Avian population consequences of climate change are most severe for long-distance migrants in seasonal habitats.

One consequence of climate change is an increasing mismatch between timing of food requirements and food availability. Such a mismatch is primarily expected in avian long-distance migrants because of their complex annual cycle, and in habitats with a seasonal food peak. Here we show that insectivorous long-distance migrant species in The Netherlands declined strongly (1984-2004) in forests, a habitat characterized by a short spring food peak, but that they did not decline in less seasonal marshes. Also, within generalist long-distance migrant species, populations declined more strongly in forests than in marshes. Forest-inhabiting migrant species arriving latest in spring declined most sharply, probably because their mismatch with the peak in food supply is greatest. Residents and short-distance migrants had non-declining populations in both habitats, suggesting that habitat quality did not deteriorate. Habitat-related differences in trends were most probably caused by climate change because at a European scale, long-distance migrants in forests declined more severely in western Europe, where springs have become considerably warmer, when compared with northern Europe, where temperatures during spring arrival and breeding have increased less. Our results suggest that trophic mismatches may have become a major cause for population declines in long-distance migrants in highly seasonal habitats.

Climate change and unequal phenological changes across four trophic levels: constraints or adaptations?

1. Climate change has been shown to affect the phenology of many organisms, but interestingly these shifts are often unequal across trophic levels, causing a mismatch between the phenology of organisms and their food. 2. We consider two alternative hypotheses: consumers are constrained to adjust sufficiently to the lower trophic level, or prey species react more strongly than their predators to reduce predation. We discuss both hypotheses with our analyses of changes in phenology across four trophic levels: tree budburst, peak biomass of herbivorous caterpillars, breeding phenology of four insectivorous bird species and an avian predator. 3. In our long-term study, we show that between 1988 and 2005, budburst advanced (not significantly) with 0.17 d yr(-1), while between 1985 and 2005 both caterpillars (0.75 d year(-1)) and the hatching date of the passerine species (range for four species: 0.36-0.50 d year(-1)) have advanced, whereas raptor hatching dates showed no trend. 4. The caterpillar peak date was closely correlated with budburst date, as were the passerine hatching dates with the peak caterpillar biomass date. In all these cases, however, the slopes were significantly less than unity, showing that the response of the consumers is weaker than that of their food. This was also true for the avian predator, for which hatching dates were not correlated with the peak availability of fledgling passerines. As a result, the match between food demand and availability deteriorated over time for both the passerines and the avian predators. 5. These results could equally well be explained by consumers' insufficient responses as a consequence of constraints in adapting to climate change, or by them trying to escape predation from a higher trophic level, or both. Selection on phenology could thus be both from matches of phenology with higher and lower levels, and quantifying these can shed new light on why some organisms do adjust their phenology to climate change, while others do not.

Food-limitation in a generalist predator.

Investigating food-limitation in generalist predators is difficult, because they can switch to alternative prey, when one of their staple prey becomes scarce. Apart from data on the dynamics of the predator population, a robust study requires: (i) a documentation of the predator's entire prey base; and (ii) an experimental or natural situation, where profitable dietary shifts are impossible, because several preferred prey species decline simultaneously. Here, we provide a detailed description of how food-supply has limited a generalist avian top predator, the northern goshawk Accipiter gentilis. In our study area, populations of several principal goshawk prey species crashed simultaneously during 1975-2000, whereas other extrinsic factors remained essentially unchanged. The breeding and non-breeding segments of the local goshawk population declined markedly, associated with a significant increase in nest failures. Brood size of successful pairs remained unaffected by changes in prey availability. Breeding recruitment ceased at a time when potential replacement birds ('floaters') were still present, providing a rare empirical demonstration of an 'acceptance threshold' in raptor territory choice. To investigate how goshawk diet changed in response to varying food-supplies, we make novel use of an analytical tool from biodiversity research-'abundance-biomass-comparison curves' (ABC curves). With increasing levels of food-stress, the dominance of principal prey species in the diet decreased, and the number of small-bodied prey species increased, as did intra-guild predation. Our finding that breeder and non-breeder segments declined in concert is unexpected. Our results carry the management implication that, in food-limited raptor populations, externally induced breeder mortality can rapidly depress population size, as losses are no longer buffered when floaters reject breeding opportunities.