Allometric relationships and character evolution in Onthophagus taurus (Coleoptera: Scarabaeidae)

Allometric relationships in primary sexual traits (male and female genitalia), secondary sexual traits (male horns and female carinae), and non-sex-related traits (external body traits, epipharynx traits) were studied in the dung beetle Onthophagus taurus. Model II regressions of log-transformed data were used to quantify relationships, with pronotum width as regressor and indicator of overall body size. Slopes (allometric values) for the different trait categories were significantly different, with secondary sexual traits showing the highest values (higher than 1.0), followed by external body traits (slightly lower than 1.00) and epipharynx traits (around 0.2). Primary sexual traits and body size were mostly uncorrelated and genital sizes were virtually constant. Allometries of secondary sexual traits were quite different in the two sexes: the relationship between male horn length and pronotum width was approximately sigmoidal, while that between female carina length and pronotum width was linear. External body traits had significantly higher allometric values in females than in males. Our results suggest that traits in the different categories are under different kinds of selection. Genital allometries can be explained on the basis of sexual selection by cryptic female choice or by the lock-and-key hypothesis. Among secondary sexual characters, male horn morphology seems to be mostly "environmentally" determined and sexual selection would affect only a component of the developmental mechanism of horn expression. External body characters are likely under natural selection, even though a few traits could be sex-related. Finally, in both sexes, internal epipharynx traits seem to be subject to the same selective pressure, probably because males and females use the same feeding niche. The constancy of genital sizes in O. taurus suggests that in developmental processes, more is invested in primary sexual traits (to produce genitalia of the proper size) than in secondary sexual or body traits. Males receiving low quantities of food may incur costs associated with a small horn or small body size, but do not incur costs associated with small genitalia. Females probably share the same developmental pattern.