Colony size affects nestling immune function: a cross-fostering experiment in a colonial waterbird

Elevated transmission rate of pathogens and parasites is considered one of the major costs of sociality in birds. However, greater risk of infection in colonial birds might be compensated by specific immune adaptations. Here, we predicted that nestlings raised in larger colonies should invest more in their immune function. To test this hypothesis, we manipulated colony size and conduced cross-fostering experiment in a colonial waterbird, the common tern Sterna hirundo. Establishment of different size colonies under uniform environmental conditions was induced by providing large and small patches of attractive nesting area for terns (floating rafts). Then, pairs of clutches were swapped between large and small tern colonies, and skin-swelling response to phytohaemagglutinin (PHA) was assessed for nestlings from experimental and control broods. Contrary to our expectations, we found a negative effect of foster colony size on nestling PHA response (nestlings raised in the larger colony had lower PHA response). In addition, nestling PHA response correlated negatively with heterophil/lymphocyte ratio used as a measure of physiological stress. This suggested that low PHA response of nestlings raised in the larger colony could be mediated by an elevated level of social stress. We suggest that depression of immune function via social stress may constitute a strong selective pressure against large colony size in the common tern, and possibly in other colonial species. We also recommend that this largely overlooked cost of sociality should be considered in the further studies on the evolution and ecology of avian coloniality. ELECTRONIC SUPPLEMENTARY MATERIAL: The online version of this article (10.1007/s00442-019-04402-3) contains supplementary material, which is available to authorized users.