Spatial heterogeneity ensures long-term stability in vegetation and Fritillaria meleagris flowering in Uppsala Kungsäng, a semi-natural meadow

Semi-natural grasslands are becoming increasingly rare, and their vegetation may be affected by environmental changes and altered management. At Kungsängen Nature Reserve, a wet to mesic semi-natural meadow near Uppsala, Sweden, we analysed long-term changes in the vegetation using data from 1940, 1982, 1995 and 2016. We also analysed the spatial and temporal dynamics in the Fritillaria meleagris population based on countings of flowering individuals in 1938, 1981–1988 and 2016–2021. Between 1940 and 1982 the wet part of the meadow became wetter, which led to an increased cover of Carex acuta and pushed the main area of flowering of F. meleagris up towards the mesic part. Annual variation in the flowering propensity of F. meleagris (in May) was affected by temperature and precipitation in the phenological phases of growth and bud initiation (June in the previous year), shoot development (September in the previous year) and initiation of flowering (March–April). However, the response to weather was in opposite directions in the wet and mesic parts of the meadow, and the flowering population showed large year-to-year variation but no long-term trend. Variation in management (poorly documented) led to changes in different parts of the meadow, but the overall composition of the vegetation, species richness and diversity changed little after 1982. Species richness and species composition of the meadow vegetation, and the long-term stability of the F. meleagris population are maintained by the variation in wetness, highlighting the importance of spatial heterogeneity as an insurance against biodiversity loss in semi-natural grasslands and nature reserves generally.