Aims: Ecological theories predict that assembly processes shape communities so that co-existing species may either be functionally more dissimilar ("divergence") or more similar ("convergence") than expected by chance. Two important factors that are rarely considered in combination are spatial scale and successional stage. Our aim is to identify different functional patterns during succession across spatial scales and to discuss the likely underlying assembly processes. We expect to find convergence due to environmental filtering at early succession, especially at the largest scales, and a tendency towards divergence driven by competitive interactions as succession unfolds and at smaller spatial scales. Location: Protected Landscape Area "Bohemian Karst", Czech Republic. Methods:We studied three succession stages (two, eight and 55 years after abandonment). Each field was sampled using 40 1-m 2 quadrats containing 100 pins in a regular grid. We analysed trait dispersion at several combinations of grain and extent and characterised trait similarity using phylogeny and four traits: specific leaf area, leaf dry matter content, height, and seed mass analysed individually and combined using Gower distance (multi-trait dissimilarity). Results:In the youngest field, three of the traits individually and the multi-trait dissimilarity showed convergence regardless of scale. Phylogeny showed convergence at most spatial scales of the youngest stage. The other two fields showed greater divergence predominantly in the oldest field at the smallest spatial scale. Conclusions:The results are mainly congruent with theoretical expectations and provide a valuable example of directional changes from trait convergence to divergence along succession. The scale-invariant convergence in the early stages of succession seems to be the result of environmental filtering and weaker competitive exclusion. At later stages and particularly at smaller scales, divergence becomes more common, likely because competition limits the trait similarity of species.These results highlight the importance of predictable (non-random) changes in ecological succession.

As the almost 200-year palaeontological research revealed, the geographical distribution of various fossil mollusk faunas in deposits of the late Neogene Lake Pannon displays a regular pattern. The lake basin was filled by lateral accretion of sediments, resulting in condensed sedimentary successions in the distal parts of the basin and successively younger shallow-water deposits from the margins towards the basin center. Exposed intra-basin basement highs, however, broke this strict pattern when they acted as sediment sources during the lake’s lifetime. The Mecsek Mts in southern Hungary was such an island in Lake Pannon during the early late Miocene. Deposition of the 200 m thick Sarmatian–Pannonian sedimentary succession in Pécs-Danitzpuszta at the foot of the Mecsek Mts was thus controlled by local tectonic and sedimentary processes, resulting in a unique succession of facies and mollusk faunas. A typical, restricted marine Sarmatian fauna is followed by a distinct freshwater or oligohaline interval, which, according to micropalaeontological evidence, still belongs to the Sarmatian. Although poor preservation of fossils does not allow firm conclusions, it seems that freshwater Sarmatian snails were the ancestors of the brackish-water-adapted early Pannonian pulmonate snail taxa. The successive “Sarmatian-type” dwarfed cockle fauna is similar to those widely reported from the Sarmatian–Pannonian boundary in various parts of the Pannonian Basin; however, a thorough taxonomic study of its species is still lacking. The bulk of the sedimentary succession corresponds to the sublittoral to profundal “white marls,” which are widespread in the southern Pannonian Basin. In Croatia and Serbia, they are divided into the Lymnocardium praeponticum or Radix croatica Zone (11.6–11.4 Ma) below, and the Congeria banatica Zone (11.4–9.7 Ma) above; this division can be applied to the Pécs-Danitzpuszta succession as well. Sedimentation of the calcareous marl, however, ceased at Pécs-Danitzpuszta at about 10.5–10.2 Ma ago (during the younger part of the Lymnocardium schedelianum Chron), when silt was deposited with a diverse sublittoral mollusk fauna. Similar faunas are known from the Vienna Basin, southern Banat, and other marginal parts of the Pannonian Basin System, but not from Croatia and Serbia, where deposition of the deep-water white marls continued during this time. Finally, the Pécs-Danitzpuszta succession was capped with a thick, coarse-grained sand series that contains mollusk molds and casts representing a typical littoral assemblage. This littoral fauna is well-known from easternmost Austria, northern Serbia, and northwestern Romania, but never directly from above the sublittoral L. schedelianum Zone. The fauna is characteristic for the upper part of the Lymnocardium conjungens Zone and has an inferred age of ca. 10.2–10.0 Ma. The Pécs-Danitzpuszta succession thus allows to establish the chronostratigraphic relationship between mollusk faunas that have not been observed in one succession nor in close proximity to each other in other parts of the Pannonian Basin.