Trade-off between horns and other functional traits in two Onthophagus species (Scarabaeidae, Coleoptera)

Beetle horns are extraordinarily diversified secondary sexual structures used for mate choice and male-male combat. Due to an interaction of nutritional, hormonal and genetic factors, their polyphenic development is metabolically expensive and occurs in the virtually closed system of the pre-pupal stage, after the developing larva has stopped feeding. Previous studies showed the occurrence of resource competition resulting in a trade-off between horns and other morphological structures. These studies also revealed functional associations between autoecology and horns, as a function of their physical location (i.e. head versus pronotum), and suggested that constraints imposed by trade-offs on adult morphology may have profound evolutionary consequences, such as ecological and reproductive isolation. In this study, we compared trade-off patterns between horns and other functional traits (eyes, antennae, legs, head, epipharynges and genitalia) in two congeneric species bearing horns located in the same anatomical area, but with different morphologies. Specifically, we considered Onthophagus taurus, characterised by a pair of long, lunated cephalic horns, and Onthophagus fracticornis, expressing a single cephalic horn. We demonstrated that, even when horns are located in the same physical position on the insect's body, differences in horn morphology can bring about differences in how functional traits respond to horn investment. These differences are interpretable in the light of the hierarchy of functions carried out by these structures and their component parts in each species.