Causes and consequences of inverse density-dependent territorial behaviour and aggression in a monogamous mammal.

Territoriality is an important process shaping population dynamics, and the defence of a territory is crucial for individuals to increase the duration of territory occupancy and, consequently, reproductive success. However, little is known about how the frequency of territory intrusions and subsequent territorial behaviours and aggression by territory owners are affected by external factors, such as population density. This is important because it can affect mate change (the replacement of one pair member) and dispersal, a key ecological process. The aim of this study was to investigate the behavioural and spatial response of territory owners to intruder pressure as a function of population density in a territorial, monogamous mammal, the Eurasian beaver (Castor fiber). Using a combination of GPS technology, scent experiments, camera trap data and tail scar observations from an individual-based long-term study, we investigated the factors influencing spatial movement patterns by territory owners in response to a simulated intruder and the factors affecting territory intrusions. We found consistent inverse density-dependent patterns in territorial behaviours and evidence of conspecific aggression. At lower densities, territory owners detected more simulated intrusions, showed more territorial reactions and experienced increased conspecific aggression as indicated by tail scars, suggesting increased intruder pressure. Inverse density-dependent territorial behaviour and aggression suggest a potential mechanistic link between inverse density-dependent natal dispersal and mate change. At low population densities, increased dispersal amplifies intruder pressure, leading to the observed increases in territorial behaviours, conspecific aggression and previously observed mate turnover, which in turn might increase natal dispersal. Our study demonstrates how population density can affect the behaviour and space use of individuals, which is important for territory occupancy and fitness.