Forest disturbance increases functional diversity but decreases phylogenetic diversity of an arboreal tropical ant community.

Tropical rainforest trees host a diverse arthropod fauna that can be characterised by their functional diversity (FD) and phylogenetic diversity (PD). Human disturbance degrades tropical forests, often coinciding with species invasion and altered assembly that leads to a decrease in FD and PD. Tree canopies are thought to be particularly vulnerable, but rarely investigated. Here, we studied the effects of forest disturbance on an ecologically important invertebrate group, the ants, in a lowland rainforest in New Guinea. We compared an early successional disturbed plot (secondary forest) to an old-growth plot (primary forest) by exhaustively sampling their ant communities in a total of 852 trees. We expected that for each tree community (1) disturbance would decrease FD and PD in tree-dwelling ants, mediated through species invasion. (2) Disturbance would decrease ant trait variation due to a more homogeneous environment. (3) The main drivers behind these changes would be different contributions of true tree-nesting species and visiting species. We calculated FD and PD based on a species-level phylogeny and 10 ecomorphological traits. Furthermore, we assessed by data exclusion the influence of species, which were not nesting in individual trees (visitors) or only nesting species (nesters), and of non-native species on FD and PD. Primary forests had higher ant species richness and PD than secondary forest. However, we consistently found increased FD in secondary forest. This pattern was robust even if we decoupled functional and phylogenetic signals, or if non-native ant species were excluded from the data. Visitors did not contribute strongly to FD, but they increased PD and their community weighted trait means often varied from nesters. Moreover, all community-weighted trait means changed after forest disturbance. Our finding of contradictory FD and PD patterns highlights the importance of integrative measures of diversity. Our results indicate that the tree community trait diversity is not negatively affected, but possibly even enhanced by disturbance. Therefore, the functional diversity of arboreal ants is relatively robust when compared between old-growth and young trees. However, further study with higher plot-replication is necessary to solidify and generalise our findings.

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.

Floral and reproductive traits are an independent dimension within the plant economic spectrum of temperate central Europe.

Major dimensions of plant ecological strategies have been widely studied bringing forward the concept of 'economic spectra' of plants. Sexual reproductive traits, 'floral traits', have been largely neglected in this context, despite their strong link to fitness. Here, we aimed at integrating floral traits into the dimensionality of plant form and function so far dominated by vegetative traits. We used principal component analyses and constructed trait networks to assess the correlation structure of leaf, belowground, plant size-related, and floral traits. We studied forbs within two independent datasets; one compiled from central European trait databases and one sampled in the Austrian Alps. Floral traits defined the second dimension of trait variability within both datasets, while plant size determined the first dimension. Floral traits were largely independent from the leaf economic spectrum. Flower size, however, positively scaled with plant size and leaf size. Mating system was the most well-connected trait across modules of plant tissue/organ types. The independence of floral traits was consistent also after accounting for phylogenetic relationships between species. Floral traits explained a unique part of the variation in plant form and function and thus, likely play a distinctive ecological role within the whole plant economic spectrum.

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.

Alternative plant designs: consequences for community assembly and ecosystem functioning.

BACKGROUND: Alternative organism designs (i.e. the existence of distinct combinations of traits leading to the same function or performance) are a widespread phenomenon in nature and are considered an important mechanism driving the evolution and maintenance of species trait diversity. However, alternative designs are rarely considered when investigating assembly rules and species effects on ecosystem functioning, assuming that single trait trade-offs linearly affect species fitness and niche differentiation. SCOPE: Here, we first review the concept of alternative designs, and the empirical evidence in plants indicating the importance of the complex effects of multiple traits on fitness. We then discuss how the potential decoupling of single traits from performance and function of species can compromise our ability to detect the mechanisms responsible for species coexistence and the effects of species on ecosystems. Placing traits in the continuum of organism integration level (i.e. traits hierarchically structured ranging from organ-level traits to whole-organism traits) can help in choosing traits more directly related to performance and function. CONCLUSIONS: We conclude that alternative designs have important implications for the resulting trait patterning expected from different assembly processes. For instance, when only single trade-offs are considered, environmental filtering is expected to result in decreased functional diversity. Alternatively, it may result in increased functional diversity as an outcome of alternative strategies providing different solutions to local conditions and thus supporting coexistence. Additionally, alternative designs can result in higher stability of ecosystem functioning as species filtering due to environmental changes would not result in directional changes in (effect) trait values. Assessing the combined effects of multiple plant traits and their implications for plant functioning and functions will improve our mechanistic inferences about the functional significance of community trait patterning.

Stabilizing effects in temporal fluctuations: management, traits, and species richness in high-diversity communities.

The loss of biodiversity is thought to have adverse effects on multiple ecosystem functions, including the decline of community stability. Decreased diversity reduces the strength of the portfolio effect, a mechanism stabilizing community temporal fluctuations. Community stability is also expected to decrease with greater variability in individual species populations and with synchrony of their fluctuations. In semi-natural meadows, eutrophication is one of the most important drivers of diversity decline; it is expected to increase species fluctuations and synchrony among them, all effects leading to lower community stability. With a 16-year time series of biomass data from a temperate species-rich meadow with fertilization and removal of the dominant species, we assessed population biomass temporal (co)variation under different management types and competition intensity, and in relation to species functional traits and to species diversity. Whereas the effect of dominant removal was relatively small (with a tendency toward lower stability), fertilization markedly decreased community stability (i.e., increased coefficient of variation in the total biomass) and species diversity. On average, the fluctuations of individual populations were mutually independent, with a slight tendency toward synchrony in unfertilized plots, and a tendency toward compensatory dynamics in fertilized plots and no effects of removal. The marked decrease of synchrony with fertilization, contrary to the majority of the results reported previously, follows the predictions of increased compensatory dynamics with increased asymmetric competition for light in a more productive environment. Synchrony increased also with species functional similarity stressing the importance of shared ecological strategies in driving similar species responses to weather fluctuations. As expected, the decrease of temporal stability of total biomass was mainly related to the decrease of species richness, with its effect remaining significant also after accounting for fertilization. The weakening of the portfolio effect with species richness decline is a crucial driver of community destabilization. However, the positive effect of species richness on temporal stability of total biomass was not due to increased compensatory dynamics, since synchrony increased with species richness. This shows that the negative effect of eutrophication on community stability does not operate through increasing synchrony, but through the reduction of diversity.

Historical biome distribution and recent human disturbance shape the diversity of arbuscular mycorrhizal fungi.

The availability of global microbial diversity data, collected using standardized metabarcoding techniques, makes microorganisms promising models for investigating the role of regional and local factors in driving biodiversity. Here we modelled the global diversity of symbiotic arbuscular mycorrhizal (AM) fungi using currently available data on AM fungal molecular diversity (small subunit (SSU) ribosomal RNA (rRNA) gene sequences) in field samples. To differentiate between regional and local effects, we estimated species pools (sets of potentially suitable taxa) for each site, which are expected to reflect regional processes. We then calculated community completeness, an index showing the fraction of the species pool present, which is expected to reflect local processes. We found significant spatial variation, globally in species pool size, as well as in local and dark diversity (absent members of the species pool). Species pool size was larger close to areas containing tropical grasslands during the last glacial maximum, which are possible centres of diversification. Community completeness was greater in regions of high wilderness (remoteness from human disturbance). Local diversity was correlated with wilderness and current connectivity to mountain grasslands. Applying the species pool concept to symbiotic fungi facilitated a better understanding of how biodiversity can be jointly shaped by large-scale historical processes and recent human disturbance.

CLO-PLA: a database of clonal and bud-bank traits of the Central European flora.

This dataset presents comprehensive and easy-to-use information on 29 functional traits of clonal growth, bud banks, and lifespan of members of the Central European flora. The source data were compiled from a number of published sources (see the reference file) and the authors' own observations or studies. In total, 2,909 species are included (2,745 herbs and 164 woody species), out of which 1,532 (i.e., 52.7% of total) are classified as possessing clonal growth organs (1,480, i.e., 53.9%, if woody plants are excluded). This provides a unique, and largely unexplored, set of traits of clonal growth that can be used in studies on comparative plant ecology, plant evolution, community assembly, and ecosystem functioning across the large flora of Central Europe. It can be directly imported into a number of programs and packages that perform trait-based and phylogenetic analyses aimed to answer a variety of open and pressing ecological questions.

Applying the dark diversity concept to nature conservation.

Linking diversity to biological processes is central for developing informed and effective conservation decisions. Unfortunately, observable patterns provide only a proportion of the information necessary for fully understanding the mechanisms and processes acting on a particular population or community. We suggest conservation managers use the often overlooked information relative to species absences and pay particular attention to dark diversity (i.e., a set of species that are absent from a site but that could disperse to and establish there, in other words, the absent portion of a habitat-specific species pool). Together with existing ecological metrics, concepts, and conservation tools, dark diversity can be used to complement and further develop conservation prioritization and management decisions through an understanding of biodiversity relativized by its potential (i.e., its species pool). Furthermore, through a detailed understanding of the population, community, and functional dark diversity, the restoration potential of degraded habitats can be more rigorously assessed and so to the likelihood of successful species invasions. We suggest the application of the dark diversity concept is currently an underappreciated source of information that is valuable for conservation applications ranging from macroscale conservation prioritization to more locally scaled restoration ecology and the management of invasive species.

Trait assembly in grasslands depends on habitat history and spatial scale.

During the past century, grasslands in Europe have undergone marked changes in land-use, leading to a decline in plant diversity both at local and regional scales, thus possibly also affecting the mechanisms of species sorting into local communities. We studied plant species assembly in grasslands with differing habitat history and hypothesised that trait divergence prevails in historical grasslands due to niche differentiation and trait convergence prevails in more dynamic grasslands due to competitive exclusion and dispersal limitation. We tested these hypotheses in 35 grassland complexes in Estonia, containing neighbouring grassland habitats with different land-use histories: continuously managed open historical grassland, currently overgrown former grassland and young developing grassland. We assessed species assembly patterns in each grassland type for finer scale-a 2 × 2 m plot scale from a local community pool and for broader scale-a local community from the habitat species pool for that grassland stage and observed changes in trait means at finer scale. We found that grasslands with long management history are assembled differently from former grasslands or young developing grasslands. In historical grasslands, divergence or random patterns prevailed at finer scale species assembly while in former or developing grasslands, mostly convergence patterns prevailed. With increasing scale convergence patterns become more prevalent in all grassland types. We conclude that land-use history is an important factor to consider when assessing grassland functional trait assembly, particularly at small scales. Understanding the mechanisms behind species assembly and their relationship with land-use history is vital for habitat conservation and restoration.

Functional diversity through the mean trait dissimilarity: resolving shortcomings with existing paradigms and algorithms.

While an increasing number of indices for estimating the functional trait diversity of biological communities are being proposed, there is a growing demand by ecologists to clarify their actual implications and simplify index selection. Several key indices relate to mean trait dissimilarity between species within biological communities. Among them, the most widely used include (a) the mean species pairwise dissimilarity (MPD) and (b) the Rao quadratic entropy (and related indices). These indices are often regarded as redundant and promote the unsubstantiated yet widely held view that Rao is a form of MPD. Worryingly, existing R functions also do not always simplify the use and differentiation of these indices. In this paper, we show various distinctions between these two indices that warrant mathematical and biological consideration. We start by showing an existing form of MPD that considers species abundances and is different from Rao both mathematically and conceptually. We then show that the mathematical relationship between MPD and Rao can be presented simply as Rao = MPD × Simpson, where the Simpson diversity index is defined as 1 - dominance. We further show that this relationship is maintained for both species abundances and presence/absence. This evidence dismantles the paradigm that the Rao diversity is an abundance-weighted form of MPD and indicates that both indices can differ substantially at low species diversities. We discuss the different interpretations of trait diversity patterns in biological communities provided by Rao and MPD and then provide a simple R function, called "melodic," which avoids the unintended results that arise from existing mainstream functions.

Reinforcing loose foundation stones in trait-based plant ecology.

The promise of "trait-based" plant ecology is one of generalized prediction across organizational and spatial scales, independent of taxonomy. This promise is a major reason for the increased popularity of this approach. Here, we argue that some important foundational assumptions of trait-based ecology have not received sufficient empirical evaluation. We identify three such assumptions and, where possible, suggest methods of improvement: (i) traits are functional to the degree that they determine individual fitness, (ii) intraspecific variation in functional traits can be largely ignored, and (iii) functional traits show general predictive relationships to measurable environmental gradients.

Effects of long- and short-term management on the functional structure of meadows through species turnover and intraspecific trait variability.

The functional structures of communities respond to environmental changes by both species replacement (turnover) and within-species variation (intraspecific trait variability; ITV). Evidence is lacking on the relative importance of these two components, particularly in response to both short- and long-term environmental disturbance. We hypothesized that such short- and long-term perturbations would induce changes in community functional structure primarily via ITV and turnover, respectively. To test this we applied an experimental design across long-term mown and abandoned meadows, with each plot containing a further level of short-term management treatments: mowing, grazing and abandonment. Within each plot, species composition and trait values [height, shoot biomass, and specific leaf area (SLA)] were recorded on up to five individuals per species. Positive covariations between the contribution of species turnover and ITV occurred for height and shoot biomass in response to both short- and long-term management, indicating that species turnover and intraspecific adjustments selected for similar trait values. Positive covariations also occurred for SLA, but only in response to long-term management. The contributions of turnover and ITV changed depending on both the trait and management trajectory. As expected, communities responded to short-term disturbances mostly through changes in intraspecific trait variability, particularly for height and biomass. Interestingly, for SLA they responded to long-term disturbances by both species turnover and intraspecific adjustments. These findings highlight the importance of both ITV and species turnover in adjusting grassland functional trait response to environmental perturbation, and show that the response is trait specific and affected by disturbance regime history.

A global meta-analysis of the relative extent of intraspecific trait variation in plant communities.

Recent studies have shown that accounting for intraspecific trait variation (ITV) may better address major questions in community ecology. However, a general picture of the relative extent of ITV compared to interspecific trait variation in plant communities is still missing. Here, we conducted a meta-analysis of the relative extent of ITV within and among plant communities worldwide, using a data set encompassing 629 communities (plots) and 36 functional traits. Overall, ITV accounted for 25% of the total trait variation within communities and 32% of the total trait variation among communities on average. The relative extent of ITV tended to be greater for whole-plant (e.g. plant height) vs. organ-level traits and for leaf chemical (e.g. leaf N and P concentration) vs. leaf morphological (e.g. leaf area and thickness) traits. The relative amount of ITV decreased with increasing species richness and spatial extent, but did not vary with plant growth form or climate. These results highlight global patterns in the relative importance of ITV in plant communities, providing practical guidelines for when researchers should include ITV in trait-based community and ecosystem studies.

Functional species pool framework to test for biotic effects on community assembly.

Functional trait differences among species are increasingly used to infer the effects of biotic and abiotic processes on species coexistence. Commonly, the trait diversity observed within communities is compared to patterns simulated in randomly generated communities based on sampling within a region. The resulting patterns of trait convergence and divergence are assumed to reveal abiotic and biotic processes, respectively. However, biotic processes such as competition can produce both trait divergence and convergence, through either excluding similar species (niche differences, divergence) or excluding dissimilar species (weaker competitor exclusion, convergence). Hence, separating biotic and abiotic processes that can produce identical patterns of trait diversity, or even patterns that neutralize each other, is not feasible with previous methods. We propose an operational framework in which the functional trait dissimilarity within communities (FDcomm) is compared to the corresponding trait dissimilarity expected from the species pool (i.e., functional species pool diversity, FDpool). FDpool includes the set of potential species for a site delimited by the operating environmental and dispersal limitation filters. By applying these filters, the resulting pattern of trait diversity is consistent with biotic processes, i.e., trait divergence (FDcomm > FDpool) indicates niche differentiation, while trait convergence (FDcomm < FDpool) indicates weaker competitor exclusion. To illustrate this framework, with its potential application and constraints, we analyzed both simulated and field data. The functional species pool framework more consistently detected the simulated trait diversity patterns than previous approaches. In the field, using data from plant communities of typical Northern European habitats in Estonia, we found that both niche-based and weaker competitor exclusion influenced community assembly, depending on the traits and community considered. In both simulated and field data, we demonstrated that only by estimating the species pool of a site is it possible to differentiate the patterns of trait dissimilarity produced by operating biotic processes. The framework, which can be applied with both functional and phylogenetic diversity, enables a reinterpretation of community assembly processes. Solving the challenge of defining an appropriate reference species pool for a site can provide a better understanding of community assembly.

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.

Within-species variation of seed traits of dune engineering species across a European climatic gradient.

Within-species variation is a key component of biodiversity and linking it to climatic gradients may significantly improve our understanding of ecological processes. High variability can be expected in plant traits, but it is unclear to which extent it varies across populations under different climatic conditions. Here, we investigated seed trait variability and its environmental dependency across a latitudinal gradient of two widely distributed dune-engineering species (Thinopyrum junceum and Calamagrostis arenaria). Seed germination responses against temperature and seed mass were compared within and among six populations exposed to a gradient of temperature and precipitation regimes (Spiekeroog, DE; Bordeaux, FR; Valencia, ES; Cagliari, IT, Rome, IT; Venice, IT). Seed germination showed opposite trends in response to temperature experienced during emergence in both species: with some expectation, in populations exposed to severe winters, seed germination was warm-cued, whereas in populations from warm sites with dry summer, seed germination was cold-cued. In C. arenaria, variability in seed germination responses disappeared once the seed coat was incised. Seed mass from sites with low precipitation was smaller than that from sites with higher precipitation and was better explained by rainfall continentality than by aridity in summer. Within-population variability in seed germination accounted for 5 to 54%, while for seed mass it was lower than 40%. Seed trait variability can be considerable both within- and among-populations even at broad spatial scale. The variability may be hardly predictable since it only partially correlated with the analyzed climatic variables, and with expectation based on the climatic features of the seed site of origin. Considering seed traits variability in the analysis of ecological processes at both within- and among-population levels may help elucidate unclear patterns of species dynamics, thereby contributing to plan adequate measures to counteract biodiversity loss.

Cushions of Thylacospermum caespitosum (Caryophyllaceae) do not facilitate other plants under extreme altitude and dry conditions in the north-west Himalayas.

BACKGROUND: Cushion plants are commonly considered as keystone nurse species that ameliorate the harsh conditions they inhabit in alpine ecosystems, thus facilitating other species and increasing alpine plant biodiversity. A literature search resulted in 25 key studies showing overwhelming facilitative effects of different cushion plants and hypothesizing greater facilitation with increased environmental severity (i.e. higher altitude and/or lower rainfall). At the same time, emerging ecological theory alongside the cushion-specific literature suggests that facilitation might not always occur under extreme environmental conditions, and especially under high altitude and dryness. METHODS: To assess these hypotheses, possible nursing effects of Thylacospermum caespitosum (Caryophyllaceae) were examined at extremely high altitude (5900 m a.s.l.) and in dry conditions (precipitation <100 mm year(-1)) in Eastern Ladakh, Trans-Himalaya. This is, by far, the highest site, and the second driest, at which the effects of cushions have been studied so far. KEY RESULTS: In accordance with the theoretical predictions, no nursing effects of T. caespitosum on other alpine plants were detected. The number and abundance of species were greater outside cushions than within and on the edge of cushions. None of the 13 species detected was positively associated with cushions, while nine of them were negatively associated. Plant diversity increased with the size of the area sampled outside cushions, but no species-area relationship was found within cushions. CONCLUSIONS: The results support the emerging theoretical prediction of restricted facilitative effects under extreme combinations of cold and dryness, integrating these ideas in the context of the ecology of cushion plants. This evidence suggests that cases of missing strong facilitation are likely to be found in other extreme alpine conditions.

Reconciling trait based perspectives along a trait-integration continuum.

Trait based ecology has developed fast in the last decades, aiming to both explain mechanisms of community assembly, and predict patterns in nature, such as the effects of biodiversity shifts on key ecosystem processes. This body of work has stimulated the development of several conceptual frameworks and analytical methods, as well as the production of trait databases covering a growing number of taxa and organizational levels (from individuals to guilds). However, this breeding ground of novel concepts and tools currently lacks a general and coherent framework, under which functional traits can help ecologists organize their research aims, and serve as the common currency to unify several scientific disciplines. Specifically, we see a need to bridge the gaps between community ecology, ecosystem ecology, and evolutionary biology, in order to address the most pressing environmental issues of our time. To achieve this integration goal, we define a trait-integration continuum, which reconciles alternative trait definitions and approaches in ecology. This continuum outlines a coherent progression of biological scales, along which traits interact and hierarchically integrate from genetic information, to whole organism fitness-related traits, to trait syndromes and functional groups. Our conceptual scheme proposes that lower-level trait integration is closer to the inference of ecoevolutionary mechanisms determining population and community properties, whereas higher-level trait integration is most suited to the prediction of ecosystem processes. Within these two extremes, trait integration varies on a continuous scale, which relates directly to the inductive-deductive loop that should characterize the scientific method. With our proposed framework, we aim to facilitate scientists in contextualising their research based on the trait-integration levels that matter most to their specific goals. Explicitly acknowledging the existence of a trait-integration continuum is a promising way for framing the appropriate questions, thus obtaining reliable answers and results that are comparable across studies and disciplines.