Condition-dependent pheromone signaling by male rock lizards: more oily scents are more attractive.

Pheromones of vertebrates are often a mixture of several chemicals with different properties and messages, and their production seems condition dependent. Thus, pheromones are a good, but little studied, example of multiple sexual signals. Femoral gland secretions of male rock lizards Iberolacerta cyreni contain steroids that may act as pheromones, but there are also many other lipids, such as oleic acid, whose allocation to secretions may be costly because it has to be diverted from body fat reserves. This suggests that oleic acid could also have some function in secretions. Chemical analyses showed that proportions of oleic acid in femoral secretions of males were positively related to body condition of males, suggesting that the oleic acid secreted may reflect the amount of body fat reserves of a male. Tongue-flick bioassays showed that females were able to detect by chemosensory cues alone differences in proportions of oleic acid in secretions of males. Scents of males with more oleic acid elicited stronger chemosensory responses by females. Further tests with chemical standards confirmed that females distinguished oleic acid, and changes in its concentration, from other chemicals that are naturally found in secretions of males. Moreover, choice trials of scent-marked substrates showed that females were more attracted to areas that were experimentally manipulated to increase the proportion of oleic acid in natural scent marks of males. We suggest that oleic acid in femoral secretions might be a reliable advertisement of a male's body condition, which females could use to select high-quality mates in conjunction with information provided by other chemicals. Alternatively, scent marks with more oleic acid might be simply more attractive to females if chemosensory responses of females to scent of males were originated by a preexisting sensory bias for food chemicals such as the oleic acid. Nevertheless, this sensory trap might have evolved into an honest signal because the elaboration of the signal seems differentially costly for males with different body conditions.