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Aim To examine the phylogeography and population structure of three dung beetle species of the genus Trypocopris (Coleoptera, Geotrupidae). We wanted to test whether genetic differences and genealogies among populations were in accordance with morphologically described subspecies and we aimed to establish times of divergence among subspecies to depict the appropriate temporal framework of their phylogeographical differentiation. We also wished to investigate the historical demographic events and the relative influences of gene flow and drift on the distribution of genetic variability of the different populations.Location Europe (mostly Italy).Methods We collected adult males from dung pats from 15 Italian localities over the period [2000][2001][2002]. For sequence analysis, some dried specimens from Albania, Croatia, Slovakia and Spain were also used. We applied cytochrome oxidase I mitochondrial DNA sequencing and the amplified fragment length polymorphism (AFLP) technique to determine whether phylogeographical patterns within the three species support the proposed hypotheses of subspecies designations, and to detect further structure among populations that might mediate diversification. Results and main conclusionsThe results show a high concordance between the distribution of mtDNA variation and the main morphological groups recognized as subspecies, which thus may represent independent evolutionary units. The degree of mitochondrial divergence suggests that speciation events occurred during the Pliocene, while diversification of the main subspecific lineages took place in the Pleistocene, from c. 0.3 to 1.5 Ma. Mitochondrial and nuclear data also reveal that there is phylogeographical structuring among populations within each of the main groups and that both contemporary and historical processes determined this pattern of genetic structure. Geographical populations form monophyletic clades in both phylogenetic and network reconstructions. Despite the high levels of intrapopulational diversity, F ST values indicate moderate but significant genetic differentiation among populations, and a Bayesian clustering analysis of the AFLP data clearly separates the geographical populations. Nucleotide and gene diversity estimates reveal interspecific differences in the degree of diversification among populations that may be related to the different ecological requirements of the three species.

Distress signals produced by dung beetles of the genus Trypocopris (Coleoptera, Geotrupidae) were analysed to test whether interspecific and intraspecific acoustic variability are species‐ and subspecies‐specific and to ascertain to what extent bioacoustic parameters depend upon the morphology of the stridulatory organs (pars stridens). Bioacoustic analyses showed that the three species were clearly differentiated, despite the fact that disturbance stridulations presented the same stereotyped spectrographic pattern. Within each species, most of the subspecies and populations considered were also bioacoustically distinguishable. Subspecies and populations within each species were differentiated with regard to body size and stridulatory organ, and the length of the pars stridens was positively correlated with the width of the coxa, in turn positively correlated with body size. A few spectrographic measures were significantly constrained by the morphology of the stridulatory apparatus; in particular the duration of sound emission was positively correlated with the length of the apparatus and the sub‐pulse rate was negatively related to the distance between two consecutive crests. For T. pyrenaeus, with the largest number of populations sampled, there was no significant correlation between morphological and spectrographic distances, but there were significant positive correlations between morphological and geographical distances and between morphological and genetic distances. It is hypothesized that genetic differentiation might directly affect variability of the stridulatory apparata which would also be indirectly influenced and constrained by external morphological traits (like the width of the coxa and body size). Stridulatory organs, in turn, would affect the ways a few stridulatory traits (especially the temporal ones) change in time and space.