Optimal Defense Theory in an ant–plant mutualism: Extrafloral nectar as an induced defence is maximized in the most valuable plant structures

Plants allocate defences in order to decrease costs and maximize benefits against herbivores. The Optimal Defense Theory (ODT) predicts that continuously expressed (i.e. constitutive) defences are expected in structures of high value, whereas defences that are expressed or that increase their expression only after damage or upon risk of damage (i.e. induced defences) are expected in structures of low value. Although there are several studies evaluating ODT predictions, few studies have successfully tested them as a way of measuring ecological investment in extrafloral nectary (EFN)-mediated ant–plant interactions. Here we compared extrafloral nectar production and ant attractiveness to EFNs located on vegetative versus reproductive plant structures on Qualea multiflora plants subjected to different levels of simulated herbivory. We then addressed the following predictions emerging from the ODT: (a) extrafloral nectar produced in inflorescence EFNs will have higher volumes and calories and will attract more ants than extrafloral nectar produced in leaf EFNs; (b) extrafloral nectar production (volume and calories) and ant attendance will increase after simulated herbivory in leaf EFNs but not in inflorescence EFNs; (c) higher simulated leaf herbivory will induce higher extrafloral nectar production in EFNs on leaves and (d) more attractive extrafloral nectar (higher volume and calories) will attract more ants. Extrafloral nectar volume and calorie content, as well as ant abundance, were higher in EFNs of inflorescences compared to EFNs of leaves both before and after simulated herbivory, consistent with one of our predictions. However, EFNs on both leaves and inflorescences, not on leaves only, were induced by simulated herbivory, a pattern opposite to our prediction. Plants subjected to higher levels of leaf damage produced more and higher calorie extrafloral nectar, but showed similar ant abundance. Finally, more attractive extrafloral nectar attracted more ants. Synthesis. Our results show that extrafloral nectar production before and after simulated herbivory, as well as ant recruitment, varies according to the plant structure on which EFNs are located. Our study is the first to show that ant recruitment via extrafloral nectar follows predictions from Optimal Defense Theory, and that the ant foraging patterns may be shaped by the plant part attacked and the level of damage it receives.