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Gastro-intestinal nematode (GIN) parasites play an important role in the ecological dynamics of many animal populations. Recent studies suggest fine-scale spatial variation in GIN infection dynamics are important in wildlife systems, but the environmental drivers underlying this variation remain poorly understood.We used data from over two decades of GIN parasite egg counts, host space use, and spatial vegetation data from a long-term study of Soay sheep on St Kilda to test how spatial autocorrelation and vegetation in an individual’s home range predict parasite burden across three age groups. We developed a novel approach to quantify the plant functional traits present in a home range to describe the quality of vegetation present.Effects of space and vegetation varied between age classes. In immature lambs, strongyle parasite faecal egg counts (FEC) were spatially structured, being highest in the north and south of our study area. Independent of host body weight and spatial autocorrelation, plant functional traits predicted parasite egg counts. Higher egg counts were associated with more digestible and preferred plant functional traits, suggesting the association could be driven by host density and habitat preference.In contrast, we found no evidence that parasite FEC were related to plant functional traits in the host home range in yearlings or adult sheep. Adult FEC were spatially structured, with highest burdens in the north-east of our study area, while yearling FEC showed no evidence of spatial structuring.Our findings support the importance of fine-scale environmental variation for wildlife disease ecology and provides new evidence that such effects may vary across demographic groups within a population. Parasite burdens in immature individuals appear more readily influenced by fine-scale spatial variation in the environment, highlighting the importance of such heterogeneity for our understanding of wildlife epidemiology and health.

Responses to climate change have often been found to lag behind the rate of warming that has occurred. In addition to dispersal limitation potentially restricting spread at leading range margins, the persistence of species in new and unsuitable conditions is thought to be responsible for apparent time-lags.Soil seed banks can allow plant communities to temporarily buffer unsuitable environmental conditions, but their potential to slow responses to long-term climate change is largely unknown. As local forest cover can also buffer the effects of a warming climate, it is important to understand how seed banks might interact with land cover to mediate community responses to climate change.We first related species-level seed bank persistence and distribution-derived climatic niches for 840 plant species. We then used a database of plant community data from grasslands, forests and intermediate successional habitats from across Europe to investigate relationships between seed banks and their corresponding herb layers in 2763 plots in the context of climate and land cover.We found that species from warmer climates and with broader distributions are more likely to have a higher seed bank persistence, resulting in seed banks that are composed of species with warmer and broader climatic distributions than their corresponding herb layers. This was consistent across our climatic extent, with larger differences (seed banks from even warmer climates relative to vegetation) found in grasslands.Synthesis. Seed banks have been shown to buffer plant communities through periods of environmental variability, and in a period of climate change might be expected to contain species reflecting past, cooler conditions. Here, we show that persistent seed banks often contain species with relatively warm climatic niches and those with wide climatic ranges. Although these patterns may not be primarily driven by species' climatic adaptations, the prominence of such species in seed banks might still facilitate climate-driven community shifts. Additionally, seed banks may be related to ongoing trends regarding the spread of widespread generalist species into natural habitats, while cool-associated species may be at risk from both short- and long-term climatic variability and change. Responses to climate change have often been found to lag behind the rate of warming that has occurred. In addition to dispersal limitation potentially restricting spread at leading range margins, the persistence of species in new and unsuitable conditions is thought to be responsible for apparent time-lags. Soil seed banks can allow plant communities to temporarily buffer unsuitable environmental conditions, but their potential to slow responses to long-term climate change is largely unknown. As local forest cover can also buffer the effects of a warming climate, it is important to understand how seed banks might interact with land cover to mediate community responses to climate change. We first related species-level seed bank persistence and distribution-derived climatic niches for 840 plant species. We then used a database of plant community data from grasslands, forests and intermediate successional habitats from across Europe to investigate relationships between seed banks and their corresponding herb layers in 2763 plots in the context of climate and land cover. We found that species from warmer climates and with broader distributions are more likely to have a higher seed bank persistence, resulting in seed banks that are composed of species with warmer and broader climatic distributions than their corresponding herb layers. This was consistent across our climatic extent, with larger differences (seed banks from even warmer climates relative to vegetation) found in grasslands. Synthesis. Seed banks have been shown to buffer plant communities through periods of environmental variability, and in a period of climate change might be expected to contain species reflecting past, cooler conditions. Here, we show that persistent seed banks often contain species with relatively warm climatic niches and those with wide climatic ranges. Although these patterns may not be primarily driven by species' climatic adaptations, the prominence of such species in seed banks might still facilitate climate-driven community shifts. Additionally, seed banks may be related to ongoing trends regarding the spread of widespread generalist species into natural habitats, while cool-associated species may be at risk from both short- and long-term climatic variability and change.

Many ecosystems are grazed by livestock or large, wild herbivores and exist as mosaics of different vegetation communities. Changing grazing could have an impact on heterogeneity as well as on composition. A long-term, large-scale grazing experiment that maintained existing low-intensity sheep grazing, tripled it, removed it and partially substituted sheep grazing by cattle grazing was set up on a mosaic of upland vegetation types. The impact of changing grazing regimes was assessed in terms of changes in temporal and spatial species and functional beta diversity. Removal of grazing had the highest impact on species replacement, whilst increased grazing was closest to maintaining the original species complement. Wet heath and Molina mire had the lowest turnover, but wet heath showed the highest changes in unidirectional abundance as it contained species capable of increasing in abundance in response to changing grazing intensity. Agrostis-Festuca and Nardus grasslands displayed the highest level of balanced species replacement reflecting their more dynamic vegetation. In functional terms, there was no clear separation of communities based on their grazing preference, all were relatively resistant to change but Nardus grassland was the most resistant to the removal of grazing. The increased offtake associated with increased grazing led to a degree of homogenisation as grazing tolerant species associated with preferred communities increased in the unpreferred ones. Decisions about grazing management of the uplands involve many trade-offs, and this study identified potential trade-offs between stability and homogenisation to add to existing ones on the biodiversity of different groups of species and on ecosystem services.

Plant mycorrhizal status (a trait indicating the ability to form mycorrhizas) can be a useful plant trait for predicting changes in vegetation influenced by increased fertility. Mycorrhizal fungi enhance nutrient uptake and are expected to provide a competitive advantage for plants growing in nutrient-poor soils; while in nutrient-rich soils, mycorrhizal symbiosis may be disadvantageous. Some studies in natural systems have shown that mycorrhizal plants can be more frequent in P and N-poor soils (low nutrient availability) or Ca and Mg-high (high pH) soils, but empirical support is still not clear. Using vegetation and soil data from Scottish coastal habitats, and Latvian and Czech grasslands, we examined whether there is a link between plant mycorrhizal status and plant-available P, N, Ca and Mg. We performed the max test analysis (to examine the central tendency) and a combination of quantile regression and meta-analysis (to examine tendencies in different quantiles) on both community and plant species data combined with plant phylogenies. We consistently found no changes in mycorrhizal status at the community and species levels along the gradients of plant-available P, N, Ca and Mg in the central tendency and in almost all quantiles across all datasets. Thus, we found no support for the hypotheses that herbaceous species which are able to form mycorrhizas are more frequent in nutrient-poor and high pH environments. Obligatory, facultatively and non-mycorrhizal herbaceous species appear to assemble randomly along the gradients of nutrient availability in several European herbaceous habitats, suggesting that all these strategies perform similarly under non-extreme soil nutrient conditions.

SummaryMonitoring the prevalence and abundance of parasites over time is important for addressing their potential impact on host life-histories, immunological profiles, and their influence as a selective force. Only long-term ecological studies have the potential to shed light on both the temporal trends in infection prevalence and abundance and the drivers of such trends, because of their ability to dissect drivers that may be confounded over shorter time scales. Despite this, only a relatively small number of such studies exist. Here, we analysed changes in the prevalence and abundance of gastrointestinal parasites in the wild Soay sheep population of St Kilda across 31 years. The host population density has increased across the study, and population density is known to increase parasite transmission, but we found that density and year explained temporal variation in parasite prevalence and abundance independently. Prevalence of both strongyle nematodes and coccidian microparasites increased during the study, and this effect varied between lambs, yearlings and adults. Meanwhile, abundance of strongyles was more strongly linked to host density than to temporal (yearly) dynamics, while abundance of coccidia showed a strong temporal trend without any influence of density. Strikingly, coccidian abundance increased threefold across the course of the study in lambs, while increases in yearlings and adults were negligible. Our decades-long, intensive, individual-based study will enable the role of environmental change and selection pressures in driving these dynamics to be determined, potentially providing unparalleled insight into the drivers of temporal variation in parasite dynamics in the wild.Key findingsWe studied temporal trends in gastrointestinal parasites of wild sheep over 31 yearsYear and host population density explained temporal variation in parasites independentlyPrevalence of both strongyle nematodes and coccidia varied across the study periodAbundance of strongyles was more closely linked to host density than yearAbundance of coccidia increased threefold over time in lambs, but did not vary in adults

Aim: To quantify the responses of alpha and beta diversity to multivariate gradients, incorporating variation in environmental and management variability in coastal dune slacks. Location: United Kingdom dune slacks. Methods: Plant community composition, plant nutrient status and soil characteristics were measured for 164 quadrats in 41 dune slacks across 12 coastal sand dune systems. Data were collated on climate and atmospheric deposition. Hydrological regimes at daily resolution were modelled and calibrated using daily-to-monthly site measurements, from which we calculated quadrat-level hydrological metrics. Alpha diversity (richness, Shannon diversity and Pielou's evenness) metrics and beta diversity (turnover and nestedness) for species and genera were calculated across three spatial levels from sand dune system (highest) to dune slack to quadrat (lowest). Results: Diversity patterns depended on the spatial and taxonomic level considered. At smaller spatial levels (between dune slacks and between quadrats), alpha and beta diversity varied along gradients driven by soil characteristics, water table depth and atmospheric deposition. At larger spatial levels (between sand dune systems), patterns of beta diversity were a consequence of plant nutrient status. There was little variability in alpha diversity along this same gradient, with only small changes in Pielou's species evenness. Patterns at a coarser taxonomic level (genus) mirrored those at the species level. Main conclusion: We show that patterns of variation in plant diversity are dependent on the spatial level considered, but taxonomic level made little difference in understanding these patterns. Therefore, if we do not consider patterns across different spatial levels, important environmental and management drivers could be missed. The high biodiversity value and degree of threat to these European protected habitats makes such understanding invaluable for their conservation.

SummaryEcological 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, specially across different ecosystems.To address this, we compiled a global collection of long-term permanent vegetation records (>7000 plots from 78 datasets) from a wide range of habitats and combined this with existing trait databases. We tested whether the observed inter-annual variability in species abundance (coefficient of variation) was related to multiple individual traits and multivariate axes of trait variations (PCoA axes).We found that species with greater leaf dry matter content and seed mass were consistently more stable over time (lower variability in species abundance) although other leaf traits played a significant role as well, albeit weaker. Using multivariate axes did not improve predictions by specific traits.Our results confirm existing theory, providing compelling empirical evidence on the importance of specific traits, which point at ecological trade-offs in different resource use and dispersal strategies, on the stability of plant populations worldwide.

Aim\ud \ud Climate and land use are key determinants of biodiversity, with past and ongoing changes posing serious threats to global ecosystems. Unlike most other organism groups, plant species can possess dormant life‐history stages such as soil seed banks, which may help plant communities to resist or at least postpone the detrimental impact of global changes. This study investigates the potential for soil seed banks to achieve this.\ud \ud \ud \ud Location\ud \ud Europe.\ud \ud \ud \ud Time period\ud \ud 1978–2014.\ud \ud \ud \ud Major taxa studied\ud \ud Flowering plants.\ud \ud \ud \ud Methods\ud \ud Using a space‐for‐time/warming approach, we study plant species richness and composition in the herb layer and the soil seed bank in 2,796 community plots from 54 datasets in managed grasslands, forests and intermediate, successional habitats across a climate gradient.\ud \ud \ud \ud Results\ud \ud Soil seed banks held more species than the herb layer, being compositionally similar across habitats. Species richness was lower in forests and successional habitats compared to grasslands, with annual temperature range more important than mean annual temperature for determining richness. Climate and land‐use effects were generally less pronounced when plant community richness included seed bank species richness, while there was no clear effect of land use and climate on compositional similarity between the seed bank and the herb layer.\ud \ud \ud \ud Main conclusions\ud \ud High seed bank diversity and compositional similarity between the herb layer and seed bank plant communities may provide a potentially important functional buffer against the impact of ongoing environmental changes on plant communities. This capacity could, however, be threatened by climate warming. Dormant life‐history stages can therefore be important sources of diversity in changing environments, potentially underpinning already observed time‐lags in plant community responses to global change. However, as soil seed banks themselves appear, albeit less, vulnerable to the same changes, their potential to buffer change can only be temporary, and major community shifts may still be expected.

Analysing temporal patterns in plant communities is extremely important to quantify the extent and the consequences of ecological changes, especially considering the current biodiversity crisis. Long-term data collected through the regular sampling of permanent plots represent the most accurate resource to study ecological succession, analyse the stability of a community over time and understand the mechanisms driving vegetation change. We hereby present the LOng-Term Vegetation Sampling (LOTVS) initiative, a global collection of vegetation time-series derived from the regular monitoring of plant species in permanent plots. With 79 data sets from five continents and 7,789 vegetation time-series monitored for at least 6 years and mostly on an annual basis, LOTVS possibly represents the largest collection of temporally fine-grained vegetation time-series derived from permanent plots and made accessible to the research community. As such, it has an outstanding potential to support innovative research in the fields of vegetation science, plant ecology and temporal ecology., The authors acknowledge institutional support as follows. Nicola J. Day: Te Apārangi Royal Society of New Zealand (Rutherford Postdoctoral Fellowship). Jiří Danihelka: Czech Science Foundation (project no. 19-28491X) and Czech Academy of Sciences (project no. RVO 67985939). Francesco de Bello: Spanish Plan Nacional de I+D+i (project PGC2018-099027-B-I00). Eric Garnier: La Fage INRA experimental station. Tomáš Herben: GAČR grant 20-02901S. Anke Jentsch: German Federal Ministry of Education and Research (grant 031B0516C - SUSALPS) and Oberfrankenstiftung (grant OFS FP00237). Norbert Juergens: German Federal Ministry of Education and Research (grant 01LG1201N - SASSCAL ABC). Frédérique Louault and Katja Klumpp: AnaEE-France (ANR-11-INBS-0001). Robin J. Pakeman: Strategic Research Programme of the Scottish Government's Rural and Environment Science and Analytical Services Division. Meelis Pärtel: Estonian Research Council (PRG609) and European Regional Development Fund (Centre of Excellence EcolChange). Josep Peñuelas: Spanish Government (grant PID2019-110521GB-I00), Fundación Ramon Areces (grant ELEMENTAL-CLIMATE), Catalan Government (grant SGR 2017-1005), and European Research Council (Synergy grant ERC-SyG-2013-610028, IMBALANCE-P). Ute Schmiedel: German Federal Ministry of Education and Research (Promotion numbers 01LC0024, 01LC0024A, 01LC0624A2, 01LG1201A, 01LG1201N). Hana Skálová: GAČR grant 20-02901S. Karsten Wesche: International Institute Zittau, Technische Universität Dresden. Susan K. Wiser: New Zealand Ministry for Business, Innovation and Employment’s Strategic Science Investment Fund. Ben A. Woodcock: NERC and BBSRC (NE/N018125/1 LTS-M ASSIST - Achieving Sustainable Agricultural Systems). Enrique Valencia: Program for attracting and retaining talent of Comunidad de Madrid (no. 2017-T2/AMB-5406) and Community of Madrid and Rey Juan Carlos University (Young Researchers R&D Project. Ref. M2165 – INTRANESTI). Truman P. Young: National Science Foundation (LTREB DEB 19-31224).

There is a drive to improve the sustainability of agricultural systems including the biodiversity component. Cultivar mixtures offer yield benefits from the same land area, but the mechanisms behind this overyielding have not been completely worked out. One potential mechanism is improved competition with weeds. We use an experimental approach of varying barley (Hordeum vulgare L.) genotypic and phenotypic diversity to test the hypothesis that increases in diversity have an impact on weed growth strategies and community assembly, or if responses are driven by barley performance. There was no effect of increasing barley mixture diversity on weed traits, either in terms of species means or the community-weighted mean. However, Functional Richness of the weed community decreased with increasing barley mixture diversity mainly as a result of reduced specific leaf area Functional Richness. This pattern was driven by a reduction in species richness of the weed community rather than by reduced variation within species. Whilst barley phenotype had different impacts on weed traits, there were no specific mixtures of phenotypes or genotypes that had consistent effects on community assembly or weed species responses. The competitive exclusion of weed species could have agronomic and environmental benefits, through better targeting or less frequent use of herbicides. Growing crop mixtures is one of many strategies available to improve agricultural sustainability and resilience, and one that has clear benefits. ISSN:0043-1737 ISSN:1365-3180

Soil seed banks represent a critical but hidden stock for potential future plant diversity on Earth. Here we compiled and analyzed a global dataset consisting of 15,698 records of species diversity and density for soil seed banks in natural plant communities worldwide to quantify their environmental determinants and global patterns. Random forest models showed that absolute latitude was an important predictor for diversity of soil seed banks. Further, climate and soil were the major determinants of seed bank diversity, while net primary productivity and soil characteristics were the main predictors of seed bank density. Moreover, global mapping revealed clear spatial patterns for soil seed banks worldwide; for instance, low densities may render currently species-rich low latitude biomes (such as tropical rain-forests) less resilient to major disturbances. Our assessment provides quantitative evidence of how environmental conditions shape the distribution of soil seed banks, which enables a more accurate prediction of the resilience and vulnerabilities of plant communities and biomes under global changes., Soil seed banks are reservoirs of plant biodiversity. Here the authors compile a global dataset of soil seed banks in natural plant communities and report a spatially explicit analysis of environmental controls of seed bank density and diversity.

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.

Plasticity is a widely used concept in plant sciences, but there is inconsistency over its interpretation and measurement. One aspect of plasticity – adaptive plasticity – may be particularly important in shaping plant fitness and reproductive success and represents the first line of a plants defence to environmental change. Here, we define adaptive plasticity, highlight its importance to plant growth and survival, and suggest appropriate approaches for its measurement. We argue that a focus on adaptive plasticity could help address some fundamental challenges in plant ecology and evolutionary biology, including developing insight into climate-change resilience of natural populations and crops.

Analysing temporal patterns in plant communities is extremely important to quantify the extent and the consequences of ecological changes, especially considering the current biodiversity crisis. Long-term data collected through the regular sampling of permanent plots represent the most accurate resource to study ecological succession, analyse the stability of a community over time and understand the mechanisms driving vegetation change. We hereby present the LOng-Term Vegetation Sampling (LOTVS) initiative, a global collection of vegetation time-series derived from the regular monitoring of vascular plants in permanent plots. With 79 datasets from five continents and 7789 vegetation time-series monitored for at least six years and mostly on an annual basis, LOTVS possibly represents the largest collection of temporally fine-grained vegetation time-series derived from permanent plots and made accessible to the research community. As such, it has an outstanding potential to support innovative research in the fields of vegetation science, plant ecology and temporal ecology.

Questions: Hydrological niche segregation is widespread and has been found across a range of different habitats. Different plant species can occupy distinct hydrological niches, and as a result fine-scale variability in hydrology can structure plant communities. However, these patterns may not be as clear in habitats where differences in hydrology are more short-lived, such as coastal dune slacks. We explored the extent that the hydrological regime structures dune slack plant communities. Location: Ainsdale Coastal Sand Dune National Nature Reserve, UK. Methods: Six hundred quadrats were surveyed, 100 in each of six coastal dune slacks. Water table levels are recorded monthly in each slack. Metrics summarising hydrological regime were calculated and adjusted for each quadrat based on elevation. We tested the relationship between water table depth, plant communities and topography across and within dune slacks. Results: Half (three) of the slacks showed a significant influence of hydrology on plant community composition. The three that did not were the ones that varied least topographically and contained less diverse plant communities. We also provide indirect evidence of niche segregation by modelling species-specific responses between mean water table depth and probability of species presence. Conclusions: We demonstrate that hydrology is a dominant driver of plant community composition across dune slacks. However, plant communities are not always structured by hydrology, demonstrating the complexity of vegetation patterns. Topographic variation appears to impact plant community patterns, as do successional processes, highlighting the need to create diverse habitats for slack restoration and management.

Better understanding of the mechanistic basis of plant plasticity will enhance efforts to breed crops resilient to predicted climate change. However, complexity in plasticity's conceptualisation and measurement may hinder fruitful crossover of concepts between disciplines that would enable such advances. We argue active adaptive plasticity is particularly important in shaping the fitness of wild plants, representing the first line of a plant's defence to environmental change. Here, we define how this concept may be applied to crop breeding, suggest appropriate approaches to measure it in crops, and propose a refocussing on active adaptive plasticity to enhance crop resilience. We also discuss how the same concept may have wider utility, such as in ex situ plant conservation and reintroductions., Trends in Plant Science, 27 (7), ISSN:1360-1385, ISSN:1878-4372

The long-term project on St Kilda, including field assistant Jill Pilkington, has been largely funded by the UK Natural Environment Research Council and for one period by the Wellcome Trust. 1. The population of Soay sheep on the island of Hirta in the Outer Hebrides has been the subject of continuous study for more than 35 years. This paper focuses on the botanical aspects of the plant-herbivore interaction, showing how the vegetation affects and is affected by the sheep. 2. Grazing impacts on biomass and spatial structure varied across plant communities, with Holcus/Agrostis grasslands affected most and Wet Heath least, consistent with the hypothesis that herbivore impacts are proportional to plant productivity. 3. Within plant communities, the negative relationships between sheep numbers and plant abundance (sward height, gap/tussock cover and biomass) were significant in March but not significant in August, as expected if sheep numbers are limited by food supply in winter. 4. In most species, flower stem density declined with increasing sheep numbers. There were no examples where unpalatable plants showed increased flowering (e.g. from competitor release under selective grazing). 5. Plant production in temporary grazing exclosures (above ground net primary production) was greatest in Holcus/Agrostis grassland (12.6t/ha dry matter/year), lower in Nardus grassland (5.1t/ha/yr) and least in Plantago sward (1.3t/ha/yr) associated with differences in historical nutrient supply and microclimate. 6. The net effect of grazing on plant species richness was positive: a few highly palatable species were excluded, but small-scale coexistence of grazing tolerant species was enhanced by defoliation. 7. The Soay sheep population fluctuated from 908 (in 1988) to 2208 (in 2009), increasing by an average of 39 extra animals per year over the period 1985-2011. Between 2011 and 2020, the population fluctuated less widely and showed no trend. 8. Population change (ln(N(t+1)/N(t)) was inversely density dependent but positively correlated with plant production in Holcus/Agrostis grassland which increased during the study. 9. This plant-herbivore interaction is highly resilient, and though some species (Festuca rubra and Ranunculus acris) declined in the Holcus/Agrostis grassland, there was no significant upward trend in the abundance of unpalatable plant species. 10. Synthesis. Implications for future studies and analyses of plant-herbivore data. Concentrating on estimating primary productivity and herbivore offtake, rather than simply measuring change in plant biomass, is likely to provide greatly improved explanatory power for understanding herbivore population dynamics. Postprint

Plant mycorrhizal status (a trait indicating the ability to form mycorrhizas) can be a useful plant trait for predicting changes in vegetation influenced by increased fertility. Mycorrhizal fungi enhance nutrient uptake and are expected to provide a competitive advantage for plants growing in nutrient-poor soils; while in nutrient-rich soils, mycorrhizal symbiosis may be disadvantageous. Some studies in natural systems have shown that mycorrhizal plants can be more frequent in P and N-poor soils (low nutrient availability) or Ca and Mg-high (high pH) soils, but empirical support is still not clear. Using vegetation and soil data from Scottish coastal habitats, and Latvian and Czech grasslands, we examined whether there is a link between plant mycorrhizal status and plant-available P, N, Ca and Mg. We performed the max test analysis (to examine the central tendency) and a combination of quantile regression and meta-analysis (to examine tendencies in different quantiles) on both community and plant species data combined with plant phylogenies. We consistently found no changes in mycorrhizal status at the community and species levels along the gradients of plant-available P, N, Ca and Mg in the central tendency and in almost all quantiles across all datasets. Thus, we found no support for the hypotheses that herbaceous species which are able to form mycorrhizas are more frequent in nutrient-poor and high pH environments. Obligatory, facultatively and non-mycorrhizal herbaceous species appear to assemble randomly along the gradients of nutrient availability in several European herbaceous habitats, suggesting that all these strategies perform similarly under non-extreme soil nutrient conditions.

Environmental policy decisions should be based on robust indicators of changes in the environment. In Scotland these have been formalised as Ecosystem Health Indicators. Indicators work best where there is a direct link between what is measured and environmental change; changes in indices of species diversity or abundance provide alerts to environmental change but do not in themselves provide evidence of mechanisms. An alternative approach is to replace species occurrence data with species habitat or attribute values to create an indicator that links changes in species occurrences to environmental drivers. A series of indicators based on biological recording data, Ellenberg habitat preference values and mean climate niche was created by taking all records for bryophytes in Scotland (522187 records, 1960–2016) and analysing changes in mean attribute values through time for each 10 km × 10 km grid square. The resulting mean attribute values were analysed at the country level and showed clear quadratic (humped) trends for Ellenberg Nitrogen (N) and Reaction (R), an index of environmental acidity. There are linear decreasing trends for Ellenberg Light (L) and Precipitation, and linear increasing trends for January and July Temperature. Ellenberg Moisture (F) showed a clear trend, but it was not interpretable and appeared influenced by recording effort. Disaggregating to region was partly successful, but reduced samples sizes meant that for many regions the national trend was not significant. Ellenberg N and the two Temperature indices appear to be useful indicators as they are interpretable and performed better in the downscaling. Changes in L were also interpretable, but it performed poorly in downscaling to regional values. Ellenberg R appeared to be driven by changes in N as the two values are highly correlated across species. The downward trend in Precipitation ran counter to increases in rainfall suggesting that temperature driven evaporation is more likely to be driving changes in bryophyte distribution and whilst the indicator performed well it is difficult to explain to non-experts. Opportunistically collected data, like plant records, have great potential to be employed as indicators, especially as citizen scientists are involved and state resources can be used for data archiving and analysis. Also, conversion to attribute values removes many of the biases brought by differences in recorder effort and it directly links changes in species occurrences to environmental drivers.

Marginal agricultural land, which in the UK refers to upland grazings in particular, is going to see changes in management driven by markets, subsidies, grants and environmental change with implications for biodiversity. Using a large‐scale, long‐term grazing experiment in the UK uplands we assessed the impact of intensification (tripling sheep numbers), abandonment (removal of sheep) and grazer diversification (partial replacement of sheep by cattle) on vegetation composition in a heterogenous area of grassland. Species benefiting from increased grazing included sweet vernal grass (Anthoxanthum odoratum), mat grass (Nardus stricta) and deer grass (Trichophorum cespitosum). Species that benefitted from the removal of grazing included bog asphodel (Narthecium ossifragum), bracken (Pteridium aquilinum) and blaeberry (Vaccinium myrtillus). Responses differed between vegetation communities; more productive acid grassland communities showed little change when grazing was removed, whilst less productive mire communities contained species, capable of increasing after grazing removal. Increased grazing and, to a lesser extent, the introduction of cattle increased species diversity. Synthesis and applications. Marginal agricultural grassland will likely see management change driven by markets, subsidies and environmental change. Vegetation change in these relatively infertile grasslands is slow and features shuffling dominance amongst species in the initial vegetation. Initial structural changes affect other trophic levels in this experiment; the slow change in composition will affect the system over longer timescales. Management decisions in the uplands encompass complex trade‐offs between production, biodiversity and a range of ecosystem services. Predicting the consequences of decisions is difficult given the slow dynamics of unproductive habitats.

International audience; The global scientific community has become increasingly diverse over recent decades, but is this ongoing development also reflected among top‐publishing authors and potential scientific leaders? We surveyed 13 leading journals in ecology, evolution, and conservation to investigate the diversity of the 100 top‐publishing authors in each journal between 1945 and 2019. Out of 1051 individual top‐publishing authors, only 11% are women. The United States, the United Kingdom, Australia, Germany, and Canada account for more than 75% of top‐publishing authors, while countries of the Global South (as well as Russia, Japan, and South Korea) were strikingly underrepresented. The number of top‐publishing authors who are women and/or are from the Global South is increasing only slowly over time. We outline transformative actions that scientific communities can take to enhance diversity, equity and inclusion at author, leadership, and society level. The resulting promotion of scientific innovation and productivity is essential for the development of global solutions in conservation science.

© 2021 The Authors., Aims: Understanding fine-grain diversity patterns across large spatial extents is fundamental for macroecological research and biodiversity conservation. Using the GrassPlot database, we provide benchmarks of fine-grain richness values of Palaearctic open habitats for vascular plants, bryophytes, lichens and complete vegetation (i.e., the sum of the former three groups). Location: Palaearctic biogeographic realm. Methods: We used 126,524 plots of eight standard grain sizes from the GrassPlot database: 0.0001, 0.001, 0.01, 0.1, 1, 10, 100 and 1,000 m and calculated the mean richness and standard deviations, as well as maximum, minimum, median, and first and third quartiles for each combination of grain size, taxonomic group, biome, region, vegetation type and phytosociological class. Results: Patterns of plant diversity in vegetation types and biomes differ across grain sizes and taxonomic groups. Overall, secondary (mostly semi-natural) grasslands and natural grasslands are the richest vegetation type. The open-access file ”GrassPlot Diversity Benchmarks” and the web tool “GrassPlot Diversity Explorer” are now available online (https://edgg.org/databases/GrasslandDiversityExplorer) and provide more insights into species richness patterns in the Palaearctic open habitats. Conclusions: The GrassPlot Diversity Benchmarks provide high-quality data on species richness in open habitat types across the Palaearctic. These benchmark data can be used in vegetation ecology, macroecology, biodiversity conservation and data quality checking. While the amount of data in the underlying GrassPlot database and their spatial coverage are smaller than in other extensive vegetation-plot databases, species recordings in GrassPlot are on average more complete, making it a valuable complementary data source in macroecology., GrassPlot development has been supported by the Bavarian Research Alliance (BayIntAn_UBT_2017_58), the Eurasian Dry Grassland Group (EDGG) and the International Association for Vegetation Science (IAVS); IB, CorM, JAC, IGM, DGM, MHe, DL and MTo were supported by the Basque Government (IT936‐16); CorM, IAx, MCh, JDa, PD, MHá, ZL, ZPr, EŠ and LT were supported by the Czech Science Foundation (19‐28491X); TR was supported by the Estonian Research Council (PUT1173); RJP was funded by the Strategic Research Programme of the Scottish Government’s Rural and Environmental Science and Analytical Services Division”; SBa was supported by the GINOP‐2.3.2‐15‐2016‐00019 project; GFi was partially supported by the MIUR initiative “Department of excellence” (Law 232/2016)"; BJA was funded by the Spanish Research Agency (grant AEI/ 10.13039/501100011033); AK, VB, IM, DS, IV and DV were supported by the National Research Foundation of Ukraine (2020.01/0140); MP and AH were supported by the Estonian Research Council (PRG874, PRG609), and the European Regional Development Fund (Centre of Excellence EcolChange); Data collection of HCP was funded by FORMAS (Swedish Research Council for Environment, Agricultural Science and Spatial Planning) and The Swedish Institute; JR was supported by the Czech Science Foundation (grant No. 20‐09895S) and the long‐term developmental project of the Czech Academy of Sciences (RVO 67985939); ATRA was funded by the Grant of Excellence Departments, MIUR‐Italy (ARTICOLO 1, COMMI 314 – 337 LEGGE 232/2016); JMA was supported by Carl Tryggers stiftelse för vetenskaplig forskning and Qatar Petroleum; AAli was supported by the Jiangsu Science and Technology Special Project (Grant No. BX2019084), and Metasequoia Faculty Research Startup Funding at Nanjing Forestry University (Grant No. 163010230), and he is currently supported by Hebei University through Faculty Research Startup Funding Program; ZB was supported by the NKFI K 124796 grant; The GLORIA‐ Aragón project of JLBA was funded by the Dirección General de Cambio Climático del Gobierno de Aragón (Spain); MCs and LDem were supported by DG Environment through the European Forum on Nature Conservation and Pastoralism and Barbara Knowles Fund, in collaboration with Pogány‐havas Association, Romania; JDa was partially supported by long‐term research development project no. RVO 67985939 of the Czech Academy of Sciences; BD and OV were supported by the NKFI KH 126476, NKFI KH 130338, NKFI FK 124404 and NKFI FK 135329 grants; BD, OV and AKe were supported by the Bolyai János Scholarship of the Hungarian Academy of Sciences; BE was funded by the Environmental Department of the Tyrolean Federal State Government, the MAB Programme of the Austrian Academy of Science, the Mountain Agriculture Research Unit and the Alpine Research Centre Obergurgl of Innsbruck University. The GLORIA projects of BE were funded by the EU project no. EVK2‐CT‐2000‐00056, the Earth System Sciences Program of the Austrian Academy of Sciences (project MEDIALPS), the Amt für Naturparke, Autonome Provinz Bozen‐Südtirol, the Südtiroler Wissenschaftsfonds and the Tiroler Wissenschaftsfonds; RGG was supported by the Spanish Ministry of Research to sample GLORIA sites in central Spain (CGL 2008‐00901/BOS) and present works by the Autonomous Region of Madrid (REMEDINAL TE‐CM, S2018/EMT‐4338); MJ was supporteLatviaed by Latvia Grant No. 194051; NP and SŠ were partly supported by the Slovenian Research Agency, core fundings P1‐0403 and J7‐1822.

Empirical critical loads are widely used to quantify and manage the ecological impacts of reactive nitrogen (N) deposition. Critical load values aim to identify a level of N deposition below which significant harmful effects do not occur according to present knowledge. Critical loads have been primarily based on experiments, but these are few in number and have well-known limitations, so there is a strong imperative to test and validate values with other forms of evidence. We assembled data on the spatial variability in vegetation communities in the United Kingdom and used Threshold Indicator Taxa Analyses (TITAN) to investigate linkages between species changes and modelled current and cumulative N deposition. Our analyses focused on five datasets: acid grasslands, alpine habitats, coastal fixed dunes, dune slacks and wet grasslands. In four of these habitats there was evidence for a significant decline in the cover of at least one species (a ‘species-loss change-point’) occurring below the critical load, and often at very low levels of N deposition. In all of the habitats there was evidence for clustering of many individual species-loss change-points, implying a community change-point analogous to an ecological threshold. Three of these community change-points occurred below the critical load and the remaining two overlapped with the critical load range. Studies using similar approaches are now increasingly common, with similar results. Across 19 similar analyses there has been evidence for plant species loss change-points below the critical load in 18 analyses, and community-level species loss change-points below the critical load in 13 analyses. None of these analyses has shown community change-points above the critical load. Field data increasingly suggest that many European critical loads are too high to confidently prevent loss of sensitive species.

The stability of ecological communities is critical for the stable provisioning of ecosystem services, such as food and forage production, carbon sequestration, and soil fertility. Greater biodiversity is expected to enhance stability across years by decreasing synchrony among species, but the drivers of stability in nature remain poorly resolved. Our analysis of time series from 79 datasets across the world showed that stability was associated more strongly with the degree of synchrony among dominant species than with species richness. The relatively weak influence of species richness is consistent with theory predicting that the effect of richness on stability weakens when synchrony is higher than expected under random fluctuations, which was the case in most communities. Land management, nutrient addition, and climate change treatments had relatively weak and varying effects on stability, modifying how species richness, synchrony, and stability interact. Our results demonstrate the prevalence of biotic drivers on ecosystem stability, with the potential for environmental drivers to alter the intricate relationship among richness, synchrony, and stability., We were supported by US NSF Grants DEB-8114302, DEB8811884, DEB-9411972, DEB-0080382, DEB-0620652, DEB-1234162, and DEB0618210; the Nutrient Network (https://nutnet.org/) experiment from NSF Research Coordination Network Grant NSF-DEB-1042132; the New Zealand National Vegetation Survey Databank; and Institute on the Environment Grant DG-0001-13. Data (Dataset 56, SI Appendix, Supplementary Text S4) owned by NERC Database Right/Copyright NERC. Further support was provided by the Jornada Basin Long-Term Ecological Research project, Cedar Creek Ecosystem Science Reserve, and the University of Minnesota. The Rothamsted Long-term Experiments National Capability is supported by UK Biotechnology and Biological Sciences Research Council Grant BBS/E/C/000J0300 and the Lawes Agricultural Trust. This research was funded by Czech Science Foundation Grant GACR16-15012S and Czech Academy of Sciences Grant RVO 67985939. E.V. was funded by 2017 Program for Attracting and Retaining Talent of Comunidad de Madrid Grant 2017-T2/AMB-5406.

Questions. Compensatory dynamics are described as one of the main mechanisms that increase community stability, e.g., where decreases of some species on a year‐to‐year basis are offset by an increase in others. Deviations from perfect synchrony between species (asynchrony) have therefore been advocated as an important mechanism underlying biodiversity effects on stability. However, it is unclear to what extent existing measures of synchrony actually capture the signal of year‐to‐year species fluctuations in the presence of long‐term directional trends in both species abundance and composition (species directional trends hereafter). Such directional trends may lead to a misinterpretation of indices commonly used to reflect year‐to‐year synchrony. Methods. An approach based on three‐term local quadrat variance (T3) which assesses population variability in a three‐year moving window, was used to overcome species directional trend effects. This “detrending” approach was applied to common indices of synchrony across a worldwide collection of 77 temporal plant community datasets comprising almost 7,800 individual plots sampled for at least six years. Plots included were either maintained under constant “control” conditions over time or were subjected to different management or disturbance treatments. Results. Accounting for directional trends increased the detection of year‐to‐year synchronous patterns in all synchrony indices considered. Specifically, synchrony values increased significantly in ~40% of the datasets with the T3 detrending approach while in ~10% synchrony decreased. For the 38 studies with both control and manipulated conditions, the increase in synchrony values was stronger for longer time series, particularly following experimental manipulation. Conclusions. Species’ long‐term directional trends can affect synchrony and stability measures potentially masking the ecological mechanism causing year‐to‐year fluctuations. As such, previous studies on community stability might have overemphasised the role of compensatory dynamics in real‐world ecosystems, and particularly in manipulative conditions, when not considering the possible overriding effects of long‐term directional trends., We thank multiple entities for the financial support necessary to obtain the different databases: the U.S. National Science Foundation under grant numbers DEB‐8114302, DEB‐8811884, DEB‐9411972, DEB‐0080382, DEB‐0620652, DEB‐1234162, DEB‐9707477, DEB‐0316402, DEB‐08‐16453, and DEB‐12‐56034, DEB‐0618210, the Nutrient Network (http://www.nutnet.org) experiment from the National Science Foundation Research Coordination Network (NSF‐DEB‐1042132), the New Zealand National Vegetation Survey Databank, the Spanish MINECO (Project CGL2014‐53789‐R), the Madrid Regional Government (Projects REMEDINAL‐3 and REMEDINAL‐TE), the European Research Council Synergy grant 610028 (IMBALANCE‐P), the Institute on the Environment (DG‐0001‐13), the SOERE‐ACBB financed through French National Agency for Research (ANAEE‐F, ANR‐11‐INBS‐0001), the Estonian Research Council (IUT 20‐28, IUT 20‐29), Czech Science Foundation (GAČR 17‐05506S and 19‐28491X), the European Regional Development Fund (Centre of Excellence EcolChange), the German Federal Environmental Foundation (DBU) for a grant to the NABU Hamburg (management experiment Calamagrostis epigejos), and the German Federal Ministry of Education and Research within the framework of the project BIOTA Southern Africa (promotion numbers 01LC0024, 01LC0024A and 01LC0624A2), Task 159 of SASSCAL (promotion number 01LG1201) and the Scottish Government's Rural and Environmental Science and Analytical Services division. Acknowledgement Data owned by NERC© Database Right/Copyright NERC. Further support was provided by the Jornada Basin Long‐Term Ecological Research (LTER) project, Cedar Creek Ecosystem Science Reserve and the University of Minnesota. We also thank the Lawes Agricultural Trust and Rothamsted Research for data from the e‐RA database. The Rothamsted Long‐term Experiments National Capability (LTE‐NCG) is supported by the UK Biotechnology and Biological Sciences Research Council (Grant BBS/E/C/000J0300) and the Lawes Agricultural Trust.

The use of functional information in the form of species traits plays an important role in explaining biodiversity patterns and responses to environmental changes. Although relationships between species composition, their traits, and the environment have been extensively studied on a case-by-case basis, results are variable, and it remains unclear how generalizable these relationships are across ecosystems, taxa and spatial scales. To address this gap, we collated 80 datasets from trait-based studies into a global database for metaCommunity Ecology: Species, Traits, Environment and Space; “CESTES”. Each dataset includes four matrices: species community abundances or presences/absences across multiple sites, species trait information, environmental variables and spatial coordinates of the sampling sites. The CESTES database is a live database: it will be maintained and expanded in the future as new datasets become available. By its harmonized structure, and the diversity of ecosystem types, taxonomic groups, and spatial scales it covers, the CESTES database provides an important opportunity for synthetic trait-based research in community ecology., Measurement(s)species abundance • species trait • environmental feature • latitude • longitudeTechnology Type(s)digital curationFactor Type(s)year of data collection • type of ecosystem • level of human disturbanceSample Characteristic - Environmentterrestrial natural environment • fresh water • marine biome • anthropogenic terrestrial biome • natural environment • area of mixed forest • cultivated environmentSample Characteristic - LocationEarth (planet) Machine-accessible metadata file describing the reported data: 10.6084/m9.figshare.11317790

1. The intensity of pastoral management in areas of High Nature Value farming is declining in some regions of Europe but increasing in others. This affects open habitats of conservation concern, such as the British uplands, where bird species that benefit from low-intensity grazing may be most sensitive to such polarization. While experimental manipulations of livestock grazing intensities have improved our understanding of upland breeding bird responses in the short term, none have examined the long-term impacts of altered management on reproductive success. 2. Using a replicated landscape-scale experiment that started in 2003, we investigated the effects of four grazing treatments (intensive sheep; low-intensity sheep; low-intensity mixed sheep and cattle; and no grazing) on the breeding productivity of meadow pipits Anthus pratensis, the most common upland passerine. Surveys were carried out systematically during early (2003 and 2004) and late (2015 and 2016) sampling periods of the experiment to compare the short- and long-term effects of grazing treatments on breeding density and productivity of pipits specifically, but also on the overall bird community. 3. Pipit breeding density was lowest under low-intensity sheep grazing while the highest egg-stage nest survival was observed in the same treatment, although no significant treatment effects were detected on overall nest survival or fledgling output. There were no significant differences in treatment effects between the sampling periods on any breeding variable, but overall nest survival was lower in the later sampling period across all treatments. 4. Breeding bird species richness differed between treatments in the later sampling period, with highest species richness in the ungrazed treatment. 5. Synthesis and applications. Livestock grazing management can have different outcomes for different upland birds. Our results showed that, with time, meadow pipit breeding productivity tended to be higher when sheep grazing intensity was reduced and/or mixed with cattle, and lower when livestock were removed, but not significantly so. Removal of grazing, however, can significantly increase bird species richness. The long-term experiment showed an overall decline in fledglings regardless of grazing treatments, potentially a result of increased predator numbers harboured by nearby developing woodland, highlighting the importance of considering wider landscape processes in grazing management decisions.

The use of functional information in the form of species traits plays an important role in explaining biodiversity patterns and responses to environmental changes. Although relationships between species composition, their traits, and the environment have been extensively studied on a case-by-case basis, results are variable, and it remains unclear how generalizable these relationships are across ecosystems, taxa and spatial scales. To address this gap, we collated 80 datasets from trait-based studies into a global database for metaCommunity Ecology: Species, Traits, Environment and Space; “CESTES”. Each dataset includes four matrices: species community abundances or presences/absences across multiple sites, species trait information, environmental variables and spatial coordinates of the sampling sites. The CESTES database is a live database: it will be maintained and expanded in the future as new datasets become available. By its harmonized structure, and the diversity of ecosystem types, taxonomic groups, and spatial scales it covers, the CESTES database provides an important opportunity for synthetic trait-based research in community ecology., This work was funded by the German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig (funded by the German Research Foundation; DFG FZT 118). We thank Jitendra Gaikwad for his precious help with the technical aspects of the database storage and management through the iDiv Biodiversity Portal (https://idata.idiv.de/, https://doi.org/10.25829/idiv.286-21-2695) and the iDiv IT Support for their help in the CESTES website development (https://icestes.github.io/). The datasets provided by A. Jeliazkov were collected with financial support from the Fédération d’Ile-de-France pour la Recherche en Environnement (FIRE FR-3020). The dataset provided by R. Carvalho and F.L. Tejerina-Garro had financial support from the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq No. 471283/2006-1) granted to FLTG. The datasets provided by R. van Klink, J.-Y. Humbert, R. Arlettaz and M.H.M. Menz were collected with financial support from the Swiss National Science Foundation (grants 31003A_125398/2 and 31003A_149656) awarded to RA.

The predicted long lag time between a decrease in atmospheric deposition and a measured response in vegetation has generally excluded the investigation of vegetation recovery from the impacts of atmospheric deposition. However, policy-makers require such evidence to assess whether policy decisions to reduce emissions will have a positive impact on habitats. Here we have shown that 40 years after the peak of SOx emissions, decreases in SOx are related to significant changes in species richness and cover in Scottish Calcareous, Mestrophic, Nardus and Wet grasslands. Using a survey of vegetation plots across Scotland, first carried out between 1958 and 1987 and resurveyed between 2012 and 2014, we test whether temporal changes in species richness and cover of bryophytes, Cyperaceae, forbs, Poaceae, and Juncaceae can be explained by changes in sulphur and nitrogen deposition, climate and/or grazing intensity, and whether these patterns differ between six grassland habitats: Acid, Calcareous, Lolium, Nardus, Mesotrophic and Wet grasslands. The results indicate that Calcareous, Mesotrophic, Nardus and Wet grasslands in Scotland are starting to recover from the UK peak of SOx deposition in the 1970's. A decline in the cover of grasses, an increase in cover of bryophytes and forbs and the development of a more diverse sward (a reversal of the impacts of increased SOx) was related to decreased SOx deposition. However there was no evidence of a recovery from SOx deposition in the Acid or Lolium grasslands. Despite a decline in NOx deposition between the two surveys we found no evidence of a reversal of the impacts of increased N deposition. The climate also changed significantly between the two surveys, becoming warmer and wetter. This change in climate was related to significant changes in both the cover and species richness of bryophytes, Cyperaceae, forbs, Poaceae and Juncaceae but the changes differed between habitats.

Despite the important roles ground beetles (Coleoptera: Carabidae) play in ecosystems, the highly valued ecosystem services they provide, and ample descriptive documentation of their phenology, the relative impact of various environmental factors on carabid phenology is not well studied. Using the long-term pitfall trap capture data from 12 terrestrial Environmental Change Network (ECN) sites from the UK, we examined how changing climate influenced the phenology of common carabids, and the role particular climate components had on phenological parameters. Of the 28 species included in the analyses, 19 showed earlier start of their activity. This advance was particularly pronounced in the spring, supporting the view that early phenophases have a greater tendency to change and these changes are more directly controlled by temperature than later ones. Autumn activity extended only a few cases, suggesting a photoperiod-driven start of hibernation. No association was found between life-history traits and the ability of species to change their phenology. Air temperatures between April and June were the most important factors determining the start of activity of each species, whilst late season precipitation hastened the cessation of activity. The balance between the advantages and disadvantages of changing phenology on various levels is likely to depend on the species and even on local environmental criteria. The substantially changing phenology of Carabidae may influence their function in ecosystems and the ecosystem services they provide.

Aims: To test a recently developed method of assessing functional beta-diversity - as the difference in location of the convex hull in trait space - as a means of assessing the impact of species loss and colonization over time on functional composition. This approach was tested using data from a survey and subsequent resurvey of Scottish coastal vegetation.Location: Scotland, UK.Results: Fixed dunes, slacks, unimproved grassland and mires showed the highest functional overlap through time (high correspondence in convex hull position between two surveys), while salt marsh, improved grassland and heath showed the lowest. Fixed dune was the most stable in terms of retaining the highest proportion of the original convex hull, while salt marsh was least stable. Salt marsh also showed the highest new functional space occupied, whilst mires and slacks showed the least colonization by functionally distinct species. Generally sites on the west coast of Scotland were most stable, while those on the east showed the greatest functional changes. Assessing functional beta-diversity provided a different picture to assessing species beta-diversity; high species turnover might have little impact on the functional characteristics of the vegetation, whereas the invasion of one functionally different species can have a sizeable impact on ecosystem processes.Conclusions: The method proved effective in highlighting habitats and sites where species changes had the largest impact on the functional space occupied by the plant assemblage. The method provides complementary information to that derived from other types of analysis, including about the potential needs for improved management or full-scale restoration.

We thank Scottish Natural Heritage for funding this work via a PhD studentship to JM. The National Trust for Scotland provided the field site and fenced the experiment. In particular we thank Kate Sampson and Lindsay Mackinlay at NTS staff for their assistance in establishing this experiment. RJM and RJP were funded through the 2011-2016 and 2016-2021 Strategic Research Programmes of the Scottish Government.

Background: Intensive farming affects farmland biodiversity, and some arable plants in particular. Increasing crop genetic diversity can increase crop productivity or resilience and could also benefit rare arable plants. Aims: We examined whether barley presence, sowing density and genetic diversity impacted the rare plant Valerianella rimosa and explored possible underlying mechanisms. Methods: In a field study near Dundee, Scotland, we sowed plots of five single barley genotypes, and all five genotypes combined, at three densities; we also had barley-free plots. Valerianella seeds were sown into half of all plots. Measured responses included early-season cover and harvest biomass of barley and common weeds, abiotic parameters (soil moisture, light) and establishment, biomass and seed production by V. rimosa. Results: Barley presence promoted V. rimosa establishment early in the growing season, but without barley density or genetic diversity effects. By harvest, the impact of barley presence on V. rimosa abundance was lost; there were no effects on Valerianella seed production. Barley negatively impacted common weeds, but V. rimosa did not benefit from any indirect facilitation by barley, being bigger without barley. Conclusions: Early beneficial effects of barley on V. rimosa abundance appear offset by late-season competition. However, limited impacts of barley on V. rimosa reproductive success, and negative impacts on common weeds, indicate crops might play a role in conservation management of rare arable plants by creating space in the weed community. This work was supported by the Rural & Environment Science & Analytical Services Division of the Scottish Government through the Strategic Research Programmes 2011–2016 and 2016–2021. CS was supported by the Swiss National Science Foundation (PZ00P3_148261 and PA00P3_136474).

Analysis of changes in functional traits and functional diversity offers a means of identifying the main environmental drivers of change, the impacts of that change on community assembly and the consequence of change on ecosystem function and service delivery. Changes in key traits and functional diversity in Scottish grasslands were analysed for a resurvey of 561 grasslands sampled an average of 40 years apart and these changes correlated with changes in environmental drivers. There were clear differences between the responses of different grassland habitats; more productive habitats showed evidence of increased agricultural utilisation, whereas more unproductive habitats showed evidence of reduced exploitation. This was echoed in an increase in Functional Richness in unproductive habitats and a decrease in productive ones; the latter indicating increased habitat filtering in the heavily utilised, productive habitats. There were also overall shifts to later-flowering species and less availability of resources for pollinators in less productive habitats. Models relating changes in traits to the environment had little power, but did indicate the importance of pollution and that recovery from the peak in livestock numbers between the surveys had not fully worked through. Similarly, changes in functional diversity were poorly explained by changes in environmental drivers, though increased rainfall had positive impacts on Functional Evenness and Richness. Functional Divergence (measured by Rao’s Q) was, however, strongly linked to changes in species traits over time. Scottish grasslands have diverged in how they have been utilised; more productive grasslands have been exploited more, resulting in shifts towards species with more exploitative growth strategies, a narrower range of growth and investment strategies, but a reduction in competitive dominance. The less productive grasslands appear to be utilised less, with shifts towards more conservative growth strategies and reduced habitat filtering, and the potential to deliver greater ecosystem multifunctionality. Resource declines for pollinators appeared to be universal across all grassland habitats.

A growing awareness of the destructive effects of non-native invasive species has led to a massive increase in removal programmes around the world. However, little is generally known about what happens to sites following the removal of the invasives and the implicit assumption that the native community will return, unaided, to pre-invasion conditions is often left untested. We assessed recovery of the native understorey plant community following removal of the non-native invasive Rhododendron ponticum L. from Scottish Atlantic oak woodland. We recorded understorey community composition in sites covering a gradient of increasing R. ponticum density, and across a separate series of sites covering a chronosequence of time since R. ponticum removal. We then compared both of these series to the target community found in uninvaded sites. We also analysed differences in soil chemistry between the sites to test for chemical legacy effects of invasion in the soil. Native understorey cover declined as R. ponticum density increased, with bryophytes dropping to less than a third of the cover present in uninvaded sites and forbs and grasses being completely extirpated under dense stands. Cleared sites showed no evidence of returning to the target community, even after 30 years of recovery, and instead formed a bryophyte-dominated “novel community,” containing few of the typical oak woodland vascular plants. Contrary to expectation, soil pH, C:N ratio and nutrient concentrations (N, P, K, Ca and Mg) were not affected by the invasion of R. ponticum, and chemical legacy effects in the soil were not responsible for the failure of the native community to revert to pre-invasion conditions. Instead, we hypothesise that the rapid formation of a bryophyte mat, coupled with the often substantial distances to potential seed sources, hindered vascular plant recolonisation. Synthesis and applications. Clear evidence of invasion history can be detected in the understorey plant community even decades after the successful removal of the invasive Rhododendron ponticum L. This finding demonstrates that native communities may be unable to recover effectively of their own accord following invasive species removal, and will require further management interventions in order to achieve restoration goals.

The accumulation of biodiversity in space and time has been modelled extensively using the species–area relationship and the species–time relationship, respectively. Recently, these models have been combined into time–area curves in order to investigate spatiotemporal scaling of species richness. This study expands on previous research by applying these spatiotemporal models to functional diversity. Understanding spatiotemporal dynamics of ecological traits is important due to their crucial role in ecosystem functioning and mediating species responses to environmental change. We present a new function based on the semi-logarithmic species–area relationship, which was applied with a power function to vegetation survey data from Scottish machair grassland for both species richness and two measures of functional diversity. When taking a whole-study approach using non-linear mixed effects models, the semi-logarithmic function used here shows a positive time–area interaction for species richness, contrasting with the negative interaction of the power law found in previous investigations. Although there was a negative time–area interaction for functional diversity measures at the whole-study scale, parameter estimates were inconsistent at the individual site level. Overall, the results reveal differing spatiotemporal dynamics of species and their traits and suggest that the appropriate scale for space-for-time substitutions depends on the aspect of biodiversity being investigated. The new model developed in this study, and the novel application to functional diversity, opens up future possible research into spatiotemporal dynamics of biodiversity.

The soil seed bank plays an important role in determining what plant species emerge following the removal of monodominant invasive species. A depleted seed bank may provide a substantial barrier to site restoration, however, little is known about what changes occur in the seed bank during invasion and following clearance. This study used greenhouse germination trials to quantify and compare the seed bank present in 30 Scottish Atlantic oak woodland sites under three treatments: (1) sites featuring dense stands of the non-native invasive shrub Rhododendron ponticum; (2) sites that had been previously subject to dense R. ponticum stands but which had been cleared; and (3) uninvaded control sites. Seed banks of densely invaded and cleared sites had significantly lower species richness than those of uninvaded control sites with a lower abundance of graminoid, and to a lesser extent forb seeds than the uninvaded controls. The seed bank community composition differed significantly between the three site types. Uninvaded sites featured a wide array of species, densely invaded sites were dominated by R. ponticum seeds and cleared sites were dominated by birch (Betula sp.) seeds. Cleared sites contained very few R. ponticum seeds indicating that once effective clearance had been achieved, re-invasion would be unlikely to occur from the soil seed bank. Our findings revealed that the soil seed bank present in cleared sites was very different from the seed bank of uninvaded control sites, with clear implications for site restoration.