BACKGROUND: Effective communication between sexual partners is essential for successful reproduction. Avian parents with biparental incubation need to know how to negotiate, when and who will incubate, and how to harmonize partner exchange at the nest. Although considerable effort has been dedicated to studies of incubation rhythms, few studies have investigated how behavioural signals serve to tighten cooperation between parents. Moreover, existing studies are almost exclusively restricted to species in which long distances between incubating and non-incubating parents prevent continuous communication during incubation. Thus, the most frequently described parental exchange system is a simple model characterized by the return of the non-incubating parent to the nest itself. Here, we propose more complex parental exchange behaviour in the Northern Lapwing (Vanellus vanellus), a territorial species capable of continuous partner communication during incubation and with a highly variable male contribution to incubation. RESULTS: Northern Lapwing females regularly vocalized shortly before departing from the nest, while males mostly left the nest quietly. Responsiveness of the male to female vocalization, perhaps in combination with her flying away from the nest, helped to synchronize incubation care by increasing the probability of exchange, and also by shortening the exchange gaps. In contrast, a male-to-female exchange gap most often occurred after the male quietly flew away from the nest. The frequency of female vocal signalling was not correlated with the male incubation effort on a between-nest scale, but the highest probability of a female-to-male exchange occurred after vocal signalling by females with the most nest-attentive males. Conversely, lowered effort by females to vocalize in the night was accompanied by lower willingness of males to incubate. CONCLUSIONS: Our results suggest that (1) that the incubating parent can communicate with the non-incubating partner using sex-specific behavioural signals, and this helps to synchronize parental exchange on the nest, (2) this signalling may combine acoustic and visual cues, and (3) the efficiency of this signalling might influence the overall nest attendance. The presumption that the repertoire of behavioural signals during reproduction will be much more complex in territorial species that are capable of continuous communication between the partners during the incubation period should be further tested.

Faculty of Environmental Sciences Czech University of Life Sciences Prague Kamýcká 129 165 00 Praha Suchdol Czech Republic

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