The songs of male pied flycatchers: exploring the legacy of the fathers.

Singing is a key element of songbirds' behavioral repertoire, particularly for males, which sing during the breeding season to defend resources against other males and to attract females. Different song traits may convey honest information about males' qualities or conditions, which may be used by females to select their mates. Traits under strong sexual selection have an important component of additive genetic variation (i.e., the main genetic inheritance from parents), and so relatively high heritability; therefore, it can be expected that song traits also do. Although the act of singing is an innate behavior, and thus, genetically determined, songbirds need to learn their songs and therefore the genetic contribution to song traits may be reduced by the effect of environmental factors. We tested this hypothesis in seven song traits recorded in the long-distance migratory bird, the pied flycatcher (Ficedula hypoleuca). From a 23-year database (1992-2015), we obtained songs for 28 father-son pairs, and for each song trait we applied parent-offspring regressions to estimate heritability. The type of syllables sung are learned from tutors, and here we also determined the cultural contribution of fathers to the song repertoires of their sons, by quantifying the percentage of syllables that sons shared with their fathers, and compared this with what sons shared with other males in the population (e.g., neighbors). The heritabilities of song traits were highly variable (ranging from -0.22 to 0.56), but most of these were around zero and none of them were significant. These results indicate that the seven song traits are most likely determined by environmental factors. Sons shared more syllables with their fathers than with neighbors (21% vs. 3%), suggesting that fathers are important song tutors during the nestling period. We conclude that there is a cultural inheritance from fathers to their sons' syllable repertoires, but there is no strong evidence for a genetic contribution of fathers to the seven song traits studied.