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.

Inverse relationship between species competitiveness and intraspecific trait variability may enable species coexistence in experimental seedling communities.

Theory suggests that intraspecific trait variability may promote species coexistence when competitively inferior species have higher intraspecific trait variability than their superior competitors. Here, we provide empirical evidence for this phenomenon in tree seedlings. We evaluated intraspecific variability and plastic response of ten traits in 6750 seedlings of ten species in a three-year greenhouse experiment. While we observed no relationship between intraspecific trait variability and species competitiveness in competition-free homogeneous environments, an inverse relationship emerged under interspecific competition and in spatially heterogeneous environments. We showed that this relationship is driven by the plastic response of the competitively inferior species: Compared to their competitively superior counterparts, they exhibited a greater increase in trait variability, particularly in fine-root traits, in response to competition, environmental heterogeneity and their combination. Our findings contribute to understanding how interspecific competition and intraspecific trait variability together structure plant communities.

Contrasting impacts of non-native and threatened species on morphological, life history, and phylogenetic diversity in bird assemblages.

Human activities have altered the species composition of assemblages through introductions and extinctions, but it remains unclear how those changes can affect the different facets of biodiversity. Here we assessed the impact of changes in species composition on taxonomic, functional, and phylogenetic diversity across 281 bird assemblages worldwide. To provide a more nuanced understanding of functional diversity, we distinguished morphological from life-history traits. We showed that shifts in species composition could trigger a global decline in avian biodiversity due to the high number of potential extinctions. Moreover, these extinctions were not random but unique in terms of function and phylogeny at the regional level. Our findings demonstrated that non-native species cannot compensate for these losses, as they are both morphologically and phylogenetically close to the native fauna. In the context of the ongoing biodiversity crisis, such alterations in the functional and phylogenetic structure of bird assemblages could heighten ecosystem vulnerability.

Functional trait trade-offs define plant population stability across different biomes.

Ecological theory posits that temporal stability patterns in plant populations are associated with differences in species' ecological strategies. However, empirical evidence is lacking about which traits, or trade-offs, underlie species stability, especially across different biomes. We compiled a worldwide collection of long-term permanent vegetation records (greater than 7000 plots from 78 datasets) from a large range of habitats which we combined with existing trait databases. We tested whether the observed inter-annual variability in species abundance (coefficient of variation) was related to multiple individual traits. We found that populations with greater leaf dry matter content and seed mass were more stable over time. Despite the variability explained by these traits being low, their effect was consistent across different datasets. Other traits played a significant, albeit weaker, role in species stability, and the inclusion of multi-variate axes or phylogeny did not substantially modify nor improve predictions. These results provide empirical evidence and highlight the relevance of specific ecological trade-offs, i.e. in different resource-use and dispersal strategies, for plant populations stability across multiple biomes. Further research is, however, necessary to integrate and evaluate the role of other specific traits, often not available in databases, and intraspecific trait variability in modulating species stability.

Changes in the functional diversity of modern bird species over the last million years.

Despite evidence of declining biosphere integrity, we currently lack understanding of how the functional diversity associated with changes in abundance among ecological communities has varied over time and before widespread human disturbances. We combine morphological, ecological, and life-history trait data for >260 extant bird species with genomic-based estimates of changing effective population size (Ne) to quantify demographic-based shifts in avian functional diversity over the past million years and under pre-anthropogenic climate warming. We show that functional diversity was relatively stable over this period, but underwent significant changes in some key areas of trait space due to changing species abundances. Our results suggest that patterns of population decline over the Pleistocene have been concentrated in particular regions of trait space associated with extreme reproductive strategies and low dispersal ability, consistent with an overall erosion of functional diversity. Further, species most sensitive to climate warming occupied a relatively narrow region of functional space, indicating that the largest potential population increases and decreases under climate change will occur among species with relatively similar trait sets. Overall, our results identify fluctuations in functional space of extant species over evolutionary timescales and represent the demographic-based vulnerability of different regions of functional space among these taxa. The integration of paleodemographic dynamics with functional trait data enhances our ability to quantify losses of biosphere integrity before anthropogenic disturbances and attribute contemporary biodiversity loss to different drivers over time.

Observed and dark diversity dynamics over millennial time scales: fast life-history traits linked to expansion lags of plants in northern Europe.

Global change drivers (e.g. climate and land use) affect the species and functional traits observed in a local site but also its dark diversity-the set of species and traits locally suitable but absent. Dark diversity links regional and local scales and, over time, reveals taxa under expansion lags by depicting the potential biodiversity that remains suitable but is absent locally. Since global change effects on biodiversity are both spatially and temporally scale dependent, examining long-term temporal dynamics in observed and dark diversity would be relevant to assessing and foreseeing biodiversity change. Here, we used sedimentary pollen data to examine how both taxonomic and functional observed and dark diversity changed over the past 14 500 years in northern Europe. We found that taxonomic and functional observed and dark diversity increased over time, especially after the Late Glacial and during the Late Holocene. However, dark diversity dynamics revealed expansion lags related to species' functional characteristics (dispersal limitation and stress intolerance) and an extensive functional redundancy when compared to taxa in observed diversity. We highlight that assessing observed and dark diversity dynamics is a promising tool to examine biodiversity change across spatial scales, its possible causes, and functional consequences.

The functional structure of plant communities drives soil functioning via changes in soil abiotic properties.

While biodiversity is expected to enhance multiple ecosystem functions (EFs), the different roles of multiple biodiversity dimensions remain difficult to disentangle without carefully designed experiments. We sowed plant communities with independent levels of functional (FD) and phylogenetic diversities (PD), combined with different levels of fertilization, to investigate their direct and indirect roles on multiple EFs, including plant-related EFs (plant biomass productivity, litter decomposability), soil fertility (organic carbon and nutrient pool variables), soil microbial activity (respiration and nutrient cycling), and an overall multifunctionality. We expected an increase in most EFs in communities with higher values of FD and/or PD via complementarity effects, but also the dominant plant types (using community weighted mean, CWM, independent of FD and PD) via selection effects on several EFs. The results showed strong direct effects of different dimensions of plant functional structure parameters on plant-related EFs, through either CWM or FD, with weak effects of PD. Fertilization had significant effects on one soil microbial activity and indirect effects on the other variables via changes in soil abiotic properties. Dominant plant types and FD showed only indirect effects on soil microbial activity, through litter decomposition and soil abiotic properties, highlighting the importance of cascading effects. This study shows the relevance of complementary dimensions of biodiversity for assessing both direct and cascading effects on multiple EFs.

Phenotypic dissimilarity index: Correcting for intra- and interindividual variability when quantifying phenotypic variation.

In trait-based ecology, phenotypic variation (PVar) is often quantified with measures expressing average differences between populations standardized in the range 0-1. However, these measures disregard the within-population trait variability. In addition, some of them cannot be partitioned between populations. These aspects can either alter their interpretation or limit their applicability. To overcome these problems, we propose a new measure, the phenotypic dissimilarity (PhD) index, to quantify PVar between populations in scenarios of varying within-population interindividual trait variability. PhD can also quantify within-population PVar while accounting for intraindividual trait variability. Using simulated and real data, we show that using the PhD index becomes important when the within-population trait variability is not negligible, as in all ecological studies. By accounting for within-population trait variability, the PhD index does not overestimate PVar across an environmental gradient compared to other estimators. Traits inherently vary within species. Accounting for such variability is essential to understanding species' phenotypic responses to environmental cues. The proposed PhD index will provide ecologists with a tool for quantifying PVar within species and compare it between species at different levels of biological organization. We provide an R function to calculate the PhD index.

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.

Competition-induced transgenerational plasticity influences competitive interactions and leaf decomposition of offspring.

Phenotypic plasticity, within and across generations (transgenerational plasticity), allows organisms and their progeny to adapt to the environment without modification of the underlying DNA. Recent findings suggest that epigenetic modifications are important mediators of such plasticity. However, empirical studies have, so far, mainly focused on plasticity in response to abiotic factors, overlooking the response to competition. We tested for within-generation and transgenerational phenotypic plasticity triggered by plant-plant competition intensity, and we tested whether it was mediated via DNA methylation, using the perennial, apomictic herb Taraxacum brevicorniculatum in four coordinated experiments. We then tested the consequences of transgenerational plasticity affecting competitive interactions of the offspring and ecosystem processes, such as decomposition. We found that, by promoting differences in DNA methylation, offspring of plants under stronger competition developed faster and presented more resource-conservative phenotypes. Further, these adjustments associated with less degradable leaves, which have the potential to reduce nutrient turnover and might, in turn, favour plants with more conservative traits. Greater parental competition enhanced competitive abilities of the offspring, by triggering adaptive phenotypic plasticity, and decreased offspring leaf decomposability. Our results suggest that competition-induced transgenerational effects could promote rapid adaptations and species coexistence and feed back on biodiversity assembly and nutrient cycling.

Diversity of parental environments increases phenotypic variation in Arabidopsis populations more than genetic diversity but similarly affects productivity.

BACKGROUND AND AIMS: The observed positive diversity effect on ecosystem functioning has rarely been assessed in terms of intraspecific trait variability within populations. Intraspecific phenotypic variability could stem both from underlying genetic diversity and from plasticity in response to environmental cues. The latter might derive from modifications to a plant's epigenome and potentially last multiple generations in response to previous environmental conditions. We experimentally disentangled the role of genetic diversity and diversity of parental environments on population productivity, resistance against environmental fluctuations and intraspecific phenotypic variation. METHODS: A glasshouse experiment was conducted in which different types of Arabidopsis thaliana populations were established: one population type with differing levels of genetic diversity and another type, genetically identical, but with varying diversity levels of the parental environments (parents grown in the same or different environments). The latter population type was further combined, or not, with experimental demethylation to reduce the potential epigenetic diversity produced by the diversity of parental environments. Furthermore, all populations were each grown under different environmental conditions (control, fertilization and waterlogging). Mortality, productivity and trait variability were measured in each population. KEY RESULTS: Parental environments triggered phenotypic modifications in the offspring, which translated into more functionally diverse populations when offspring from parents grown under different conditions were brought together in mixtures. In general, neither the increase in genetic diversity nor the increase in diversity of parental environments had a remarkable effect on productivity or resistance to environmental fluctuations. However, when the epigenetic variation was reduced via demethylation, mixtures were less productive than monocultures (i.e. negative net diversity effect), caused by the reduction of phenotypic differences between different parental origins. CONCLUSIONS: A diversity of environmental parental origins within a population could ameliorate the negative effect of competition between coexisting individuals by increasing intraspecific phenotypic variation. A diversity of parental environments could thus have comparable effects to genetic diversity. Disentangling the effect of genetic diversity and that of parental environments appears to be an important step in understanding the effect of intraspecific trait variability on coexistence and ecosystem functioning.

Diverging functional strategies but high sensitivity to an extreme drought in tropical dry forests.

Extreme drought events have negative effects on forest diversity and functioning. At the species level, however, these effects are still unclear, as species vary in their response to drought through specific functional trait combinations. We used long-term demographic records of 21,821 trees and extensive databases of traits to understand the responses of 338 tropical dry forests tree species to ENSO2015 , the driest event in decades in Northern South America. Functional differences between species were related to the hydraulic safety-efficiency trade-off, but unexpectedly, dominant species were characterised by high investment in leaf and wood tissues regardless of their leaf phenological habit. Despite broad functional trait combinations, tree mortality was more widespread in the functional space than tree growth, where less adapted species showed more negative net biomass balances. Our results suggest that if dry conditions increase in this ecosystem, ecological functionality and biomass gain would be reduced.

Synchrony matters more than species richness in plant community stability at a global scale.

The stability of ecological communities is critical for the stable provisioning of ecosystem services, such as food and forage production, carbon sequestration, and soil fertility. Greater biodiversity is expected to enhance stability across years by decreasing synchrony among species, but the drivers of stability in nature remain poorly resolved. Our analysis of time series from 79 datasets across the world showed that stability was associated more strongly with the degree of synchrony among dominant species than with species richness. The relatively weak influence of species richness is consistent with theory predicting that the effect of richness on stability weakens when synchrony is higher than expected under random fluctuations, which was the case in most communities. Land management, nutrient addition, and climate change treatments had relatively weak and varying effects on stability, modifying how species richness, synchrony, and stability interact. Our results demonstrate the prevalence of biotic drivers on ecosystem stability, with the potential for environmental drivers to alter the intricate relationship among richness, synchrony, and stability.

Temporal lags in observed and dark diversity in the Anthropocene.

Understanding biodiversity changes in the Anthropocene (e.g. due to climate and land-use change) is an urgent ecological issue. This important task is challenging because global change effects and species responses are dependent on the spatial scales considered. Furthermore, responses are often not immediate. However, both scale and time delay issues can be tackled when, at each study site, we consider dynamics in both observed and dark diversity. Dark diversity includes those species in the region that can potentially establish and thrive in the local sites' conditions but are currently locally absent. Effectively, dark diversity connects biodiversity at the study site to the regional scales and defines the site-specific species pool (observed and dark diversity together). With dark diversity, it is possible to decompose species gains and losses into two space-related components: one associated with local dynamics (species moving from observed to dark diversity and vice versa) and another related to gains and losses of site-specific species pool (species moving to and from the pool after regional immigration, regional extinction or change in local ecological conditions). Extinction debt and immigration credit are useful to understand dynamics in observed diversity, but delays might happen in species pool changes as well. In this opinion piece we suggest that considering both observed and dark diversity and their temporal dynamics provides a deeper understanding of biodiversity changes. Considering both observed and dark diversity creates opportunities to improve conservation by allowing to identify species that are likely to go regionally extinct as well as foreseeing which of the species that newly arrive to the region are more likely to colonize local sites. Finally, by considering temporal lags and species gains and losses in observed and dark diversity, we combine phenomena at both spatial and temporal scales, providing a novel tool to examine biodiversity change in the Anthropocene.

Author Correction: Contrasting functional structure of saproxylic beetle assemblages associated to different microhabitats.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Contrasting functional structure of saproxylic beetle assemblages associated to different microhabitats.

Saproxylic beetles depend on diverse microhabitats that are exploited by different species assemblages. We focused on anlyse the functional trait patterns and functional diversity components of two main assemblages that were collected with window traps (WTs) and hollow emergence traps (HETs) respectively, between three protected areas of the Iberian Peninsula. For that we measured phenological, physiological, morphological, and ecological traits. Results showed that the main microhabitats exploited by each assemblage (WT and HET) constrain most of the morphological traits and trophic guilds. In addition, relative elytra length and predator guild, together with adult activity period, responded to differences at the habitat level (among study areas). HET assemblages were less taxonomically diverse but more functionally diverse than those of WTs, enhancing the functional relevance of tree hollows. Additionally, niche filtering dominated WT assemblages, which were characterised by a narrower functional space and a higher redundancy. In contrast, in the HET assemblages the coexistence of functionally dissimilar species is driven by the niche heterogeneity. HET and WT assemblages differed in the functional space occupied by each within areas, but both assemblages reflected coincident patterns among areas that pointed to a reduction of functional space with management.