No experimental evidence for local competition in the nestling phase as a driving force for density-dependent avian clutch size

In birds, local competition for food between pairs during the nestling phase may affect nestling growth and survival. A decrease in clutch size with an increase in breeding density could be an adaptive response to this competition. To investigate whether breeding density causally affected the clutch size of great tits (Parus major), we manipulated breeding density in three out of eight study plots by increasing nest-box densities. We expected clutch size in these plots to be reduced compared to that in control plots. We analysed both the effects of variation in annual mean density (between-year comparisons) and experimental density (within-year comparison between plots) on clutch size variation, the occurrence of second broods and nestling growth. We examined within-female variation in clutch size to determine whether individual responses explain the variation over years. Over the 11 years, population breeding density increased (from 0.33 to 0.50 pairs ha(-1)) while clutch size and the occurrence of second broods decreased (respectively from 10.0 to 8.5 eggs and from 0.39 to 0.05), consistent with a negative density-dependent effect for the whole population. Nestling growth showed a declining but nonsignificant trend over years. The decline in population clutch size over years was primarily explained by changes occurring within individuals rather than selective disappearance of individuals laying large clutches. Within years, breeding density differed significantly between manipulated plots (0.16 pairs ha(-1) vs. 0.77 pairs ha(-1)) but clutch size, occurrence of second broods and nestling growth were not affected by the experimental treatment, resulting in a discrepancy between the effects of experimental and annual variation in density on reproduction. We discuss two hypotheses that could explain this discrepancy: (i) the decline in breeding performance over time was not due to density, but resulted from other, unknown factors. (ii) Density did cause the decline in breeding performance, but this was not due to local competition in the nestling phase. Instead, we suggest that competition acting in a different phase (e.g. before egg laying or after fledgling) was responsible for the density effect on clutch size among years.