Summary 1. Species assemblages of diatoms, rotifers, chydorids, planktonic crustaceans and chironomids were studied in 235 alpine lakes in the Alps, Pyrenees, Tatras (Western Carpathians), Retezat (Southern Carpathians) and Rila Mountains (Balkans). 2. For all taxonomic groups we found a hierarchical structure in the community assemblage using distinct scales of lake clustering (number of k‐means groups) based on species composition similarity (Hellinger distance). We determined the optimal partition in assemblage types (i.e. number of lake clusters) for each taxonomic group by maximising the sum of the taxon indicative value (IndVal) and performed discriminant analyses, using environmental variables not conditioned by geographical patterns. Relevant environmental variables differed among and within taxonomic groups. Therefore the assemblages respond to a complex environmental mosaic, with the exception of diatom assemblages, which followed an acid–base gradient. 3. The significant environmental variables could be grouped into four general factors: lake size, tropho‐dynamic status, acid–base balance and ice‐cover duration (i.e., altitudinal gradient). Lake size was significant for the highest number of assemblage types; however, the most significant factor differed among taxonomic groups: acid–base balance for diatoms, lake size for rotifers, ice‐cover duration for chydorids and planktonic crustaceans and tropho‐dynamic status for chironomids. No single environmental typology accounted for the assemblage structure of all taxonomic groups. 4. However, defining ecological thresholds as values within environmental gradients at which the rate of change in assemblages is accelerated relative to points distant from that threshold, we were able to find specific threshold values for each of the four main general environmental factors identified, which were relevant across several taxonomic groups: 3 ha for lake area; 0.6 mg L−1 for dissolved organic carbon; 190 days for ice‐cover duration and 200 μeq L−1 for acid neutralising capacity. Above and below these values ecosystem organisation change substantially. They have direct applications in establishing lake typologies for environmental quality and biodiversity conservation programmes, and in improving predictions about global change impacts.

The scarcity of high-resolution empirical data directly tracking diversity over time limits our understanding of speciation and extinction dynamics and the drivers of rate changes. Here, we analyze a continuous species-level fossil record of endemic diatoms from ancient Lake Ohrid, along with environmental and climate indicator time series since lake formation 1.36 million years (Ma) ago. We show that speciation and extinction rates nearly simultaneously decreased in the environmentally dynamic phase after ecosystem formation and stabilized after deep-water conditions established in Lake Ohrid. As the lake deepens, we also see a switch in the macroevolutionary trade-off, resulting in a transition from a volatile assemblage of short-lived endemic species to a stable community of long-lived species. Our results emphasize the importance of the interplay between environmental/climate change, ecosystem stability, and environmental limits to diversity for diversification processes. The study also provides a new understanding of evolutionary dynamics in long-lived ecosystems.

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Due to the ubiquity and high dispersal capacity of unicellular eukaryotes, their often extraordinary diversity found in isolated and long-lived ecosystems such as ancient lakes is typically attributed to multiple colonization events rather than to in situ speciation. However, respective evolutionary studies are very scarce and the often high number of species flocks in ancient lakes across multicellular taxa raises the question whether unicellular species, such as diatoms, may radiate as well. Here, we use an integrative approach that includes molecular data from benthic diatom species of the genus Aneumastus endemic to ancient Lake Ohrid, fossil data obtained from the sediment record of a recent deep-drilling project and biogeographical information to test if this group, indeed, constitutes a species flock. Molecular-clock and phylogenetic analyses indicate a young monophyletic group of several endemic species. Molecular, fossil and biogeographical data strongly suggest a rapid intralacustrine diversification, which was possibly triggered by the emergence of novel habitats. This finding is the first evidence for a species flock in diatoms and suggests that in situ speciation is also a relevant evolutionary process for unicellular eukaryotes in isolated ecosystems.

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