Ancient origin of endemic Iberian earth-boring dung beetles (Geotrupidae).

The earth-boring dung beetles belong to the family Geotrupidae that includes more than 350 species classified into three subfamilies Geotrupinae, Lethrinae, and Taurocerastinae, mainly distributed across temperate regions. Phylogenetic relationships within the family are based exclusively on morphology and remain controversial. In the Iberian Peninsula there are 33 species, 20 of them endemic, which suggests that these lineages might have experienced a radiation event. The evolution of morphological adaptations to the Iberian semi-arid environments such as the loss of wings (apterism) or the ability to exploit alternative food resources is thought to have promoted diversification. Here, we present a phylogenetic analysis of 31 species of Geotrupidae, 17 endemic to the Iberian Peninsula, and the remaining from southeastern Europe, Morocco, and Austral South America based on partial mitochondrial and nuclear gene sequence data. The reconstructed maximum likelihood and Bayesian inference phylogenies recovered Geotrupinae and Lethrinae as sister groups to the exclusion of Taurocerastinae. Monophyly of the analyzed geotrupid genera was supported but phylogenetic relationships among genera were poorly resolved. Ancestral character-state reconstruction of wing loss evolution, dating, and diversification tests altogether showed neither evidence of a burst of cladogenesis of the Iberian Peninsula group nor an association between apterism and higher diversification rates. Loss of flight did not accelerate speciation rates but it was likely responsible for the high levels of endemism of Iberian geotrupids by preventing their expansion to central Europe. These Iberian flightless beetle lineages are probably paleoendemics that have survived since the Tertiary in this refuge area during Plio-Pleistocene climatic fluctuations by evolving adaptations to arid and semi-arid environments.