Table 1. Population trends in relation to DD0Yr (regression coefficient of DD0 at MAD on year). The effects of DD0Yr on population trends are obtained from a model including a binary factor migration strategy but excluding the effect of the non-significant interaction. Full models are reported in electronic supplementary material, tables S1 and S2. Finland DD0Yr data were averaged between Hanko and Jurmo. Population trends for the seven countries and for Finland (a) are derived from BirdLife International [34]. Population trends used in model Finland (b) are derived from Va¨isa¨nen [35]. The number of short-distance or longdistance migrants for which population trend data were available was 70 and 42 for the seven countries, 65 and 39 for Finland (a), and 35 and 27 for Finland (b).to common ancestry, led to consistent results (table 1; see electronic supplementary material). Moreover, analyses run on DD23Yr or DDþ3Yr both on ‘raw’ species data and while controlling for phylogenetic effects largely confirmed the results of analyses based on DD0Yr (see electronic supplementary material, tables S1 and S2). Several previous studies of migratory birds have provided evidence that migrants have advanced their arrival to the breeding areas during recent decades [19,20]. Conversely, very few studies have investigated whether the rate of change in migration and breeding phenology has compensated for generalized phenological shifts owing to climate change [10,11,16]. Here, we have adopted an approach based on degree-days accumulation to investigate whether the progress of spring at the time when migratory birds arrive to their breeding areas has changed over the last five decades. This approach is novel and can be extended to studies of the ecological consequences of phenological response to climate change in studies of birds and other taxa, owing to the ubiquitous nature of the effects of temperature on the phenology of plants and ectothermic organisms (see §1). Advancement in arrival date to the breeding areas, which has indeed occurred for most species, has not fully compensated for climate change, and larger degree-day values at arrival in recent years may imply that birds have become ecologically mismatched. This is the case because the phenophases of plants and ectothermic animals, including those relevant to bird
ecology, are mostly regulated by temperature [24–29,52]. The present study thus rests on the assumption that long-term increase in degree-days at arrival causes an (increasing) ecological mismatch. This would not be the case if birds were arriving too early (i.e. at too low degree-days) in the past, as an increase in degree-days at arrival would ensure smaller, rather than larger, ecological mismatch. Although no unequivocal reference for optimal conditions (in terms of degree-days at arrival) is known for any bird species, we consider the possibility that ecological matching increases with increasing degree-days at arrival a remote one. First, selection for earlier arrival and breeding has been repeatedly
documented in bird studies [17], implying that birds arrive on average too late, rather than too early. Second, birds are advancing their arrival date, whereas they should be expected to delay it if they were arriving too early. Third, temperature changes have been occurring at faster pace during recent decades than in the previous ones [4] (see also figure 2). If birds show a constant latency in responding to climate change, more rapid recent climate change per se justifies the expectation of larger ecological mismatch.We thus consider the assumption that higher degree-days at arrival are associated with stronger ecological mismatch as warranted. Obviously, the extent of ecological mismatch arising because of a unit increase in degree-days may widely vary among species according to their trophic level and habitat. However, we found no statistical evidence that the effect of DDiYr on population trend depended on diet or habitat, suggesting that any negative consequence of change in degree-days at arrival on population trends was independent of variation in major ecological traits of the species. The ability of birds to cope with climate change by adjusting their migration phenology may be constrained by the timing of life-cycle phases preceding migration, which may limit the scope for phenotypically plastic response, by different climatic variation in the wintering areas and en route, and/or by depleted genetic variance in migration traits, which can hinder microevolutionary response [14]. All these scenarios are compatible with the observation that LDMs, having advanced their arrival less, have accumulated a larger ‘thermal delay’ when
compared with SDMs. We have also shown that species with larger ‘thermal delay’ at arrival have undergone the largest population decline, while species with small-to-moderate thermal delay have barely shown a decline. Previous studies have shown that LDMs have