Adaptive phenological mismatches of birds and their food in a warming world

Climate change has profound ecological effects in birds, with the clearest effect a shift in timing, or phenology, of avian reproduction. To assess the consequences of these shifts, we performed a literature search and compared the rates of phenological change in the reproduction of birds with that of the food for their offspring. While in some areas the rate of change of the birds and their food was similar, there were also areas where the birds' shift lagged behind that of their food. In these cases, this will lead to a phenological mismatch, which will affect the fitness of the brood. There are two hypotheses explaining why climate change leads to mismatched reproduction: either the cues used no longer accurately predict the peak in food abundance (the cues hypothesis) or the fitness costs of egg production and/or incubation of laying early enough to match reproduction are substantial in early spring and are not compensated by the fitness benefits of a better matched reproduction (constraint hypothesis). In the latter case, the phenological mismatch is adaptive. We present a simple mathematical model to show that this may be the case if there are fitness costs of egg laying and/or incubation under cold conditions and if the temperatures that determine the peak in food abundance increase stronger than the temperatures affecting the costs of egg laying and incubation, as is the case in the Netherlands. Whether or not a phenological mismatch is adaptive has important consequences for natural selection acting on timing of reproduction. If the mismatch is not adaptive, timing of reproduction will be under direct natural selection, while, if the mismatch is adaptive, selection is likely to be on the costs of egg production, possibly on egg size or adult size. In all cases, a mismatch is expected to have negative population consequences and, especially when the mismatch is adaptive, these consequences cannot be reduced by a response to natural selection on timing directly. This makes experimental studies on laying date, which can determine whether the mismatch is adaptive, of crucial importance.