Why do all host species not show defense against avian brood parasitism: evolutionary lag or equilibrium?

Avian brood parasitism reduces the reproductive success of hosts and is therefore expected to select for host defenses against parasitism, such as an ability to reject parasitic eggs. Field studies have shown that some hosts recognize and reject parasitism, whereas others do not, and the degree of the defense varies from population to population. One long-standing debate concentrates on the differences in the distribution of host defenses observed in hosts parasitized by the brown-headed cowbird and the common cuckoo. The cowbird's hosts show either few or nearly perfect defenses, whereas the cuckoo's hosts have defenses varying from none to complete, with most falling in between the two extremes. To explore the mechanisms underlying this pattern, I constructed a mathematical model in which host defense is assumed to be genetically determined and analyzed how the host defense is established under parasitic pressure. The model shows that differences in the defense-level distribution can be attributed to the difference in the parasite's breeding strategy, generalized or specialized: hosts parasitized by generalists show perfect, none, or intermediate levels of the defense depending on the host abundances, whereas hosts parasitized by specialists always exhibit either none or intermediate levels of the defense if the parasite lacks counter defenses such as egg mimicry. This result provides a testable explanation for the existence of accepter species of the brown-headed cowbird, which might reconcile the previously conflicting hypotheses.