Several plant-herbivore hypotheses are based on the assumption that plants cannot simultaneously allocate resources to growth and defence. We studied seasonal patterns in allocation to growth and putatively defensive compounds by monitoring several chemical and physical traits in the leaves of mountain birch from early June (budburst) to late September (leaf senescence). We found significant seasonal changes in all measured characteristics, both in terms of concentrations (mg g) and amounts (mg leaf). Changes were very rapid in the spring, slow in the middle of the season, and there was another period of fast changes in the senescing leaves. Co-occurring changes in physical leaf traits and concentrations of several compounds indicated a seasonal decline in foliage suitability for herbivores. Concentrations of protein and free amino acids declined through the growing season whereas individual sugars showed variable seasonal patterns. The seasonal trends of phenolic groups differed drastically: concentrations of soluble proanthocyanidins increased through the season, whereas cell wall-bound proanthocyanidins, gallotannins and flavonoid glycosides declined after an initial increase in young leaves. We failed to find proof that the seasonal accumulation of phenolics would have been seriously compromised by leaf or shoot growth, as assumed by the growth/differentiation balance hypothesis and the protein competition model hypothesis. On the contrary, there was a steady increase in the total amount of phenolics per leaf even during the most active leaf growth.

Understanding the early evolution of aposematic (warning) coloration has been a challenge for scientists, as a new conspicuous morph in a population of cryptic insects would have a high predation risk and would probably die out before local predators learnt to avoid it. Fisher presented the idea of aggregation benefit through the survival of related individuals; however, his theory has been strongly debated as the mechanisms that favour grouping have never been explored experimentally with the incorporation of detectability costs. Here we create a comprehensive 'novel world' experiment with the great tit (Parus major) as a predator to explore simultaneously the predation-related benefits and costs for aposematic aggregated prey, manipulating both group size and signal strength. Our results show that grouping would have been highly beneficial for the first aposematic prey individuals surrounded by naive predators, because (1) detectability risk increased only asymptotically with group size; (2) additional detectability costs due to conspicuous signals were marginal in groups; (3) even naive predators deserted the group after detecting unpalatability (dilution effect); and (4) avoidance learning of signal was faster in groups. None of these mechanisms require kin selection.

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The quality of tree leaves as food for herbivores changes rapidly especially during the spring and early summer. However, whether the quality of an individual tree in relation to other trees in the population changes during the growing season and between years is less clear. We studied the seasonal and annual stability of chemical and physical traits affecting leaf quality for herbivores. Rankings of trees in terms of the contents of two major groups of phenolics in their leaves, hydrolyzable tannins and proanthocyanidins (condensed tannins), were very stable from the early spring to the end of the growing season. There were also strong positive within-season correlations in the levels of some other groups of phenolics in the leaves (kaempferol glycosides, myricetin glycosides and p-coumaroylquinic acid derivatives). The contents of individual sugars and the sum content of protein-bound amino acids showed patterns of seasonal consistency in mature leaves, but not in young developing leaves. The seasonal correlations in leaf water content and toughness were also strongest in mature leaves. The correlations between two years at corresponding times of the growing season were strongly positive for the major groups of phenolics throughout the season, but were more variable for the contents of proteins and some sugars. Leaf toughness and water content showed strong positive correlations in mature leaves. Despite the consistency of tree ranking in terms of leaf phenolics, the relative resistance status of trees may, however, change during a growing season because there was a negative correlation between the content of hydrolyzable tannins (early-season resistance compounds) in leaves early in the season and the content of proanthocyanidins (late-season resistance compounds) late in the season, and vice versa. Thus, assuming that phenolics affect herbivore preference and performance, different plants may suffer damage at different times of the growing season, and the overall variation between trees in the fitness consequences may be low. In addition, the adaptation of herbivorous insects to mountain birch foliage in general, as well as to specific tree individuals, may be constrained by variation in the relative resistance status of the trees.