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1. Home ranging is a near-ubiquitous phenomenon in the animal kingdom. Understanding the behavioural mechanisms that give rise to observed home range patterns is thus an important general question, and mechanistic home range analysis (MHRA) provides the tools to address it. However, such analysis has hitherto been restricted to scent-marking territorial animals, so its potential breadth of application has not been tested.\ud \ud 2. Here, we apply MHRA to a population of long-tailed tits (Aegithalos caudatus), a non15 territorial passerine, in the non-breeding season where there is no clear “central place” near which birds need to remain. The aim is to uncover the principal movement mechanisms underlying observed home range formation.\ud \ud 3. Our foundational models consist of memory-mediated conspecific avoidance between flocks, combined with attraction to woodland. These are then modified to incorporate the effects of flock size and relatedness, to uncover the effect of these on the mechanisms of home range formation.\ud \ud 4. We found that a simple model of spatial avoidance, together with attraction to the central parts of woodland areas, accurately captures long-tailed tit home range patterns. Refining these models further, we show that the magnitude of spatial avoidance by a flock is negatively correlated to both the relative size of the flock (compared to its neighbour) and the relatedness of the flock with its neighbour.\ud \ud 5. Our study applies MHRA beyond the confines of scent-marking, territorial animals, so paves the way for much broader taxonomic application. These could potentially help uncover general properties underlying the emergence of animal space use patterns. This is also the first study to apply MHRA to questions of relatedness and flock size, thus broadening the potential possible applications of this suite of analytic techniques.

Kin selection is regarded as a key process in the evolution of avian cooperative breeding, and kinship influences helper decisions in many species. However, the effect of kinship on nonbreeding social organization is still poorly understood despite its potential fitness implications. Here, we investigated the origins and consequences of kin associations in nonbreeding flocks of long-tailed tits, Aegithalos caudatus, an atypical cooperative breeder where helpers are failed breeders that redirect care towards relatives living in kin neighbourhoods. We found that kinship is an important factor in initial grouping decisions; all members of a nuclear family initially joined the same flock and failed breeders chose to flock with their relatives. Flocks that merged during the nonbreeding season also contained relatives. In contrast to these findings of positive kin association, when long-tailed tits switched flocks they tended to disperse into flocks with fewer relatives, although such switches often occurred with kin. In a playback experiment, we found no evidence that aggression shown towards members of other flocks was affected by kinship, indicating that kin associations result from a preference to flock with relatives rather than a constraint on flocking with nonrelatives. Finally, using social network analysis, we show that fine-scale nonbreeding associations among individuals were positively related to kinship, and that these nonbreeding associations were reflected in helping decisions in the subsequent breeding season, in addition to the previously reported effects of kinship and proximity. We conclude that long-tailed tits prefer to associate with kin when not breeding, and suggest that by doing so they gain either nepotistic benefits within flocks or future indirect benefits during breeding.

Kin selection has played an important role in the evolution and maintenance of cooperative breeding behaviour in many bird species. However, although relatedness has been shown to affect the investment decisions of helpers in such systems, less is known about the role that kin discrimination plays in other contexts, such as communal roosting. Individuals that roost communally benefit from reduced overnight heat loss, but the exact benefit derived depends on an individual's position in the roost which in turn is likely to be influenced by its position in its flock's dominance hierarchy. We studied the effects of kinship and other factors (sex, age, body size and flock sex ratio) on an individual's roosting position and dominance status in captive flocks of cooperatively breeding long-tailed tits Aegithalos caudatus. We found that overall, kinship had little influence on either variable tested; kinship had no effect on a bird's position in its flock's dominance hierarchy and the effect of kinship on roosting position was dependent on the bird's size. Males were generally dominant over females and birds were more likely to occupy preferred roosting positions if they were male, old and of high status. In this context, the effect of kinship on social interactions appears to be less important than the effects of other factors, possibly due to the complex kin structure of winter flocks compared to breeding groups.