Colony size as a predictor of breeding behaviour in a common waterbird.

The choice of colony size may have profound consequences for individual fitness in colonially breeding birds, but at the same time it may require certain behavioural adaptations. Here, we aimed to examine behavioural divergence of common terns Sterna hirundo nesting in colonies of different size. For this purpose, we promoted establishment of small (<35 pairs) and large (>100 pairs) tern colonies under uniform ecological and environmental conditions by providing attractive patches of nesting substrate (floating rafts) at a single site. We combined video recording and GPS-tracking to assess communal and individual defence initiation rate, intra-specific aggression rate, and foraging flight characteristics. We found that birds from larger colonies more frequently engaged in communal defence and they performed longer foraging flights, while terns from smaller colonies more frequently showed individual defence behaviours. Also, intra-specific aggression rate was higher in smaller colonies, but this effect was primarily attributed to a higher proportion of edge breeding pairs, which were more aggressive. Our results suggest that various colony sizes may be associated with different behavioural syndromes, which comprise of diverse personality traits, such as social responsiveness, social tolerance, or propensity for aggression. It remains to be tested whether these behavioural differences reflect processes of phenotypic sorting among colonies of different size or whether they are a result of behavioural plasticity under different social contexts.

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.