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ABSTRACT The vertebrate skull is a complex structure, and studies of skull shape have yielded considerable insight into the evolutionary forces shaping specialized phenotypes in organisms as diverse as bats, frogs, and fossorial animals. Here, we used phylogenetic comparative analyses of CT scans of male skulls from 57 species ofSceloporuslizards to explore patterns of skull evolution in a group of generalist taxa. We found that most interspecific variation is in terms of skull elongation such that some species have long, narrow skulls, whereas others exhibit more compact and robust skulls. We also found strong links to overall body size, with evolutionary shifts to larger bodies being associated with more compact skulls and slower evolutionary rates. This is the opposite of the pattern in most mammals in which larger bodied species have longer snouts, and more like the pattern in frogs in which function has played a more important evolutionary role. Also, unlike other vertebrates, the jaw, anterior, and posterior parts of theSceloporusskull are largely integrated, having evolved independently of each other only to a limited, albeit significant, degree. Our results emphasize the importance of body size in the evolutionary shaping of the skull and suggest that additional studies of behavioral function in a generalist group are warranted. 

Abstract Single substances within complex vertebrate chemical signals could be physiologically or behaviourally active. However, the vast diversity in chemical structure, physical properties and molecular size of semiochemicals makes identifying pheromonally active compounds no easy task. Here, we identified two volatile cyclic dipeptides, cyclo(L-Leu-L-Pro) and cyclo(L-Pro-L-Pro), from the complex mixture of a chemical signal in terrestrial vertebrates (lizard genusSceloporus), synthesised one of them and investigated their biological activity in male intra-specific communication. In a series of behavioural trials, lizards performed more chemosensory behaviour (tongue flicks, lip smacks and substrate lickings) when presented with the synthesised cyclo(L-Pro-L-Pro) chemical blend, compared to the controls, the cyclo(L-Leu-L-Pro) blend, or a combined blend with both cyclic dipeptides. The results suggest a potential semiochemical role of cyclo(L-Pro-L-Pro) and a modulating effect of cyclo(L-Leu-L-Pro) that may depend on the relative concentration of both compounds in the chemical signal. In addition, our results stress how minor compounds in complex mixtures can produce a meaningful behavioural response, how small differences in structural design are crucial for biological activity, and highlight the need for more studies to determine the complete functional landscape of biologically relevant compounds.