Summary 1.Calcareous grasslands are communities of high conservation value, often characterized by high plant species richness. These grasslands have experienced a major decline in area throughout Europe, principally resulting from agricultural intensification. Although they have been the focus of extensive previous research, few attempts have been made to examine the long-term dynamics of multiple communities at the landscape scale. 2. To assess long-term change in the structure and composition of a calcareous grassland metacommunity, 88 extant sites first surveyed by R. Good in the 1930s were resurveyed in 2009. Values of a-, b-and c-diversity were compared between the two surveys, using a one-way analysis of similarity (ANOSIM) and non-metric multidimensional scaling. Elements of metacommunity structure (EMS) analysis was used to identify metacommunity structure, and changes in metacommunity composition were related to plant traits. 3. Analyses indicated that a-diversity increased over time, with mean (±SD) species richness per site increasing from 29.31 ± 7.65 in the 1930s to 40.18 ± 16.41 in 2009. No change in b-diversity was recorded. However, c-diversity increased, with the total number of species rising from 219 in the 1930s to 280 in 2009. Species composition shifted over time, associated with a decline in 'stress-tolerant' species typical of species-rich calcareous grasslands, and an increase in species typical of mesotrophic grasslands. This was associated with an increase in mean Ellenberg N value, suggesting that eutrophication has been a driver of floristic change. 4. Elements of metacommunity structure analysis indicated that the structure of this grassland plant metacommunity was Clementsian at both survey times, indicating species sorting. Metacommunity structure was stable over time, despite changes in a-and c-diversity. Analysis of potential structuring mechanisms revealed a significant influence of elevation. 5. Synthesis. This investigation provides a rare example of the long-term dynamics of a plant metacommunity. Results indicate that substantial change has occurred in the composition of calcareous grasslands during this time, both at local and regional scales. The investigation provides evidence of the impact of environmental change on immigration and extinction processes operating in calcareous grasslands at different scales, and highlights challenges for their future conservation.

Analysis of long-term vegetation change is limited. Furthermore most studies evaluating change only examine two snapshots in time, which makes it difficult to define rates of change and accurately assess potential drivers. To assess longterm change in calcareous grassland over multiple time periods, we re-surveyed a transect study undertaken at Parsonage Down National Nature Reserve, Wiltshire, southern England in 1970 and 1990 by T. Wells. We examined differences in soil properties and species traits in each of the survey years to understand potential drivers of vegetation change, including nitrogen deposition and grazing management. There was a clear shift in species composition, combined with significant declines in species richness and diversity between 1970 and 2016, with the greater rate of change occurring between 1990 and 2016. A significant increase in soil total nitrogen was found, which was significantly associated with the decline in species diversity between 1970 and 1990. Significant changes in community-weighted mean traits were identified for plant height (increasing), specific leaf area (decreasing), grazing tolerance (decreasing) and Ellenberg N (decreasing) between 1970 and 2016. By using survey data from multiple time periods, we suggest that N deposition may have contributed Communicated by Zoltan Nagy.Electronic supplementary material The online version of this article (https://doi.().,-volV) ( 01234567 89().,-volV) towards community changes between 1970 and 1990, as indicated by the change in soil properties and the associated decline in species diversity. Vegetation change between 1990 and 2016 is likely to be largely attributable to a decline in grazing pressure, indicated by the increase in taller species and a decrease in grazing tolerance.

The extinction debt, delayed species extinctions following landscape degradation, is a widely discussed concept. But a consensus about the prevalence of extinctions debts is hindered by a multiplicity of methods and a lack of comparisons among habitats. We applied three contrasting species-area relationship methods to test for plant community extinction debts in three habitats which had different degradation histories over the last century: calcareous grassland, heathland and woodland. These methods differ in their data requirements, with the first two using information on past and current habitat area alongside current species richness, whilst the last method also requires data on past species richness. The most data-intensive, and hence arguably most reliable method, identified extinction debts across all habitats for specialist species, whilst the other methods did not. All methods detected an extinction debt in calcareous grassland, which had undergone the most severe degradation. We conclude that some methods failed to detect an extinction debt, particularly in habitats that have undergone moderate degradation. Data on past species numbers are required for the most reliable method; as such data are rare, extinction debts may be under-reported.

Assessing habitat loss effects on biodiversity is a major focus of ecological research. The relationship between habitat amount and biodiversity, postulated in the habitat amount hypothesis, is usually assessed at one point in time, which does not account for habitat loss as a temporal process. We examined habitat amount effects at two time periods, 1930s and 2010s, using plant data from three semi-natural habitats: calcareous grassland, heathland and broadleaved woodland, across Dorset, southern England. Woodlands, which changed little in area over the time period, showed minimal effects of habitat amount on species occurrence in both time periods. For grassland and heathland, which had undergone severe losses over the study period, we found the expected positive relationship in the 2010s, but the relationship was negative for these habitats in the 1930s. We explored possible reasons for this result. Total perimeter-to-area ratio (TPAR) showed positive effects in the 1930s for grassland and heathland, suggesting effects of habitat configuration, specifically edge. However, TPAR was highly correlated with habitat amount so this finding is speculative. One possible explanation for the relationships with habitat amount, and the change between the two periods could be the quality of the surrounding matrix. In the 1930s, the landscape was less intensified and was dominated by semi-natural habitats, whereas by the 2010s much had been converted to arable and intensive grasslands. We speculate that species could likely utilise the matrix to a greater degree in the 1930s compared with the 2010s when the matrix was more hostile, thereby decreasing the importance of habitat amount in the 1930s compared with the 2010s. These findings have important implications for conservation, as they show the importance of context (i.e. matrix quality) in determining the relationship between habitat amount and biodiversity.

Historical data on co-occurring taxa are extremely rare. As such, the extent to which distinct co-occurring taxa experience similar long-term patterns in species richness and compositional change (e.g., when exposed to a changing environment) is not clear.Using data from a diverse ecological community surveyed in the 1930s and resurveyed in the 2010s, we investigated whether local plant and insect assemblages displayed cross-taxon congruence-that is, spatiotemporal correlation in species richness and compositional change-across six co-occurring taxa: vascular plants, non-vascular plants, grasshoppers and crickets (Orthoptera), ants (Hymenoptera: Formicinae), hoverflies (Diptera: Syrphidae), and dragonflies and damselflies (Odonata). All taxa exhibited high levels of turnover across the ca. 80-year time period. Despite minimal observed changes at the level of the whole study system, species richness displayed widespread cross-taxon congruence (i.e., correlated temporal change) across local assemblages within the study system. Hierarchical logistic regression models suggest a role for shared responses to environmental change underlying cross-taxon correlations and highlight stronger correlations between vascular plants and their direct consumers, suggesting a possible role for biotic interactions between these groups.These results provide an illustration of cross-taxon congruence in biodiversity change using data unique in its combination of temporal and taxonomic scope, and highlight the potential for cascading and comparable effects of environmental change (abiotic and biotic) on co-occurring plant and insect communities. However, analyses of historical resurveys based on currently available data come with inherent uncertainties.As such, this study highlights a need for well-designed experiments, and monitoring programs incorporating co-occurring taxa, to determine the underlying mechanisms and prevalence of congruent biodiversity change as anthropogenic environmental change accelerates apace.