medium treatment. Soil patches were designed to be relatively larger (156 cm(2)) and smaller (39 cm(2)) than plant root extent. We found plant diversity was significantly lower in the small-scale heterogeneous treatment compared to the homogeneous treatment of the same fertility. Additionally, low fertility patches in the small-scale heterogeneous treatment had lower diversity than patches of the same size in the low fertility treatment. Shoot and root biomass were larger in the small-scale heterogeneous treatment than in the homogeneous treatment of the same fertility. Further, we found that soil resource heterogeneity may reduce diversity indirectly by increasing shoot biomass, thereby enhancing asymmetric competition for light resources. When soil resource heterogeneity occurs at small spatial scales it can lower plant diversity by increasing asymmetric competition belowground, since plants with large root systems can forage among patches and exploit soil resources. Additionally, small-scale soil heterogeneity may lower diversity indirectly, through increasing light competition, when nutrient uptake by competitive species increases shoot biomass production.

Microfragmentation concept explains non-positive environmental heterogeneity-diversity relationships.

Although recent studies have revealed that the relationship between diversity and environmental heterogeneity is not always positive, as classical niche theory predicts, scientists have had difficulty interpreting these results from an ecological perspective. We propose a new concept-microfragmentation-to explain how small-scale heterogeneity can have neutral or even negative effect on species diversity. We define microfragmentation as a community level process of splitting habitat into a more heterogeneous environment that can have non-positive effects on the diversity through habitat loss and subsequent isolation. We provide support for the microfragmentation concept with results from spatially explicit heterogeneity-diversity model simulations, in which varying sets of species (with different ratios of specialist and generalist species) were modeled at different levels of configurational heterogeneity (meaning that only the habitat structure was changed, not its composition). Our results indicate that environmental heterogeneity can affect community diversity in the same way as fragmentation at the landscape level. Although generalist species might not be seriously affected by microfragmentation, the persistence of specialist species can be seriously disturbed by small-scale patchiness. The microfragmentation concept provides new insight into community level diversity dynamics and can influence conservation and management strategies.