Phenotypic plasticity enables many damaged plants to increase nectar secretion rates from extrafloral nectaries (EFNs), or in the case of broad bean,Vicia fabaL., to produce additional EFNs, to attract natural enemies of herbivores. While plants benefit greatly from these defensive mutualisms, the costs of producing EFNs are largely unknown. We hypothesized that if EFN production is costly, then damaged plants with high resource levels would be able to produce more EFNs than plants that are resource-limited. Here, we show that this indirect inducible defence does follow this general pattern.Vicia fabaenriched with 6 or 12 g of 14 : 14 : 14 NPK fertilizer increased EFN numbers after leaf damage by 46 and 60%, respectively, compared with nutrient-poor plants. Thus, EFN production is both damage- and resource-dependent. Analogous to direct defences, production of EFNs may limit the overall loss of leaf tissue when risk of herbivory increases.

By comparing the relative sizes of anatomical structures among phenotypes, selective pressures that shape species' morphologies can be evaluated. Aphids emit droplets containing an alarm pheromone/defensive secretion from unique anatomical structures called cornicles, upon being attacked. As aphids live in colonies of high relatedness, it is uncertain whether direct or inclusive fitness benefits have chiefly promoted cornicle evolution. Morphological measurements for apterous parthenogen, alate parthenogen, female sexual and male sexual morphs of 43 species (21 genera, one subfamily) were assessed to distinguish between the hypotheses that: (1) cornicles evolved for mechanical defence against natural enemies (direct fitness); (2) cornicles evolved for alarm signalling (inclusive fitness); or (3) cornicle length has been largely constrained by flight aerodynamics. Our results generally support the inclusive fitness hypothesis; cornicle length decreases as the relative number and relatedness of offspring decreases. As cornicle length is greatest in apterous parthenogenetic morphs, inclusive fitness benefits of protecting highly related kin may have been a key factor selecting for cornicles, and increased cornicle length, in aphids.