Evolutionary significance of antiparasite, antipredator and learning phenotypes of avian nest defence.

Avian nest defence, which is expected to serve both antiparasite and antipredator functions, may benefit or be detrimental to birds, although selective forces that potentially operate on nest defence have not been quantified as a whole. Together with fitness values, we analysed two traits of nest defence, intensity and plasticity, in two distantly related passerine species, yellow warbler (Setophaga petechia) in North America and reed warbler (Acrocephalus scirpaceus) in Europe, both favourite host species for brood parasites. Breeders that escaped parasitism were the most vocal among reed warblers, whereas there was no specific defence phenotype that predicted prevention of parasitism in yellow warblers. Breeders that escaped nest predation were, in both species, those with the most distractive response at the first exposure to a nest-threatening event, such as the experimental predation or parasitism simulated at the nest. However, increasing defence intensity benefited yellow warblers but was detrimental to reed warblers, because intense defence responses attracted predators. Adaptiveness of nest defence was revealed by nest defence phenotypes when examined in concert with the seasonal fitness (i.e. measures of reproductive success). Results revealed selective forces favoured yellow warblers with strong defence phenotypes. Opposite forces were instead revealed among reed warblers whose favoured phenotypes were strong, yet less flexible, defenders.

BEHAVIORAL DEFENSES AGAINST AVIAN BROOD PARASITISM IN SYMPATRIC AND ALLOPATRIC HOST POPULATIONS.

The brown-headed cowbird (Molothrus ater) is a widespread, obligate brood parasite of North American passerine birds. In southern Manitoba, where hosts are sympatric with cowbirds, American robins (Turdus migratorius) ejected parasitic eggs from all experimentally parasitized clutches (N = 25) and no eggs were accepted for more than four days. In contrast, robins in northern Manitoba, an area where cowbirds do not breed, accepted parasitic eggs in 33% of nests (N = 18) for at least five days. Acceptance of experimental cowbird eggs by a second host, the yellow warbler (Dendroica petechia), was similar in allopatric (100% of 20 nests) and sympatric (88.6% of 35 nests) populations, but models of a female cowbird elicited greater nest defense by warblers in the area of sympatry. Neither host rejected eggs of conspecifics, thus, rejection of cowbird eggs was not an epiphenomenon of conspecific brood parasitism. These results support the hypothesis that recognition of cowbirds and their eggs evolved as adaptations to counter cowbird parasitism and not some other selection pressure. The expression of anti-parasite defenses by some individuals within allopatric populations further suggests these traits may be controlled genetically but persist in such areas either through the continued introgression of rejecter genes from sympatric populations or because of the low cost of rejection behavior when parasitism is absent or rare.