‘Carabidologists do it all’ (Niemelä 1996a) is a phrase with which most European carabidologists are familiar. Indeed, during the last half a century, professional and amateur entomologists have contributed enormously to our understanding of the basic biology of carabid beetles. The success of the field is in no small part due to regular European Carabidologists’ Meetings, which started in 1969 in Wijster, the Netherlands, with the 14th meeting again held in the Netherlands in 2009, celebrating the 40th anniversary of the first meeting and 50 years of long-term research in the Dwingelderveld. This paper offers a subjective summary of some of the major developments in carabidology since the 1960s. Taxonomy of the family Carabidae is now reasonably established, and the application of modern taxonomic tools has brought up several surprises like elsewhere in the animal kingdom. Progress has been made on the ultimate and proximate factors of seasonality and timing of reproduction, which only exceptionally show non-seasonality. Triggers can be linked to evolutionary events and plausibly explained by the “taxon cycle” theory. Fairly little is still known about certain feeding preferences, including granivory and ants, as well as unique life history strategies, such as ectoparasitism and predation on higher taxa. The study of carabids has been instrumental in developing metapopulation theory (even if it was termed differently). Dispersal is one of the areas intensively studied, and results show an intricate interaction between walking and flying as the major mechanisms. The ecological study of carabids is still hampered by some unresolved questions about sampling and data evaluation. It is recognised that knowledge is uneven, especially concerning larvae and species in tropical areas. By their abundance and wide distribution, carabid beetles can be useful in population studies, bioindication, conservation biology and landscape ecology. Indeed, 40 years of carabidological research have provided so much data and insights, that among insects - and arguably most other terrestrial organisms - carabid beetles are one of the most worthwhile model groups for biological studies.

BackgroundThe Alpine region harbours one of the most diverse subterranean faunas in the world, with many species showing extreme morphological modifications. The ground beetles of tribe Trechini (Coleoptera, Carabidae) are among the best studied and widespread groups with abundance of troglobionts, but their origin and evolution is largely unknown.ResultsWe sequenced 3.4 Kb of mitochondrial (cox1, rrnL, trnL, nad1) and nuclear (SSU, LSU) genes of 207 specimens of 173 mostly Alpine species, including examples of all subterranean genera but two plus a representation of epigean taxa. We applied Bayesian methods and maximum likelihood to reconstruct the topology and to estimate divergence times using a priori rates obtained for a related ground beetle genus. We found three main clades of late Eocene-early Oligocene origin: (1) the genus Doderotrechus and relatives; (2) the genus Trechus sensu lato, with most anisotopic subterranean genera, including the Pyrenean lineage and taxa from the Dinaric Alps; and (3) the genus Duvalius sensu lato, diversifying during the late Miocene and including all subterranean isotopic taxa. Most of the subterranean genera had an independent origin and were related to epigean taxa of the same geographical area, but there were three large monophyletic clades of exclusively subterranean species: the Pyrenean lineage, a lineage including subterranean taxa from the eastern Alps and the Dinarides, and the genus Anophthalmus from the northeastern Alps. Many lineages have developed similar phenotypes independently, showing extensive morphological convergence or parallelism.ConclusionsThe Alpine Trechini do not form a homogeneous fauna, in contrast with the Pyrenees, and show a complex scenario of multiple colonisations of the subterranean environment at different geological periods and through different processes. Examples go from populations of an epigean widespread species going underground with little morphological modifications to ancient, geographically widespread lineages of exclusively subterranean species likely to have diversified once fully adapted to the subterranean environment.

Faille, A., Casale, A. & Ribera, I. (2010). Phylogenetic relationships of Western Mediterranean subterranean Trechini groundbeetles (Coleoptera: Carabidae). —Zoologica Scripta, 40, 282–295. Carabid beetles of tribe Trechini (Coleoptera) are one of the main groups of insects that colonized the subterranean environment. Many species of this group have developed similar morphological modifications related to the subterranean life, resulting in a characteristic Aphaenops‐like phenotype that obscures their phylogenetic relationships (depigmented, blind, elongated body and appendages, narrow head and pronotum). We present here the result of a molecular study using a combination of nuclear (small ribosomal unit, large ribosomal unit) and mitochondrial (cox1, cyb, rrnL, trnL, nad1) genes to investigate the phylogenetic placement of the highly modified subterranean genera of the tribe Trechini from the west Mediterranean area (France, Spain, Morocco and Sardinia). Our results confirm the multiple independent origin of troglomorphism among these genera, and reveal a pattern largely determined by geographical proximity. We discuss the validity of some groups proposed on the base of morphological features, and provide estimates of divergence between subterranean genera and other groups of Trechini, including epigean species of the same area. We compare the estimated age for the origin of the main groups resulting from two different calibrations, using one the standard mitochondrial mutation rate (2.3% divergence per Myr) and the other the separation between Sardinia and mainland 33 Ma. Under the first scenario, the main groups of genera would have a late Miocene origin, with a subsequent colonization of north Africa at the Pliocene–Pleistocene boundary. The assumption that the main groups originated through vicariance due to the separation of the Sardinian plate in the Oligocene results in a Messinian origin of the north African subterranean taxa, and a global mitochondrial rate reduced to 1% divergence per Myr.

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Aim Current climate, biotic habitat provision and historical events are known drivers of diversity patterns. However, these three factors are seldom evaluated together. Here, we study the influence of climate, the distribution of deciduous forests and Pleistocene climate changes on the species diversity of Carabus ground beetles in Europe. Location Continental Europe.Methods We used geographically weighted regressions (GWR) to explore geographical variation in the relationship between species richness and current climate in a spatially explicit context. Further, we analysed simultaneously the network of relationships among current temperature, climatic variability since the Last Glacial Maximum (LGM) and the distribution of deciduous forests through structural equation models (SEM). Also, we assessed dissimilarity in the composition of European faunas by means of beta diversity metrics related with true spatial replacement and nestedness. ResultsWe find that Carabus richness patterns are, at least in part, influenced by water-energy dynamics. However, the effects of current climate are also shaped by Pleistocene glaciations, as water and energy variables change in importance at the southern limits of the ice sheet during the LGM. Accordingly, this border results in abrupt shifts in the relative importance of both (1) current and past climate correlates on Carabus richness, and (2) nestedness and true turnover on the compositional changes among their assemblages. Moreover, we also detect a direct effect of the geographical distribution of deciduous forests on Carabus species richness in both northern and southern regions. Main conclusionsOur results confirm that the processes shaping diversity patterns may depend on the history of particular regions. While Carabus richness seems to be largely driven by current climate in southern Europe, in the north it appears to be more affected by the imprint of past climates. Our findings also suggest that the areas of influence of Pleistocene glaciations may depend on the idiosyncratic characteristics of particular taxa.

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