Latitudinal cline in weapon allometry and phenology of the European stag beetle

Animal body size commonly exhibits a remarkable variation in response to environmental conditions. Latitude, when correlated with temperature, rainfall and seasonality, represents one of the main determinants of variation in body size, as well as in allometry. It has long been recognised that populations of larger body size are found in colder environments (Bergmann's Rule), a cornerstone of evolutionary ecology. However, the way in which latitude might influence investment in exaggerated weapons of animals has received little attention. The European stag beetle Lucanus cervus (Linnaeus, 1758) is the focus of this study. Males of this species exhibit exaggerated mandibles, mainly used as weapons during intra-sexual conflicts. Five populations ranging from northern Italy to the southern limit of the distribution of L. cervus were analysed. Combining morphological and phenological data, latitudinal variation in body size, weapon investment and activity period of the adults were evaluated. The analysis of the allometry of mandibles strongly supported the presence of two male morphs. Large males (major morph) invest significantly more in weapons compared to males of the minor morph. Consistent with Bergmann's Rule, these results confirmed that the stag beetle body size increased at higher latitudes (N) and that this increase in size triggers an arms race which leads to further exaggeration of male weapons which is particularly evident in major males. In this morph, the mandible allometric coefficient line was steeper for the northern populations. The activity period also varied with latitude, beginning later at lower latitudes. Characterisation and comparison of adult phenologies provide valuable data to be used in the design of monitoring programmes for this threatened species and are important for modelling the species responses to climate change.