Expression of annual cycles in preen wax composition in red knots: constraints on the changing phenotype

Birds living in seasonal environments change physiology and behavior in correspondence to temporally changing environmental supplies, demands and opportunities. We recently reported that the chemical composition of uropygial gland secretions of sandpipers (Scolopacidae, order Charadriformes) changes during the breeding season from mixtures of monoesters to diesters, which fulfill specific functions related to incubation. A proper temporal match between the expression of diester preen waxes and incubation requires a flexible organization of the trait. Here we analyze the possible degrees of flexibility with reference to the functionality of better-understood molt and body mass cycles of free-living and captive red knots (Calidris canutus). The relative flexibility of seasonal cycles in preen wax composition was examined by two experimental perturbations: (1) giving birds restricted access to food and (2) monitoring them long-term under a constant photoperiodic regime. We found that wax type cannot change instantaneously, but that changing the type of wax is under similar organizational time constraints as the replacement of feathers. Just as molt and mass cycles, the seasonal rhythm of diester secretion appeared to be under endogenous control: most birds placed in a constant photoperiod still maintained seasonally changing preen waxes. Diester preen wax secretion was synchronized with the peak in body mass in spring, but became less well expressed under constant photoperiodic conditions and when food availability was limited.