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The global diversity of True Water Beetles, False Water Beetles and Phytophilous Water

External and internal head structures of adults of Orchymontiinae, Prosthetopinae, Hydraeninae and Ochthebiinae were studied and those of Ochthebius semisericeus and Limnebius truncatellus are described in detail. The results are evaluated with respect to their relevance for a reconstruction of hydraenid phylogeny and also compared with structural features found in adults of other staphyliniform families. The monophyly of Hydraenidae is supported by the presence of a plate-like, trilobed premento-hypopharyngeal extension, an unusual origin of m. tentoriohypopharyngalis, dorsal tentorial arms firmly fused with the head capsule, modified basal antennomeres, and palpigers connected by a transverse sclerotized bar. Orchymontiinae are monophyletic and the basal sister group of the remaining Hydraenidae. The presence of a ventral transverse genal bulge and of a pubescent antennal club with more than two antennomeres (reversal in some prosthetopines: e.g. Mesoceration abstrictum) are possible apomorphies of Hydraenidae excluding Orchymontiinae. Prosthetopinae are probably monophyletic and the sister group of Ochthebiinae + Hydraeninae. The latter clade is characterized by a distinct cupula formed by antennomere VI, a loose five-segmented pubescent antennal club, and a modified antennal musculature. The presence of an unusual tentorio-pharyngeal dilator is a shared derived feature of Ochthebiinae and the genus Davidraena. The monophyly of Ochthebiinae was confirmed and Ochtheosus is the sister group of the remaining ochthebiine genera, which are characterized by a perforated wall-like structure formed by the posterior tentorial arms. The absence of this tentorial modification and the fimbriate galea are plesiomorphies retained in Ochtheosus. Calobius differs strongly from other subgenera of Ochthebius and a generic status may be appropriate. The monophyly of Hydraeninae is not supported. Hydraena was confirmed as a clade and Laeliaena and Limnebius are sister groups. The latter genus is characterized by several autapomorphies. The basal position of Orchymontiinae and Prosthetopinae suggests a Gondwanan origin of Hydraenidae and a primary preference for life in running water. Important evolutionary changes of head structures are complex transformations of the antennae and related structures. Yet, the use of the antennae as accessory breathing organs is not a groundplan feature of the family. The results of this study strengthen the case of staphylinoid affinities of Hydraenidae.

Aim We test whether species of western Mediterranean aquatic Coleoptera of the 'Haenydra' lineage (Hydraenidae, Hydraena) originated through: (1) successive periods of dispersal and speciation, (2) range fragmentation by random vicariance, or (3) range fragmentation by geographic isolation owing to a general reduction of population density.Location Europe. MethodsTo discriminate between scenarios we use contrasting predictions of the relationship between phylogenetic and geographic distance. The phylogeny was based on 3 kb of four mitochondrial and two nuclear gene fragments of about half of the known species of 'Haenydra', including most western Mediterranean taxa. Divergences were estimated using a molecular clock. The relationship between phylogenetic and geographic distance was tested using bivariate plots, Mantel tests and comparison of the observed phylogeny with the one minimizing geographic distances between species, as measured using Euclidean minimum spanning trees (EMSTs). ResultsThe monophyly of 'Haenydra' was strongly supported, although its phylogenetic placement was not resolved. 'Haenydra' was estimated to be of late Miocene age, with most species originating during the Pleistocene. In two clades (Hydraena tatii and Hydraena emarginata clades) there was a significant association between geographic and phylogenetic distance, and the reconstructed phylogeny was identical to that obtained through the EMST, demonstrating a strong non-randomness of the geographic distribution of the species. In two other clades (Hydraena iberica and Hydraena bitruncata clades) there was no association between geographic and phylogenetic distance, and the observed phylogeny was not the one minimizing geographic distances. In one of the clades this seems to be due to a secondary, recent range expansion of one species (H. iberica), which erased the geographic signal of their distributions.Main conclusions We show that it is possible to obtain strong evidence of stasis of the geographic ranges of narrow-range endemic species through the study of their phylogenetic relationships and current distributions. In at least two of the studied clades, current species seem to have originated through the fragmentation of a more widely distributed species, without further range movements. A process of range expansion and fragmentation may have occurred repeatedly within the 'Haenydra' lineage, contributing to the accumulation of narrow-range endemics in Mediterranean Pleistocene refugia.