Stonechat pairs in winter territories

Species like the Robin keep territories during the winter, with males and female individuals defending their own separate territories, while in the summer, territories are defended as a pair. There are some exceptions in which pairs defend winter territories, often in different areas that summer ones. One of these species is the Stonechat. In their wintering grounds on the coast and around the Humber estuary, it is frequent to encounter Stonechat pairs in the same area, feeding near each other and following each other, with no aggression (top shot, a pair at Alkborough, Lincolnshire, 11/12/2017). Of course, winter territory defence has nothing to do with reproduction, and often happens in areas that don't even hold breeding pairs in the summer. This unusual behaviour must offer some benefit to persist in the population, maybe allowing individuals to increase their chances of survival to predation or competition and to secure sufficient food resources.
A study of wintering Stonechats in the Negev desert (Israel), shed some light onto the behaviour. A total of 89 individuals were colour banded and followed through two wintering seasons. Their territorial behaviour was noted. The most surprising result of the study is that wintering Stonechat pairs are very fluid. Pairs often form after arrival to winter areas, and individuals rarely stay together all winter, instead pairs change frequently, with an average of just 4 weeks together per pair. In fact, none of the studied pairs stayed together all winter long and individuals arrived in the area and left at different times. This rules out that breeding pairs migrate together or settle on the same winter territory. Up to a third of individual are unpaired at any given time, but all associations are always between a male and a female. There were some aggressive interactions between members of a pair, which were commonly initiated by the male. Both males and females engaged in territory defence, with males more aggressive against intruders than females.

Male Stonechat at Kilnsea, 10/12/18.

Why pair in the winter?
If pairs are not stable, don't migrate together, or don't form in preparation for the winter season, what is the point in defending territories as a pair? Males and female stonechats feed on the same prey, therefore there may be a cost on sharing a territory during the winter, when food resources may be very scarce. A possible benefit outweighing the cost may be joint territory defence: two individuals are better spotting and chasing away conspecific intruders or detecting predators and defending the territory against individuals of other species with similar food resources. In this study, individuals spent a lot of time defending the territory against Mourning Wheatears, but there was no mention of differences when pairs or individual Stonechats were involved.

Another pair at Kilnsea (13/1/2020)

Why male/female pairs?
These results don't clarify why associations are not between two males or two females defending a territory. The authors speculate that as Stonechats are sexually dimorphic, even in the winter, there may be constraints that prevent two males from living peacefully in the same territory, and the increased male aggressiveness may mean two females may be unable to defend a territory against a male/female pair.

More information
Gwinner, E., Rödl, T. and Schwabl, H. Pair Territoriality of Wintering Stonechats: Behaviour, Function and Hormones. Behav. Ecol. Sociobiol. 34, 321–327 (1994).