Mean conditions predict salt marsh plant community diversity and stability better than environmental variability

Environmental variability and the frequency of extreme events are predicted to increase in future climate scenarios; however, the role of fluctuations in shaping community composition, diversity and stability is not well understood. Identifying current patterns of association between measures of community stability and climatic means and variability will help elucidate the ways in which altered variability and mean conditions may change communities in the future. Salt marshes provide essential ecosystem services and are increasingly threatened by sea‐level rise, land‐use change, eutrophication and predator loss, yet the effects of temporal environmental variation on salt marshes remain unknown. We synthesized long‐term plant community monitoring data from 11 sites on both coasts of the United States. We used an information‐theoretic approach and linear models to determine the associations among long‐term mean conditions, interannual environmental variability, and plant community stability and diversity. We found that salt marsh community stability and diversity were more strongly related to long‐term means of temperature and precipitation than to interannual variation. Warm and wet environments had fewer species and less turnover among years. Our results suggest that communities in cool, dry environments may be more resilient to climate warming due to greater species richness and turnover. Mean conditions are sufficient to predict contemporary patterns of salt marsh plant community dynamics, but environmental variability may have stronger impacts as it increases with climate change.     

Response of marine communities to local temperature changes

As global climate change and variability drive shifts in species’ distributions, ecological communities are being reorganized. One approach to understand community change in response to climate change has been to characterize communities by a collective thermal preference, or community temperature index (CTI), and then to compare changes in CTI with changes in temperature. However, important questions remain about whether and how responsive communities are to changes in their local thermal environments. We used CTI to analyze changes in 160 marine assemblages (fish and invertebrates) across the rapidly‐changing Northeast U.S. Continental Shelf Large Marine Ecosystem and calculated expected community change based on historical relationships between species presence and temperature from a separate training dataset. We then compared interannual and long‐term temperature changes with expected community responses and observed community responses over both temporal scales. For these marine communities, we found that community composition as well as composition changes through time could be explained by species associations with bottom temperature. Individual species had non‐linear responses to changes in temperature, and these nonlinearities scaled up to a nonlinear relationship between CTI and temperature. On average, CTI increased by 0.36°C (95% CI: 0.34–0.38°C) for every 1°C increase in bottom temperature, but the relationship between CTI and temperature also depended on community composition. In addition, communities responded more strongly to interannual variation than to long‐term trends in temperature. We recommend that future research into climate‐driven community change accounts for nonlinear responses and examines ecological responses across a range of temporal and geographical scales.     

Environmental heterogeneity and biotic interactions mediate climate impacts on tropical forest regeneration

Predicting the fate of tropical forests under a changing climate requires understanding species responses to climatic variability and extremes. Seedlings may be particularly vulnerable to climatic stress given low stored resources and undeveloped roots; they also portend the potential effects of climate change on future forest composition. Here we use data for ca. 50,000 tropical seedlings representing 25 woody species to assess (i) the effects of interannual variation in rainfall and solar radiation between 2007 and 2016 on seedling survival over 9 years in a subtropical forest; and (ii) how spatial heterogeneity in three environmental factors—soil moisture, understory light, and conspecific neighborhood density—modulate these responses. Community‐wide seedling survival was not sensitive to interannual rainfall variability but interspecific variation in these responses was large, overwhelming the average community response. In contrast, community‐wide responses to solar radiation were predominantly positive. Spatial heterogeneity in soil moisture and conspecific density were the predominant and most consistent drivers of seedling survival, with the majority of species exhibiting greater survival at low conspecific densities and positive or nonlinear responses to soil moisture. This environmental heterogeneity modulated impacts of rainfall and solar radiation. Negative conspecific effects were amplified during rainy years and at dry sites, whereas the positive effects of radiation on survival were more pronounced for seedlings existing at high understory light levels. These results demonstrate that environmental heterogeneity is not only the main driver of seedling survival in this forest but also plays a central role in buffering or exacerbating impacts of climate fluctuations on forest regeneration. Since seedlings represent a key bottleneck in the demographic cycle of trees, efforts to predict the long‐term effects of a changing climate on tropical forests must take into account this environmental heterogeneity and how its effects on regeneration dynamics play out in long‐term stand dynamics.     

Movement‐mediated community assembly and coexistence

Organismal movement is ubiquitous and facilitates important ecological mechanisms that drive community and metacommunity composition and hence biodiversity. In most existing ecological theories and models in biodiversity research, movement is represented simplistically, ignoring the behavioural basis of movement and consequently the variation in behaviour at species and individual levels. However, as human endeavours modify climate and land use, the behavioural processes of organisms in response to these changes, including movement, become critical to understanding the resulting biodiversity loss. Here, we draw together research from different subdisciplines in ecology to understand the impact of individual‐level movement processes on community‐level patterns in species composition and coexistence. We join the movement ecology framework with the key concepts from metacommunity theory, community assembly and modern coexistence theory using the idea of micro–macro links, where various aspects of emergent movement behaviour scale up to local and regional patterns in species mobility and mobile‐link‐generated patterns in abiotic and biotic environmental conditions. These in turn influence both individual movement and, at ecological timescales, mechanisms such as dispersal limitation, environmental filtering, and niche partitioning. We conclude by highlighting challenges to and promising future avenues for data generation, data analysis and complementary modelling approaches and provide a brief outlook on how a new behaviour‐based view on movement becomes important in understanding the responses of communities under ongoing environmental change.     

Biodiversity and persistence of ecological communities in variable environments

Recent analyses of climate data indicate that the intensity and frequency of different weather extremes have increased. Such increased environmental variability may lead to increased species extinction rates and hence have important consequences for the long-term persistence of ecological communities. Here we use model communities in order to investigate the relationship between species richness and community persistence in a fluctuating environment. We model two scenarios: (1) correlated species responses to environmental fluctuations and (2) uncorrelated (independent) species responses. We quantify the risk and extent of species extinctions using the so-called community viability analysis. It is shown that species-rich communities are more sensitive to environmental stochasticity than species-poor communities. Specifically, per species risk of extinction is higher in species-rich communities than in species-poor ones. Moreover, for a given species richness, communities with uncorrelated species responses to environmental variation run a considerable higher risk of losing a fixed proportion of species compared with communities with correlated species responses. We discuss the compatibility of these results with the ecological insurance hypothesis.     

Climate extremes drive changes in functional community structure

The response of communities to climate extremes can be quite variable. Much of this variation has been attributed to differences in community‐specific functional trait diversity, as well as community composition. Yet, few if any studies have explicitly tested the response of the functional trait structure of communities following climate extremes (CEs). Recently in South Florida, two independent, but sequential potential CEs took place, a 2010 cold front, followed by a 2011 drought, both of which had profound impacts on a subtropical estuarine fish community. These CEs provided an opportunity to test whether the structure of South Florida fish communities following each extreme was a result of species‐specific differences in functional traits. From historical temperature (1927–2012) and freshwater inflows records into the estuary (1955–2012), we determined that the cold front was a statistically extreme disturbance, while the drought was not, but rather a decadal rare disturbance. The two disturbances predictably affected different parts of functional community structure and thus different component species. The cold front virtually eliminated tropical species, including large‐bodied snook, mojarra species, nonnative cichlids, and striped mullet, while having little affect on temperate fishes. Likewise, the drought severely impacted freshwater fishes including Florida gar, bowfin, and two centrarchids, with little effect on euryhaline species. Our findings illustrate the ability of this approach to predict and detect both the filtering effects of different types of disturbances and the implications of the resulting changes in community structure. Further, we highlight the value of this approach to developing predictive frameworks for better understanding community responses to global change.     

Hierarchical regulation of pelagic invertebrates in lakes of the northern Great Plains: a novel model for interdecadal effects of future climate change on lakes

Endorheic lakes of the northern Great Plains encompass a wide range of environmental parameters (e.g., salinity, pH, DOC, Ca, nutrients, depth) that vary 1000‐fold among sites and through the past 2000 years due to variation in basin hydrology and evaporative forcing. However, while many environmental parameters are known to individually influence zooplankton diversity and taxonomic composition, relatively little is known of the hierarchical relationships among potential controls or of how regulatory mechanisms may change in response to climate variation on diverse scales. To address these issues, we surveyed 70 lakes within a 100 000 km² prairie region to simulate the magnitude of environmental change expected to occur over 100–1000 years and to quantify the unique and interactive effects of diverse environmental parameters in regulating pelagic invertebrate community structure at that scale. Multivariate analyses showed that salinity was the principal correlate of changes in invertebrate composition among lakes, with a sequential loss of taxa between salinities of 4 and 50 g total dissolved solids L^?1 until one to two species predominated in highly saline systems. In contrast, changes in the concentrations of Ca²⁺ and other mineral nutrients exerted secondary controls of invertebrate assemblages independent of salinity, whereas lake depth provided a tertiary regulatory mechanism structuring species composition. In contrast to these large‐scale hierarchical patterns, seasonal surveys (May, July, September) of a subset of 21 lakes in each of 2003–2005 revealed that annual meteorological variation had no measurable effect on pelagic invertebrates, despite large differences in temperature, precipitation, and evaporation arising from regional droughts. Together these findings show that pelagic invertebrate communities in saline lakes are resilient to interannual variability in climate, but suggest that lakes of the northern Great Plains may provide a sensitive model to forecast centennial effects of future climate change.     

Interannual variability in species composition explained as seasonally entrained chaos

The species composition of plankton, insect and annual plant communities may vary markedly from year to year. Such interannual variability is usually thought to be driven by year-to-year variation in weather conditions. Here we examine an alternative explanation. We studied the effects of regular seasonal forcing on a multi-species predator–prey model consisting of phytoplankton and zooplankton species. The model predicts that interannual variability in species composition can easily arise without interannual variability in external conditions. Seasonal forcing increased the probability of chaos in our model communities, but squeezed these irregular species dynamics within the seasonal cycle. As a result, the population dynamics had a peculiar character. Consistent with long-term time series of natural plankton communities, seasonal variation led to a distinct seasonal succession of species, yet the species composition varied from year to year in an irregular fashion. Our results suggest that interannual variability in species composition is an intrinsic property of multi-species communities in seasonal environments.     

Multiscale modeling of spring phenology across Deciduous Forests in the Eastern United States

Phenological events, such as bud burst, are strongly linked to ecosystem processes in temperate deciduous forests. However, the exact nature and magnitude of how seasonal and interannual variation in air temperatures influence phenology is poorly understood, and model‐based phenology representations fail to capture local‐ to regional‐scale variability arising from differences in species composition. In this paper, we use a combination of surface meteorological data, species composition maps, remote sensing, and ground‐based observations to estimate models that better represent how community‐level species composition affects the phenological response of deciduous broadleaf forests to climate forcing at spatial scales that are typically used in ecosystem models. Using time series of canopy greenness from repeat digital photography, citizen science data from the USA National Phenology Network, and satellite remote sensing‐based observations of phenology, we estimated and tested models that predict the timing of spring leaf emergence across five different deciduous broadleaf forest types in the eastern United States. Specifically, we evaluated two different approaches: (i) using species‐specific models in combination with species composition information to ‘upscale’ model predictions and (ii) using repeat digital photography of forest canopies that observe and integrate the phenological behavior of multiple representative species at each camera site to calibrate a single model for all deciduous broadleaf forests. Our results demonstrate variability in cumulative forcing requirements and photoperiod cues across species and forest types, and show how community composition influences phenological dynamics over large areas. At the same time, the response of different species to spatial and interannual variation in weather is, under the current climate regime, sufficiently similar that the generic deciduous forest model based on repeat digital photography performed comparably to the upscaled species‐specific models. More generally, results from this analysis demonstrate how in situ observation networks and remote sensing data can be used to synergistically calibrate and assess regional parameterizations of phenology in models.     

Effects of climate extremes on the terrestrial carbon cycle: concepts,processes and potential future impacts

Extreme droughts, heat waves, frosts, precipitation, wind storms and other climate extremes may impact the structure, composition and functioning of terrestrial ecosystems, and thus carbon cycling and its feedbacks to the climate system. Yet, the interconnected avenues through which climate extremes drive ecological and physiological processes and alter the carbon balance are poorly understood. Here, we review the literature on carbon cycle relevant responses of ecosystems to extreme climatic events. Given that impacts of climate extremes are considered disturbances, we assume the respective general disturbance‐induced mechanisms and processes to also operate in an extreme context. The paucity of well‐defined studies currently renders a quantitative meta‐analysis impossible, but permits us to develop a deductive framework for identifying the main mechanisms (and coupling thereof) through which climate extremes may act on the carbon cycle. We find that ecosystem responses can exceed the duration of the climate impacts via lagged effects on the carbon cycle. The expected regional impacts of future climate extremes will depend on changes in the probability and severity of their occurrence, on the compound effects and timing of different climate extremes, and on the vulnerability of each land‐cover type modulated by management. Although processes and sensitivities differ among biomes, based on expert opinion, we expect forests to exhibit the largest net effect of extremes due to their large carbon pools and fluxes, potentially large indirect and lagged impacts, and long recovery time to regain previous stocks. At the global scale, we presume that droughts have the strongest and most widespread effects on terrestrial carbon cycling. Comparing impacts of climate extremes identified via remote sensing vs. ground‐based observational case studies reveals that many regions in the (sub‐)tropics are understudied. Hence, regional investigations are needed to allow a global upscaling of the impacts of climate extremes on global carbon–climate feedbacks.     

Dynamic macroecology and the future for biodiversity

Reliable projections of climate‐change impacts on biodiversity are vital in formulating conservation and management strategies that best retain biodiversity into the future. While recent modelling has focussed largely on individual species, macroecology has the potential to add significant value to these efforts, by incorporating important community‐level constraints and processes. Here we show how a new dynamic macroecological approach can project climate‐change impacts collectively across all species in a diverse taxonomic group, overcoming shortfalls in our knowledge of biodiversity, while incorporating the key processes of dispersal and community assembly. Our approach applies a recently published technique (DynamicFOAM) to predict the present composition of every community, which form the initial conditions for a new metacommunity model (M‐SET) that projects changes in composition over time, under specified climate and habitat scenarios. Applying this approach at fine resolution to plant biodiversity in Tasmania (2,051 species; 1,157,587 communities), we project high average turnover in community composition from 2010 to 2100 (mean Sorensen's dissimilarity = 0.71 (±7.0 × 10^?5)), with major reductions in species richness (32.9 (±0.02) species lost per community) and no plant species benefitting from climate change in the long term. We also demonstrate how our modelling approach can identify habitat likely to be of high value for retaining rare and poorly reserved species under climate change. Our analyses highlight the potential value of this dynamic macroecological approach, that incorporates key ecological processes in projecting climate change impacts for all species simultaneously and uses simple macroecological inputs that can be derived even for highly diverse and poorly studied taxa.     

Floristic patterns and plant traits of Mediterranean communities in fragmented habitats

Aim To contrast floristic spatial patterns and the importance of habitat fragmentation in two plant communities (grassland and scrubland) in the context of ecological succession. We ask whether plant assemblages are affected by habitat fragmentation and, if so, at what spatial scale? Does the relative importance of the niche differentiation and dispersal‐limitation mechanisms change throughout secondary succession? Is the dispersal‐limitation mechanism related to plant functional traits? Location A Mediterranean region, the massif of Albera (Spain). Methods Using a SPOT satellite image to describe the landscape, we tested the effect of habitat fragmentation on species composition, determining the spatial scale of the assemblage response. We then assessed the relative importance of dispersal‐related factors (habitat fragmentation and geographical distance) and environmental constraints (climate‐related variables) influencing species similarity. We tested the association between dispersal‐related factors and plant traits (dispersal mode and life form). Results In both community types, plant composition was partially affected by the surrounding vegetation. In scrublands, animal‐dispersed and woody plants were abundant in landscapes dominated by closed forests, whereas wind‐dispersed annual herbs were poorly represented in those landscapes. Scrubby assemblages were more dependent on geographical distance, habitat fragmentation and climate conditions (temperature, rainfall and solar radiation); grasslands were described only by habitat fragmentation and rainfall. Plant traits did not explain variation in spatial structuring of assemblages. Main conclusions Plant establishment in early Mediterranean communities may be driven primarily by migration from neighbouring established communities, whereas the importance of habitat specialization and community drift increases over time. Plant life forms and dispersal modes did not explain the spatial variation of species distribution, but species richness within the community with differing plant traits was affected by habitat patchiness.     

Big changes to a small bay: introduced species and long-term compositional shifts to the fouling community of Morro Bay (CA)

Invasive species are cited as being a threat to communities and ecosystems worldwide, yet few studies have demonstrated invader impacts at these scales. Lack of historic data makes capturing large-scale community shifts problematic. We assessed long-term changes in invertebrate composition to the fouling community of a small estuary with relatively little boat traffic and no ballast water input (Morro Bay, CA). We also compared relative invasiveness of Morro Bay to international harbors (San Francisco Bay and Los Angeles/San Diego harbors). While the proportion of introduced species has not significantly changed from historic records, introduced species now occupy 86.00 % of the primary substrate. Other community shifts include; (1) a state shift to an invasive bryozoan (Watersipora subtorquata) dominated community, (2) a decrease in Mollusc richness and, (3) substantial shifts in abundance of certain species. Compared to larger more actively used harbors, Morro Bay has proportionally fewer introduced species (12.00 %) than San Francisco Bay (50.79 %) or Los Angeles/San Diego Harbors (26.23 %). Our study documents changes to a small relatively isolated estuary with little boat traffic and no ballast water input. We discuss the potential role of invasive species and other natural and anthropogenic factors as drivers of these community wide shifts. Specifically, we discuss how reintroduction of the Southern Sea Otter (Enhydra lutris nereis), an increase in sea stars (Pisaster spp.), climate change and interaction amongst potential drivers support the patterns of shifts in the Morro Bay community.     

Disentangling how climate change can affect an aquatic food web by combining multiple experimental approaches

Predicting the biological effects of climate change presents major challenges due to the interplay of potential biotic and abiotic mechanisms. Climate change can create unexpected outcomes by altering species interactions, and uncertainty over the ability of species to develop in situ tolerance or track environmental change further hampers meaningful predictions. As multiple climatic variables shift in concert, their potential interactions further complicate ecosystem responses. Despite awareness of these complexities, we still lack controlled experiments that manipulate multiple climatic stressors, species interactions, and prior exposure of species to future climatic conditions. Particularly studies that address how changes in water availability interact with other climatic stressors to affect aquatic ecosystems are still rare. Using aquatic insect communities of Neotropical tank bromeliads, we combined controlled manipulations of drought length and species interactions with a space‐for‐time transplant (lower elevations represent future climate) and a common garden approach. Manipulating drought length and experiment elevation revealed that adverse effects of drought were amplified at the warmer location, highlighting the potential of climatic stressors to synergistically affect communities. Manipulating the presence of omnivorous tipulid larvae showed that negative interactions from tipulids, presumably from predation, arose under drought, and were stronger at the warmer location, stressing the importance of species interactions in mediating community responses to climate change. The common garden treatments revealed that prior community exposure to potential future climatic conditions did not affect the outcome. In this powerful experiment, we demonstrated how complexities arise from the interplay of biotic and abiotic mechanisms of climate change. We stress that single species can steer ecological outcomes, and suggest that focusing on such disproportionately influential species may improve attempts at making meaningful predictions of climate change impacts on food webs.     

Lack of stasis in late Cenozoic marine faunas and communities, central California

Pliocene strata in the Kettleman Hills of west-central California were deposited in the broad San Joaquin embayment as a cyclic succession of parasequences during approximately three million years. Depositional environments within each cycle ranged from relatively open marine to brackish and non-marine. Although the strata were deposited in similar, recurrent environments, the fauna changed gradually rather than during brief intervals separating periods of stasis. Although environmental gradients and community structure in the Pliocene San Joaquin Embayment and in the present-day San Francisco Bay are similar, species compositions of the faunas and of parallel communities at the two sites are markedly different. In addition, times of origination and extinction of species in the Pliocene strata of the Kettleman Hills and in San Francisco Bay do not document coordinated stasis within the Late Cenozoic. In contrast, Silurian and Devonian faunas and communities of the Appalachian Basin persisted with little change in ecological-evolutionary units that lasted for up to eight million years. Relatively brief intervals of great biotic change separate these intervals of stasis. One intriguing explanation of this pattern of coordinated stasis within an ecologic-evolutionary unit is ecological locking, in which interaction between species within the community is sufficiently strong that only major changes in the environment are able to change community and faunal composition. Probably the late Cenozoic fauna underwent rapid evolution as a result of rapidly changing environmental conditions within a complex and changing shallow, inshore marine paleogeography. In contrast, coordinated stasis in the lower Paleozoic probably resulted from negligible evolution during long periods of stable to gradually changing environments in an outer shelf setting, punctuated by brief episodes of abrupt environmental change and large-scale turnover. The independent assortment of species in late Cenozoic parallel communities indicates that ecological locking did not exist.     

Climatic variables influence the temporal dynamics of an anuran metacommunity in a nonstationary way

Understanding the temporal dynamics of communities is crucial to predict how communities respond to climate change. Several factors can promote variation in phenology among species, including tracking of seasonal resources, adaptive responses to other species, demographic stochasticity, and physiological constraints. The activities of ectothermic vertebrates are sensitive to climatic variations due to the effect of temperature and humidity on species physiology. However, most studies on temporal dynamics have analyzed multi‐year data and do not have resolution to discriminate within‐year patterns that can determine community assembly cycles. Here, we tested the temporal stability and synchrony of calling activity and also how climatic variables influence anuran species composition throughout the year in a metacommunity in the Atlantic Forest of southern Brazil. Using a multivariate method, we described how the relationship between species composition and climatic variables changes through time. The metacommunity showed a weak synchronous spatial pattern, meaning that species responded independently to environmental variation. Interestingly, species composition exhibited a nonstationary response to climate, suggesting that climate affects species composition differently depending on the season. The species‐climate relationship was stronger during the spring, summer, and winter, mainly influenced by temperature, rainfall, and humidity. Thus, temporal community dynamics seem to be mediated by species life‐history traits, in which independent fluctuations promote community stability in temporally varying environments.     

Ecological genomics meets community‐level modelling of biodiversity: mapping the genomic landscape of current and future environmental adaptation

Local adaptation is a central feature of most species occupying spatially heterogeneous environments, and may factor critically in responses to environmental change. However, most efforts to model the response of species to climate change ignore intraspecific variation due to local adaptation. Here, we present a new perspective on spatial modelling of organism–environment relationships that combines genomic data and community‐level modelling to develop scenarios regarding the geographic distribution of genomic variation in response to environmental change. Rather than modelling species within communities, we use these techniques to model large numbers of loci across genomes. Using balsam poplar (Populus balsamifera) as a case study, we demonstrate how our framework can accommodate nonlinear responses of loci to environmental gradients. We identify a threshold response to temperature in the circadian clock gene GIGANTEA‐5 (GI5), suggesting that this gene has experienced strong local adaptation to temperature. We also demonstrate how these methods can map ecological adaptation from genomic data, including the identification of predicted differences in the genetic composition of populations under current and future climates. Community‐level modelling of genomic variation represents an important advance in landscape genomics and spatial modelling of biodiversity that moves beyond species‐level assessments of climate change vulnerability.     

Disentangling environmental and spatial processes of community assembly in tropical forests from local to regional scales

Understanding patterns and mechanisms of variation in the compositional structure of communities across spatial scales is one of the fundamental challenges in ecology and biogeography. In this study, we evaluated the effects of spatial extent (i.e. size of study region) on: 1) whether community composition can be better explained by environmental (i.e. niche‐based) or spatial (e.g. dispersal‐based) processes ; and 2) how climate and soils contribute to the influence of environment on plant community composition. We surveyed community composition across a network of 398 forest plots spanning a ～4000 m elevational gradient in the Madidi region in northwestern Bolivia. Using redundancy analyses and hierarchical variation partitioning, we disentangled the effects of environmental and spatial predictors on species composition, further decomposing the environmental effect between its climatic and soil components. We repeated analyses for 200 sub‐regions ranging in spatial extent from ～250 to ～17 500 km². Our analyses show a high degree of idiosyncrasy in results that come from different sub‐regions. Despite this variability, we were able to identify various important patterns in the structure of tropical plant communities in our study system. First, even though sub‐regions varied in size by nearly two orders of magnitude, the total amount of explained variation in community composition was scale independent; at all spatial scales, environment and space accounted for about 25% of the differences in community composition among plots. Second, the measured environmental effect was higher than the spatial effect on average and in the vast majority of sub‐regions. This was true regardless of the spatial extent of analysis. Finally, we found that both climatic and soil variables accounted for significant fractions of variation, but climate was always more important than soils.     

Species divergence and trait convergence in experimental plant community assembly

Despite decades of research, it remains controversial whether ecological communities converge towards a common structure determined by environmental conditions irrespective of assembly history. Here, we show experimentally that the answer depends on the level of community organization considered. In a 9‐year grassland experiment, we manipulated initial plant composition on abandoned arable land and subsequently allowed natural colonization. Initial compositional variation caused plant communities to remain divergent in species identities, even though these same communities converged strongly in species traits. This contrast between species divergence and trait convergence could not be explained by dispersal limitation or community neutrality alone. Our results show that the simultaneous operation of trait‐based assembly rules and species‐level priority effects drives community assembly, making it both deterministic and historically contingent, but at different levels of community organization.