Background:For several regions, ecologists and taxonomists have assembled information on phenotypes for almost all species of the megadiverse angiosperms. However, testing hypotheses on regional evolutionary and biogeographic history requires highly resolved and dated phylogenies covering the same taxa, which are often lacking at regional scales. Here, we filled this knowledge gap for one of the best-studied regional floras: the native angiosperms of the Netherlands. New information:We provide a molecular phylogenetic dataset and two time-calibrated trees (based on BEAST and MrBayes approaches) of the native angiosperm flora of the Netherlands, reconstructed from publicly available DNA sequences. The resulting phylogenies include 1178 species from the 2017 species list on SynBioSys NL, a national database that provides a curated checklist of native vascular plant species occurring in the Netherlands (also core to the flora of adjacent countries), and are provided in multiple formats (Newick, Nexus), along with alignments, BEAST and MrBayes input files, and metadata linking taxa to GenBank accession numbers. This dataset offers a phylogenetic framework that is based on molecular data covering more than 95% of the native species included in the checklist, reproducible and extremely well-resolved phylogeny (99.92% of nodes resolved in the maximum clade credibility tree; reflecting the absence of polytomies; 67.9% of nodes with posterior probability above 0.95 and 92.3% above 0.50) for researchers. These data will permit the signal of evolutionary history in patterns of biodiversity across the Netherlands, such as the age structure of habitat species pools or functional groups, where a focus on native species is essential. All resources are openly available via IDiv (https://doi.org/10.25829/idiv.3600-myr3t3) under a CC-BY license.
ABSTRACT Across the planet, angiosperm diversification has been shaped by palaeoclimates and biotic interactions, particularly with pollinators, leaving signatures in the angiosperm age structure. Regional floras represent biased samples of this global diversification shaped by dispersal, extinction and environmental filtering, yet it remains unclear whether floristic age structure reflects pollination strategies or their interaction with palaeoclimates. We constructed a 93% resolved phylogeny of all native 1178 angiosperms in the Netherlands and characterized age structure of eight pollination syndromes by standardized epoch‐specific lineage diversities (stELDs) quantifying the proportional increase in lineages per 10‐Myr interval. Overall, stELDs peaked under the coldest intervals and were lowest for intermediate‐temperature intervals (geological age alone had no effect). This age structure was strongest in wind‐pollinated species. Resolving insect pollination into six syndromes revealed strong heterogeneity: lepidoptera‐pollination was most strongly associated with warmest palaeotemperatures, bumblebee and wasp‐pollination under intermediate temperatures, and fly, lepidoptera and beetle‐pollination under coldest temperatures. Paleotemperature signatures correlated positively with contemporary seasonal flowering temperatures, indicating conserved thermal niches across ecological and macroevolutionary timescales. We suggest that temperate floras reflect both recent cold‐house diversification of wind‐ and certain insect‐pollinated lineages and earlier warm‐house radiations of lineages pollinated by older insect clades. Phylogenetic age structures permit detecting such patterns.
ABSTRACTInteractions between plants and insects have long fascinated scientists. While some plants rely on insects for pollination and seed dispersal, insects rely on plants for food or as a habitat. Despite extensive research investigating pair‐wise species interactions, few studies have characterized plant and insect communities simultaneously, making it unclear if diverse plant communities are generally associated with diverse insect communities. This work aims to better understand the historical and evolutionary relationships between plant and insect phylogenetic diversity (PD) on islands. We hypothesized that phylogenetically diverse plant communities (i.e., high PD) support diverse insect communities, with the relationship varying with island isolation, area, age, and latitude. Species lists for plants and insects were compiled from the published literature, and plant PD was calculated using ´standardized mean pairwise distance´ (SES.MPD) and ´standardized mean nearest taxon distance´ (SES.MNTD). For insects, PD was estimated using the number of genera, families, and orders. We found that plant diversity in evolutionary recent times (SES.MNTD) is associated with recent insect diversity (number of genera), but no relationship was found between plant and insect diversity across whole phylogenies (plant SES.MPD vs. number of insect families). Distant islands generally support high PD of plants (high SES.MPD and SES.MNTD) and insects (low number of genera). Plant and insect PD was generally high on small islands, except for plant SES.MPD revealing no relationship with island size. Insect PD was somewhat higher on young islands (low number of families), whereas there was no relationship between island age and plant PD. Plant SES.MPD was higher on high latitude islands, yet we did not find significant relationships between the latitude and the metrics of insect PD or plant SES.MNTD. These findings suggest that protecting high plant PD may also help conserve high insect PD, with a focus on small and distant islands as potential hotspots of phylogenetic diversity across multiple taxa.
Herbivory is ubiquitous. Despite being a potential driver of plant distribution and performance, herbivory remains largely undocumented. Some early attempts have been made to review, globally, how much leaf area is removed through insect feeding. Kozlov et al., in one of the most comprehensive reviews regarding global patterns of herbivory, have compiled published studies regarding foliar removal and sampled data on global herbivory levels using a standardized protocol. However, in the review by Kozlov et al., only 15 sampling sites, comprising 33 plant species, were evaluated in tropical areas around the globe. In Brazil, which ranks first in terms of plant biodiversity, with a total of 46,097 species, almost half (43%) being endemic, a single data point was sampled, covering only two plant species. In an attempt to increase knowledge regarding herbivory in tropical plant species and to provide the raw data needed to test general hypotheses related to plant-herbivore interactions across large spatial scales, we proposed a joint, collaborative network to evaluate tropical herbivory. This network allowed us to update and expand the data on insect herbivory in tropical and temperate plant species. Our data set, collected with a standardized protocol, covers 45 sampling sites from nine countries and includes leaf herbivory measurements of 57,239 leaves from 209 species of vascular plants belonging to 65 families from tropical and temperate regions. They expand previous data sets by including a total of 32 sampling sites from tropical areas around the globe, comprising 152 species, 146 of them being sampled in Brazil. For temperate areas, it includes 13 sampling sites, comprising 59 species. Thus, when compared to the most recent comprehensive review of insect herbivory (Kozlov et al.), our data set has increased the base of available data for the tropical plants more than 460% (from 33 to 152 species) and the Brazilian sampling was increased 7,300% (from 2 to 146 species). Data on precise levels of herbivory are presented for more than 57,000 leaves worldwide. There are no copyright restrictions. Please cite this paper when using the current data in publications; the authors request to be informed how the data is used in the publications.
Aridity is a critical driver of the diversity and composition of plant communities. However, how aridity influences the phylogenetic structure of functional groups (i.e. annual and perennial species) is far less understood than its effects on species richness. As perennials have to endure stressful conditions during the summer drought, as opposed to annuals that avoid it, they may be subjected to stronger environmental filtering. In contrast, annuals may be more susceptible to interannual climatic variability. Here we studied the phylogenetic structure of the annual and perennial components of understorey plant communities, along a regional aridity gradient in Mediterranean drylands. Specifically, we asked: (1) How do species richness (S) and phylogenetic structure (PS) of annuals and perennials in plant communities respond to aridity? (2) What is the contribution of other climatic and topo-edaphic variables in predicting S and PS for both components? (3) How does the taxonomic and phylogenetic turnover of annuals and perennials vary with spatial and environmental distances? We assessed annuals' and perennials' species richness, the phylogenetic structure at deep and shallow phylogenetic levels, and taxonomic and phylogenetic turnover along spatial and environmental distances. We found no relationship between annuals' richness and aridity, whereas perennials' richness showed a unimodal pattern. The phylogenetic structure of annuals and perennials showed contrasting responses to aridity and negatively correlated with topo-edaphic variables. We found phylogenetic clustering at intermediate-to-higher aridity levels for annuals, and at lower aridity levels for perennials. Both taxonomic and phylogenetic turnover in annuals and perennials correlated with the environmental distance rather than with spatial distance between communities, suggesting adaptation to local factors. Overall, our results show a decoupling in the response of the phylogenetic structure of annual and perennial components of plant communities to aridity in Mediterranean drylands. Our findings have significant implications for land management strategies under climate change.
Questions How do ecological and evolutionary processes affect the phylogenetic alpha and beta diversity of angiosperm tree assemblages in an evolutionarily young coastal environment? Location Coastal vegetation of the Brazilian Atlantic Forest Domain (restingas), spanning 20 degrees of latitude. Methods We calculated phylogenetic alpha and beta diversity at deep and shallow levels for angiosperm tree species in 136 circular sites of 10 km diameter (hereafter assemblages). The metrics we used for alpha diversity were the mean pairwise distance (MPD) and mean nearest taxon distance (MNTD), and mean pairwise distance separating species in two assemblages (D-pw) and mean nearest taxon distance separating species in two assemblages (D-nn) for beta diversity. We then investigated the relationship between phylogenetic diversity, historical (Quaternary) and current climatic variables, and edaphic conditions along the latitudinal gradient. Results We found that MPD increased with precipitation and latitude. MNTD increased with modern-day temperature, historical temperature instability and precipitation, and it was higher in more fertile and less saline soils. D-pw did not correlate either with geographic or environmental distances between assemblages. However, D-nn was strongly correlated with both geographic and environmental distances between assemblages. Conclusions The increase in MPD with precipitation and latitude suggests the presence of old Gondwanan lineages colonising restingas from refugia at higher latitudes. The increase in MNTD with modern-day temperature, historical temperature instability, precipitation, and it being higher in more fertile and less saline soils indicates that the distance between phylogenetic closest relatives tends to increase in assemblages less affected by environmental filtering. Low D-pw suggests the presence of widespread lineages across restingas, whereas high D-nn may indicate niche conservatism and dispersal limitation of more derived lineages. Our results offer insights into how ecological and evolutionary processes act to shape current patterns of biodiversity in geologically young environments.
Predicting the trajectory of ongoing diversity loss requires knowledge of historical development of community assemblages. Long-term data from palaeoecological investigations combined with key biodiversity measures in ecology such as taxonomic richness, functional diversity (FD), phylogenetic diversity (PD) and environmental factors expressed as Ellenberg indicator values (EIVs) could provide that knowledge. We explored the modern pollen–plant (moss polster pollen vs. surrounding vegetation) diversity relationships for herbaceous and woody taxa in calcareous fens from two different regions in Estonia, NE Europe. Associations of taxonomic richness, vegetation composition, FD (including functional alpha diversity and trait composition), PD and EIVs in modern pollen vs. plant data were studied with correlation analysis, Procrustes analysis and linear regression models. To test their potential use in palaeoreconstructions, diversity measures were applied on pollen data from Kanna spring fen reflecting fen vegetation development over the last nine millennia and diversity changes through time were studied using generalized additive models. Results showed significant pollen–plant richness correlations for herbaceous taxa at vegetation estimate scales up to 6 m radius and Procrustes analysis showed significant compositional associations at all plant estimate scales (up to 100 m). Woody taxa had no significant pollen–plant richness correlations but composition relationships were significant at plant estimate scales of 6–100 m. Traits that were best reflected by pollen data (both in terms of trait composition and functional alpha diversity) among woody and herbaceous taxa were seed number, clonality, SLA and LDMC. PD of herbaceous species was reflected by pollen data. Among the EIVs, Ellenberg L and T were significantly reflected by pollen data for both woody and herbaceous communities. Palaeoreconstruction from Kanna fen indicates that trends of woody taxa are mostly related to long-term changes in climate while diversity variables of herbaceous taxa closely follow autogenic processes within the fen. We suggest that pollen-based diversity estimates should be calculated separately for woody and herbaceous taxa as they clearly represent different spatial scales. Present study suggests that linking sedimentary pollen data with FD, PD and EIVs provides possibilities to examine long-term trends in community assembly and ecosystem processes that would be undetectable from traditional pollen diagrams.
AbstractLeaf-cutting ants are dominant herbivores in Neotropical rain forests, and their colony densities increase in disturbed habitats such as forest edges. However, while it is well-established that leaf-cutting ants profit from changes to the food-plant community, the phylogenetic dimension of this ant–plant interaction remains poorly understood in fragmented forests. We studied diet composition of Atta cephalotes in the edge and interior of Atlantic forest in north-east Brazil (8°30′S, 35°50′W). We applied phylogenetic signal analysis to investigate the diet across plant lineages and performed phylogenetic generalized linear models to analyse the diet in both habitats. We found a phylogenetic signal in diet and in leaf mechanical resistance, which means that A. cephalotes selects closely related food plants with less resistant leaves. Most preferred species belong to Malpighiales, Rubiaceae and Melastomataceae. We also found that irrespective of phylogeny, ants select food plants with less resistant leaves, both in edge and interior. However, ants choose more abundant plants only in edges. High abundance of optimal diet facilitates foraging in forest edges and explains why colony densities increase in disturbed habitats. Finally, by favouring or disfavouring specific clades, leaf-cutting ants contribute to changes in the phylogenetic structure of tropical rain forests, e.g. phylogenetic impoverishment.
Why can hosts coexist with conspecifics or phylogenetically proximate neighbours despite sharing specialist enemies? Do the hosts evolve increased enemy resistance? If so, does this have costs in terms of climatic-stress resistance, or in such neighbourhoods, does climatic-stress select for resistances that are multifunctional against climate and enemies? We studied oak (Quercus petraea) descendants from provenances of contrasting phylogenetic neighbourhoods and climates in a 25-year-old common garden. We found that descendants from conspecific or phylogenetically proximate neighbourhoods had the toughest leaves and fewest leaf miners, but no reduction in climatic-stress resistance. Descendants from such neighbourhoods under cold or dry climates had the highest flavonol and anthocyanin levels and the thickest leaves. Overall, populations facing phylogenetically proximate neighbours can rapidly evolve herbivore resistance, without cost to climatic-stress resistance, but possibly facilitating resistance against cold and drought via multifunctional traits. Microevolution might hence facilitate ecological coexistence of close relatives and thereby macroevolutionary conservatism of niches.
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.
Coexisting species may be evolutionarily proximate or distant, resulting in phylogenetically poor or rich communities. This variation is often considered to result from present assembly processes. We argue that, under certain conditions, deep-past processes might control the phylogenetic diversity of communities. First, deep-past effects involve macroevolutionary processes, such as diversification rate, niche conservatism, or dispersal, in the lineages that constitute communities. Second, deep-past processes in the respective region or in the habitat type play a role, for instance, through age, area, stability, or connectivity. Third, the deep past may affect communities via trophic interactions (i.e., communities of enemies or mutualists or communities of hosts). We suggest that deep-past effects can be identified in local communities by measuring phylogenetic diversity in different species pools. We also show how community phylogenetic diversity results in positive or negative eco-evolutionary feedback, and we identify present-day conservation challenges that may profit from a deep-time perspective.
Temperate calcareous grasslands are characterized by high levels of species richness at small spatial scales. Nevertheless, many species from a habitat‐specific regional species pool may be absent from local communities and represent the ‘dark diversity’ of these sites. Here we investigate dry calcareous grasslands in northern Europe to determine what proportion of the habitat‐specific species pool is realized at small scales (i.e. how the community completeness varies) and which mechanisms may be contributing to the relative sizes of the observed and dark diversity. We test whether the absence of particular species in potentially suitable grassland sites is a consequence of dispersal limitation and/or a low ability to tolerate stress (e.g. drought and grazing).We analysed a total of 1223 vegetation plots (1 × 1 m) from dry calcareous grasslands in Sweden, Estonia and western Russia. The species co‐occurrence approach was used to estimate the dark diversity for each plot. We calculated the maximum dispersal distance for each of the 291 species in our dataset by using simple plant traits (dispersal syndrome, growth form and seed characteristics). Large seed size was used as proxy for small seed number; tall plant height and low S‐strategy type scores were used to characterise low stress‐tolerance.Levels of small‐scale community completeness were relatively low (more species were absent than present) and varied between the grasslands in different geographic areas. Species in the dark diversity were generally characterized by shorter dispersal distances and greater seed weight (fewer seeds) than species in the observed diversity. Species within the dark diversity were generally taller and had a lower tolerance of stressful conditions.We conclude that, even if temperate grasslands have high levels of small‐scale plant diversity, the majority of potentially suitable species in the regional species pool may be absent as a result of dispersal limitation and low stress‐tolerance.
QuestionHow do pollen-based functional and phylogenetic diversity help to explain post-glacial vegetation change in relation to climate and human influence?LocationEstonia and Latvia, NE Europe.MethodsWe used a data set of 1062 pollen samples from 20 sites covering the last 14500yrs to estimate plant richness, evenness, functional and phylogenetic diversity (community-weighted mean and mean pair-wise distance). We adjusted existing functional and phylogenetic diversity measures for the pollen data and tested the methods with a simulation study. The simulations showed that species-based and pollen-based diversity estimates were all significantly positively correlated.ResultsThe Late Glacial (14500-11650cal. yr BP) and the mid-Holocene (8000-4000cal. yr BP) periods showed contrasting values for most of the diversity components, and several diversity estimates were strongly associated with climate. The cold climate during the Late Glacial led to high phylogenetic diversity, and relatively low functional diversity. Climate warming during the transition from the Late Glacial to the Holocene was followed by a decrease in phylogenetic diversity but an increase in functional diversity based on plant height and seed weight. Increasing human impact in the late Holocene was associated with an increase in plant richness and decreases in functional diversity based on plant height and seed weight and in phylogenetic diversity of herbs.ConclusionsPollen-based functional and phylogenetic diversity provide novel insights into post-glacial vegetation change and its drivers. Both functional and phylogenetic diversity were closely related to climatic conditions, suggesting that trait differences play an important role in long-term community response to climate change. Our results indicate that human impact during the last two millennia has influenced functional and phylogenetic diversity negatively by suppressing plants with certain traits (functional convergence) and giving advantage to plants from certain phylogenetic lineages. We see great potential in the further development of functional and phylogenetic diversity methods for pollen data.
Summary The subdiscipline of ‘community phylogenetics’ is rapidly growing and influencing thinking regarding community assembly. In particular, phylogenetic dispersion of co‐occurring species within a community is commonly used as a proxy to identify which community assembly processes may have structured a particular community: phylogenetic clustering as a proxy for abiotic assembly, that is habitat filtering, and phylogenetic overdispersion as a proxy for biotic assembly, notably competition. We challenge this approach by highlighting (typically) implicit assumptions that are, in reality, only weakly supported, including (i) phylogenetic dispersion reflects trait dispersion; (ii) a given ecological function can be performed only by a single trait state or combination of trait states; (iii) trait similarity causes enhanced competition; (iv) competition causes species exclusion; (v) communities are at equilibrium with processes of assembly having been completed; (vi) assembly through habitat filtering decreases in importance if assembly through competition increases, such that the relative balance of the two can be thus quantified by a single parameter; and (vii) observed phylogenetic dispersion is driven predominantly by local and present‐day processes. Moreover, technical sophistication of the phylogenetic‐patterns‐as‐proxy approach trades off against sophistication in alternative, potentially more pertinent approaches to directly observe or manipulate assembly processes. Despite concerns about using phylogenetic dispersion as a proxy for community assembly processes, we suggest there are underappreciated benefits of quantifying the phylogenetic structure of communities, including (i) understanding how coexistence leads to the macroevolutionary diversification of habitat lineage‐pools (i.e. phylogenetic‐patterns‐as‐result approach); and (ii) understanding the macroevolutionary contingency of habitat lineage‐pools and how it affects present‐day species coexistence in local communities (i.e. phylogenetic‐patterns‐as‐cause approach). We conclude that phylogenetic patterns may be little useful as proxy of community assembly. However, such patterns can prove useful to identify and test novel hypotheses on (i) how local coexistence may control macroevolution of the habitat lineage‐pool, for example through competition among close relatives triggering displacement and diversification of characters, and (ii) how macroevolution within the habitat lineage‐pool may control local coexistence of related species, for example through origin of close relatives that can potentially enter in competition.
Although experiments show a positive association between vascular plant and arbuscular mycorrhizal fungal (AMF) species richness, evidence from natural ecosystems is scarce. Furthermore, there is little knowledge about how AMF richness varies with belowground plant richness and biomass. We examined relationships among AMF richness, above- and belowground plant richness, and plant root and shoot biomass in a native North American grassland. Root-colonizing AMF richness and belowground plant richness were detected from the same bulk root samples by 454-sequencing of the AMF SSU rRNA and plant trnL genes. In total we detected 63 AMF taxa. Plant richness was 1.5 times greater belowground than aboveground. AMF richness was significantly positively correlated with plant species richness, and more strongly with below- than aboveground plant richness. Belowground plant richness was positively correlated with belowground plant biomass and total plant biomass, whereas aboveground plant richness was positively correlated only with belowground plant biomass. By contrast, AMF richness was negatively correlated with belowground and total plant biomass. Our results indicate that AMF richness and plant belowground richness are more strongly related with each other and with plant community biomass than with the plant aboveground richness measures that have been almost exclusively considered to date.
We used an extensive dataset (1220 vegetation plots of 1 m(2)) to study vegetation gradients and fine-scale plant diversity in dry calcareous grasslands (including alvar grasslands) in the Baltic Sea region. The study area covers the entire European distributional range of alvar habitats: Sweden (Oland, Gotland, Gotaland), Estonia (Saaremaa, Hiiumaa, north Estonia, west Estonia), and western Russia (Izhora, lzborsk). Fine-scale plant diversity was characterized by species richness and standardized phylogenetic diversity (comparing the observed mean pairwise phylogenetic distance (MPD) with MPD values from random communities). Ordination techniques (DCA) were used to characterize the main vegetation gradient. Variables describing local environment, climate, the biogeographic composition of the plant communities, and geographic location were related to fine-scale species richness and phylogenetic diversity using variation partitioning techniques and linear mixed models. The main vegetation gradient in the dry calcareous grasslands in the Baltic Sea region had a strong geographic component, was associated with soil depth, species' stress- and disturbance-tolerance and the age of the grassland habitat. Fine-scale phylogenetic diversity and species richness were negatively associated suggesting that these two diversity components are influenced by different sets of environmental and historical parameters. Fine-scale species richness was unimodally associated with the main vegetation gradient, and the highest levels of species richness were found under intermediate environmental (disturbance, light conditions and temperature) conditions where there was a mixture of species from different biogeographic regions. In contrast to species richness, fine-scale phylogenetic diversity was negatively associated with the main vegetation gradient. The highest phylogenetic diversity was found in the extremely thin-soiled alvar grasslands in Gotaland and on the Baltic islands (especially on Oland) where the high phylogenetic diversity is likely to be a reflection of a long history of continuous openness that has allowed time for the "collection" of phylogenetically different species within these unique habitats. (C) 2012 Elsevier B.V. All rights reserved.
Numerous studies have suggested a general relationship between the degree of host specialization and body size in herbivorous animals. In insects, smaller species are usually shown to be more specialized than larger-bodied ones. Various hypotheses have attempted to explain this pattern but rigorous proof of the body sizediet breadth relationship has been lacking, primarily because the scarceness of reliable phylogenetic information has precluded formal comparative analyses. Explicitly using phylogenetic information for a group of herbivores (geometrid moths) and their host plant range, we perform a comparative analysis to study the body sizediet breadth relationship. Considering several alternative measures of body size and diet breadth, our results convincingly demonstrate without previous methodological issuesa first for any taxona positive association between these traits, which has implications for evaluating various central aspects of the evolutionary ecology of herbivorous insects. We additionally demonstrate how the methods used in this study can be applied in assessing hypotheses to explain the body sizediet breadth relationship. By analyzing the relationship in tree-feeders alone and finding that the positive relationship remains, the result suggests that the body sizediet breadth relationship is not solely driven by the type of host plant that species feed on.
Question: There are contrasting opinions about how communities assemble along a productivity gradient, particularly in relation to competitive interactions. One view is that functionally similar, and closely related species, are less likely to co-exist (limiting similarity). Alternatively, competitive exclusion may act on dissimilar species because species bearing traits associated with low competitive ability are excluded (weaker competitor exclusion). We ask if patterns of functional and phylogenetic diversity are related to changes in species diversity in response to fertility manipulations.Location: Species-rich mesophytic grassland in Estonia.Methods: The grassland has been manipulated from 2002 to 2011 to increase (fertilizer addition) and decrease productivity (sucrose addition) in small-scale (50 cm x 50 cm) plots. We linked annual increases and decreases in species diversity to changes in functional and phylogenetic diversity. We used abundance- weighted mean pair-wise functional or phylogenetic distance of all possible species pairs.Results: We found convergence in four traits (plant height, leaf distribution, lateral spread, type of reproduction) and a decrease in mean functional and phylogenetic diversity, in support of weaker competitor exclusion or habitat filtering. There was less support for limiting similarity, with divergence found for two traits associated with decreasing species diversity (leaf distribution in the sucrose treatment and lateral spread in the fertilizer treatment).Conclusions: Our results support the view that competition can lead to the exclusion ofweaker competitors, rather than increasing functional and phylogenetic diversity, as expected from the principle of limiting similarity. However, multiple assembly processes, which are generally seen as mutually exclusive, are operating simultaneously, albeit on different traits and at different stages of community assembly.