Aggressive behavioural interactions between swans (Cygnusspp.) and other waterbirds during winter: a webcam-based study

Background Our understanding of any impacts of swans on other waterbirds (including other swans), and potential effects on waterbird community structure, remain limited by a paucity of fundamental behavioural and ecological data, including which species swans interact aggressively with and how frequently such interactions occur. Methods Behavioural observations of aggression by swans and other waterbirds in winters 2018/2019 and 2019/2020, were carried out via live-streaming webcams at two wintering sites in the UK. All occurrence sampling was used to identify all aggressive interactions between conspecific or heterospecifics individuals, whilst focal observations were used to record the total time spent by swans on aggressive interactions with other swans. Binomial tests were then used to assess whether the proportion of intraspecific aggressive interactions of each species differed from 0.5 (which would indicate equal numbers of intraspecific and interspecific interactions). Zero-inflated generalized linear mixed effects models (ZIGLMMs) were used to assess between-individual variation in the total time spent by swans on aggressive interactions with other swans. Results All three swan species were most frequently aggressive towards, and received most aggression from, their conspecifics. Our 10-min focal observations showed that Whooper (Cygnus cygnus) and Bewick's Swans (C. columbianus bewickii) spent 13.8 +/- 4.7 s (means +/- 95% CI) and 1.4 +/- 0.3 s, respectively, on aggression with other swans. These durations were equivalent to 2.3% and 0.2% of the Whooper and Bewick's Swan time-activity budgets, respectively. Model selection indicated that the time spent in aggressive interactions with other swans was best-explained by the number of other swans present for Whooper Swans, and an interactive effect of time of day and winter of observation for Bewick's Swans. However, the relationship between swan numbers and Whooper Swan aggression times was not strong (R-2 = 19.3%). Conclusions Whilst swans do exhibit some aggression towards smaller waterbirds, the majority of aggression by swans is directed towards other swans. Aggression focused on conspecifics likely reflects greater overlap in resource use, and hence higher potential for competition, between individuals of the same species. Our study provides an example of how questions relating to avian behaviour can be addressed using methods of remote data collection such as live-streaming webcams.