The fifth mass extinction event (MEE) at the Cretaceous–Palaeogene (K–Pg) boundary 66 million years ago (Ma) led to massive species loss but also triggered the diversification of higher taxa. Five models have been proposed depending on whether this diversification occurred before, during or after the K–Pg boundary and the rate of species accumulation. While the effects of the K–Pg MEE on vertebrate evolution are relatively well understood, the impact on invertebrates, particularly in freshwater ecosystems, remains controversial. One example is the hyperdiverse Hydrobiidae—the most species‐rich family of freshwater gastropods. Whereas some studies place its origin in the Jurassic or even Carboniferous, most fossil records postdate the K–Pg event. We therefore used robustly time‐calibrated multi‐locus phylogenies of >400 species representing >100 hydrobiid genera to unravel its evolutionary history and patterns of diversification. We found that the family started diversifying shortly after the K–Pg boundary (∼60 Ma; 95% highest posterior density 52–69 Ma). Lineage richness gradually increased to the present and phylogenetic diversity until ∼25 Ma. These findings suggest that diversification was not initially driven by ecological opportunity. Combining the two criteria of timing and rate of diversification, a soft‐explosive diversification model of aquatic vertebrates best fits the patterns observed. We also show that most higher hydrobiid taxa (i.e. subfamilies) diversified from the Middle Oligocene to Middle Miocene (i.e. 12–28 Ma). Two of the 15 major clades delimited are described here as new subfamilies (i.e. Bullaregiinae n. subfam. and Pontobelgrandiellinae n. subfam.), whose members are restricted to subterranean waters. Our results are an important contribution to understanding how the fifth MEE has shaped evolution and patterns of biodiversity in continental aquatic systems. Given the high extinction risks faced by many hydrobiids today, they also emphasise the need to study the biodiversity of vulnerable ecosystems.

Shell features are key factors for systematics and evolutionary biology studies of freshwater gastropods. Based mostly on shell morphology, 19 species of the springsnail genus Radomaniola (family Hydrobiidae) have been described. Although the scarce differentiation of various shell dimensions suggests morphological stasis, the evolutionary dynamics of shell and other anatomical structures have not yet been explored fully. By inferring a phylogeny and a time-calibrated species tree from mitochondrial (COI and 16S) and nuclear (28S) sequences of 15 recognized species, we initially examine the species diversity of the Radomaniola dataset and then, through phylogenetic comparative methods, assess the evolutionary correlation, pace and mode of 40 continuous shell and anatomical characters. By synthesizing the results of four species delimitation methods and through morphological examinations, we recognize 21 groups, for which taxonomy is discussed here. Seven new species are described. We reveal a high degree of correlated evolution between characters of the shell, which are constrained by a single morphological optimum, consistent with the morphological stasis model. Anatomical traits diverged rapidly in an unconstrained manner or toward multiple optima. These findings indicate that in order to understand the morphological evolution of springsnails, it is essential to examine different organs in detail.

Phylogenetic analyses of the highly diverse (non-marine aquatic) gastropod family Hydrobiidae Stimpson, 1865 have revealed seven main lineages, most of which represent subfamilies. The subfamily Pseudamnicolinae Radoman, 1977, and specifically the genus Pseudamnicola Paulucci, 1878 (mainly inhabiting western and central Mediterranean regions), contributes substantially to this hydrobiid richness. Most of its congeners have been described in terms of their shell and penis features, which are of limited diagnostic value. Hence, the taxonomic status of some Pseudamnicola species needs to be revised, particularly of those inhabiting marginal regions, such as the Ponto-Caspian domain, largely occupied by the subfamily Pyrgulinae Brusina, 1882. Here we present a molecular phylogeny including species of both subfamilies along with extended morphological descriptions to confirm assignments of the Iranian species Pseudamnicola zagrosensis Glöer & Pešić, 2009; Sarkia kermanshahensis Glöer & Pešić, 2009 (originally within Pseudamnicola) and P. saboori Glöer & Pešić, 2009. Our COI-based tree rejects these assignments suggesting a new potential lineage, sister to the pyrgulinid species, and comprising three genera: Shadinia Akramowski, 1976, Intermaria gen. nov. and Persipyrgula gen. nov. These genera differ molecularly by 3.6%–8.5%, and are diagnosable by penis, female genitalia and radula features. Our findings evidence the high morphological variability of pyrgulinid species and provide insight into the origins and evolution of the freshwater Ponto-Caspian fauna.