Ontogenetic stage, plant vigor and sex mediate herbivory loads in a dioecious understory herb

Plant-herbivore interactions can be mediated by plant apparency, defensive and nutritional quality traits that change through plant ontogeny, resulting in age-specific herbivory. In dioecious species, opposing allocation patterns in defense may lead to sex-biased herbivory. Here, we examine how onto stage and plant sex determine levels of herbivore damage in understory herb Mercurialis perennis under field conditions. We analyzed variation in plant size (height, total leaf area), physical (specific leaf area) and chemical (total phenolic and condensed tannins contents) defense, and nutritional quality (total water, soluble protein and nonstructural carbohydrate contents) during the shift from reproductive to post-reproductive stage. Furthermore, we explored correlations between the analyzed traits and levels of foliar damage. Post-reproductive plants had lower levels of chemical defense, and larger leaf area removed, in spite of having lower nutritive quality. Opposing patterns of intersexual differences were detected in protein and phenolic contents during reproductive stage, while in post-reproductive stage total leaf area was sexually dimorphic. Female-biased herbivory was apparent only after reproduction. Plant size parameters combined with condensed tannins content determined levels of foliar damage during post-reproductive stage, while the only trait covarying with herbivory in reproductive stage was total nonstructural carbohydrate content. Our results support claims of optimal defense theory - sensitive stage of reproduction was better defended. We conclude that different combinations of plant traits mediated interactions with herbivores in mature stages. Differences in reproductive allocation between the sexes may not immediately translate into different levels of damage, stressing the need for considering different ontogenetic stages when exploring sex bias in herbivory.