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.

Variability in mycorrhizal status of plant species is much larger within than between plots in grassland and coastal habitats.

Community-level studies linking plant mycorrhizal status to environment usually do not account for within-plot mycorrhizal status variability; thus, patterns of plant mycorrhizal status diversity are largely unknown. Here, we assessed the relative importance of within- and between-plot variability components in mycorrhizal status and examined how plant mycorrhizal status diversity is related to soil nutrient availability. We hypothesised larger between-plot variability in mycorrhizal status and higher plant mycorrhizal status diversity in P-poor soils. To test these hypotheses, we used plant phylogenies, vegetation, soil and plant mycorrhizal status data from Czech semi-natural grasslands and Scottish coastal habitats. We divided plant mycorrhizal status diversity into divergence and evenness and tested their relations to soil P, K, Ca and Mg. Within-plot variability component of mycorrhizal status was always, on average, at least 2.2 times larger than between-plot variability in our datasets. Plant mycorrhizal status divergence was positively related to Ca (in both datasets) and Mg (only in grasslands and when accounting for phylogeny). In grasslands, the relationship between Mg and plant mycorrhizal status evenness was negative when accounting for phylogeny, while it was positive when not accounting for phylogeny. Plant mycorrhizal status diversity was not linked to P and its relation to K was inconsistent. Our results suggest that high Ca in the soil can promote coexistence of mycorrhizal, facultatively mycorrhizal and non-mycorrhizal plant species. We encourage future studies to also focus on within-plot variability in mycorrhizal status, because it appears to be highly relevant in herbaceous systems.

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.

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.

Functional trait effects on ecosystem stability: assembling the jigsaw puzzle.

Under global change, how biological diversity and ecosystem services are maintained in time is a fundamental question. Ecologists have long argued about multiple mechanisms by which local biodiversity might control the temporal stability of ecosystem properties. Accumulating theories and empirical evidence suggest that, together with different population and community parameters, these mechanisms largely operate through differences in functional traits among organisms. We review potential trait-stability mechanisms together with underlying tests and associated metrics. We identify various trait-based components, each accounting for different stability mechanisms, that contribute to buffering, or propagating, the effect of environmental fluctuations on ecosystem functioning. This comprehensive picture, obtained by combining different puzzle pieces of trait-stability effects, will guide future empirical and modeling investigations.

Are Reproductive Traits Related to Pollen Limitation in Plants? A Case Study from a Central European Meadow.

The deficiency of pollen grains for ovule fertilization can be the main factor limiting plant reproduction and fitness. Because of the ongoing global changes, such as biodiversity loss and landscape fragmentation, a better knowledge of the prevalence and predictability of pollen limitation is challenging within current ecological research. In our study we used pollen supplementation to evaluate pollen limitation (at the level of seed number and weight) in 22 plant species growing in a wet semi-natural meadow. We investigated the correlation between the pollen limitation index (PL) and floral traits associated with plant reproduction or pollinator foraging behavior. We recorded significant pollen limitation for approximately 41% of species (9 out of 22 surveyed). Seven species had a significant positive response in seed production and two species increased in seed weight after pollen supplementation. Considering traits, PL significantly decreased with the number of pollinator functional groups. The relationship of PL with other examined traits was not supported by our results. The causes of pollen limitation may vary among species with regard to (1) different reproductive strategies and life history, and/or (2) temporary changes in influence of biotic and abiotic factors at a site.

Linking Plant Functional Ecology to Island Biogeography.

The study of insular systems has a long history in ecology and biogeography. Island plants often differ remarkably from their noninsular counterparts, constituting excellent models for exploring eco-evolutionary processes. Trait-based approaches can help to answer important questions in island biogeography, yet plant trait patterns on islands remain understudied. We discuss three key hypotheses linking functional ecology to island biogeography: (i) plants in insular systems are characterized by distinct functional trait syndromes (compared with noninsular environments); (ii) these syndromes differ between true islands and terrestrial habitat islands; and (iii) island characteristics influence trait syndromes in a predictable manner. We are convinced that implementing trait-based comparative approaches would considerably further our understanding of plant ecology and evolution in insular systems.

Publisher Correction: Contrasting latitudinal patterns in phylogenetic diversity between woody and herbaceous communities.

A correction to this article has been published and is linked from the HTML and PDF versions of this paper. The error has been fixed in the paper.

Contrasting latitudinal patterns in phylogenetic diversity between woody and herbaceous communities.

Although many studies have shown that species richness decreases from low to high latitudes (the Latitudinal Diversity Gradient), little is known about the relationship between latitude and phylogenetic diversity. Here we examine global latitudinal patterns of phylogenetic diversity using a dataset of 459 woody and 589 herbaceous plant communities. We analysed the relationships between community phylogenetic diversity, latitude, biogeographic realm and vegetation type. Using the most recent global megaphylogeny for seed plants and the standardised effect sizes of the phylogenetic diversity metrics 'mean pairwise distance' (SESmpd) and 'mean nearest taxon distance' (SESmntd), we found that species were more closely-related at low latitudes in woody communities. In herbaceous communities, species were more closely-related at high latitudes than at intermediate latitudes, and the strength of this effect depended on biogeographic realm and vegetation type. Possible causes of this difference are contrasting patterns of speciation and dispersal. Most woody lineages evolved in the tropics, with many gymnosperms but few angiosperms adapting to high latitudes. In contrast, the recent evolution of herbaceous lineages such as grasses in young habitat types may drive coexistence of closely-related species at high latitudes. Our results show that high species richness commonly observed at low latitudes is not associated with high phylogenetic diversity.

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.

A Pragmatic Approach to Getting Published: 35 Tips for Early Career Researchers.

It is trite to say "publish or perish," yet many early career researchers are often at a loss on how to best get their work published. With strong competition and many manuscripts submitted, it is difficult to convince editors and reviewers to opt for acceptance. A pragmatic approach to publishing may increase one's odds of success. Here, we - a group of postdocs in the field of plant science - present specific recommendations for early career scientists on advanced levels. We cannot provide a recipe-like set of instructions with success guaranteed, but we come from a broad background in plant science, with experience publishing in a number of journals of varying topics and impact factors. We provide tips, tricks, and tools for collaboration, journal selection, and achieving acceptance.

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.

Evaluating Functional Diversity: Missing Trait Data and the Importance of Species Abundance Structure and Data Transformation.

Functional diversity (FD) is an important component of biodiversity that quantifies the difference in functional traits between organisms. However, FD studies are often limited by the availability of trait data and FD indices are sensitive to data gaps. The distribution of species abundance and trait data, and its transformation, may further affect the accuracy of indices when data is incomplete. Using an existing approach, we simulated the effects of missing trait data by gradually removing data from a plant, an ant and a bird community dataset (12, 59, and 8 plots containing 62, 297 and 238 species respectively). We ranked plots by FD values calculated from full datasets and then from our increasingly incomplete datasets and compared the ranking between the original and virtually reduced datasets to assess the accuracy of FD indices when used on datasets with increasingly missing data. Finally, we tested the accuracy of FD indices with and without data transformation, and the effect of missing trait data per plot or per the whole pool of species. FD indices became less accurate as the amount of missing data increased, with the loss of accuracy depending on the index. But, where transformation improved the normality of the trait data, FD values from incomplete datasets were more accurate than before transformation. The distribution of data and its transformation are therefore as important as data completeness and can even mitigate the effect of missing data. Since the effect of missing trait values pool-wise or plot-wise depends on the data distribution, the method should be decided case by case. Data distribution and data transformation should be given more careful consideration when designing, analysing and interpreting FD studies, especially where trait data are missing. To this end, we provide the R package "traitor" to facilitate assessments of missing trait data.

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.

Mycorrhizas in the Central European flora: relationships with plant life history traits and ecology.

Plant traits have been widely used to characterize different aspects of the ecology of plant species. Despite its wide distribution and its proven significance at the level of individuals, communities, and populations, the ability to form mycorrhizal associations has been largely neglected in these studies so far. Analyzing plant traits associated with the occurrence of mycorrhizas in plants can therefore enhance our understanding of plant strategies and distributions. Using a comparative approach, we tested for associations between mycorrhizal status and habitat characteristics, life history traits, and plant distribution patterns in 1752 species of the German flora (a major part of the Central European flora). Data were analyzed using log-linear models or generalized linear models, both accounting for phylogenetic relationships. Obligatorily mycorrhizal (OM) species tended to be positively associated with higher temperature, drier habitats, and higher pH; and negatively associated with moist, acidic, and fertile soils. Competitive species were more frequently OM, and stress tolerators were non-mycorrhizal (NM), while ruderal species did not show any preference. Facultatively mycorrhizal (FM) species showed the widest geographic and ecological amplitude. Indigenous species were more frequently FM and neophytes (recent aliens) more frequently OM than expected. FM species differed markedly from OM and NM species in almost all analyzed traits. Specifically, they showed a wider geographic distribution and ecological niche. Our study of the relationships between mycorrhizal status and other plant traits provides a comprehensive test of existing hypotheses and reveals novel patterns. The clear distinction between FM and OM + NM species in terms of their ecology opens up a new field of research in plant-mycorrhizal ecology.

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.

Ecological assembly rules in plant communities--approaches, patterns and prospects.

Understanding how communities of living organisms assemble has been a central question in ecology since the early days of the discipline. Disentangling the different processes involved in community assembly is not only interesting in itself but also crucial for an understanding of how communities will behave under future environmental scenarios. The traditional concept of assembly rules reflects the notion that species do not co-occur randomly but are restricted in their co-occurrence by interspecific competition. This concept can be redefined in a more general framework where the co-occurrence of species is a product of chance, historical patterns of speciation and migration, dispersal, abiotic environmental factors, and biotic interactions, with none of these processes being mutually exclusive. Here we present a survey and meta-analyses of 59 papers that compare observed patterns in plant communities with null models simulating random patterns of species assembly. According to the type of data under study and the different methods that are applied to detect community assembly, we distinguish four main types of approach in the published literature: species co-occurrence, niche limitation, guild proportionality and limiting similarity. Results from our meta-analyses suggest that non-random co-occurrence of plant species is not a widespread phenomenon. However, whether this finding reflects the individualistic nature of plant communities or is caused by methodological shortcomings associated with the studies considered cannot be discerned from the available metadata. We advocate that more thorough surveys be conducted using a set of standardized methods to test for the existence of assembly rules in data sets spanning larger biological and geographical scales than have been considered until now. We underpin this general advice with guidelines that should be considered in future assembly rules research. This will enable us to draw more accurate and general conclusions about the non-random aspect of assembly in plant communities.