Isolation by time and habitat and coexistence of distinct host races of the common cuckoo.

Isolation by time occurs when different populations of a single species reproduce at different times and thereby reduce the probability of interbreeding, potentially causing divergent adaptation to timing of reproduction, eventually resulting in ecological species separated by timing of reproduction. We analysed extensive data on timing of reproduction by different host races of the common cuckoo Cuculus canorus that is an obligate brood parasite laying eggs in the nests of many different species of passerine birds. Because different hosts breed at different times, specific host races of cuckoos have adapted to specific hosts by laying eggs when nests of these hosts are available, and such divergence may be further exaggerated by differences in timing of breeding among host races with similar habitat requirements. Host species accounted for a quarter of the variance in timing of breeding by the cuckoo. Common cuckoos reproduced at a similar, but narrower subset of dates as did possible hosts, showing that only a fraction of hosts with specific breeding dates were parasitized. Common cuckoo eggs laid in the 'right' kind of nests, phenotypically matching the eggs of the host, were laid later during the season than cuckoo eggs laid in the 'wrong' kind of nests where the eggs did not mimic those of the host. Pairs of sympatric cuckoo host races differed more in timing of breeding than pairs of allopatric host races, and pairs of cuckoo host races with similar breeding habitat differed more in breeding date than pairs of cuckoo host races with dissimilar habitat, as expected from reproductive character displacement. These findings are consistent with cuckoo host races being isolated by timing of breeding and habitat.