Efficiency fosters cumulative culture across species.

Recent studies in several taxa have demonstrated that animal culture can evolve to become more efficient in various contexts ranging from tool use to route learning and migration. Under recent definitions, such increases in efficiency might satisfy the core criteria of cumulative cultural evolution (CCE). However, there is not yet a satisfying consensus on the precise definition of efficiency, CCE or the link between efficiency and more complex, extended forms of CCE considered uniquely human. To bring clarity to this wider discussion of CCE, we develop the concept of efficiency by (i) reviewing recent potential evidence for CCE in animals, and (ii) clarifying a useful definition of efficiency by synthesizing perspectives found within the literature, including animal studies and the wider iterated learning literature. Finally, (iii) we discuss what factors might impinge on the informational bottleneck of social transmission, and argue that this provides pressure for learnable behaviours across species. We conclude that framing CCE in terms of efficiency casts complexity in a new light, as learnable behaviours are a requirement for the evolution of complexity. Understanding how efficiency greases the ratchet of cumulative culture provides a better appreciation of how similar cultural evolution can be between taxonomically diverse species-a case for continuity across the animal kingdom. This article is part of a discussion meeting issue 'The emergence of collective knowledge and cumulative culture in animals, humans and machines'.

Complex foraging behaviours in wild birds emerge from social learning and recombination of components.

Recent well-documented cases of cultural evolution towards increasing efficiency in non-human animals have led some authors to propose that other animals are also capable of cumulative cultural evolution, where traits become more refined and/or complex over time. Yet few comparative examples exist of traits increasing in complexity, and experimental tests remain scarce. In a previous study, we introduced a foraging innovation into replicate subpopulations of great tits, the 'sliding-door puzzle'. Here, we track diffusion of a second 'dial puzzle', before introducing a two-step puzzle that combines both actions. We mapped social networks across two generations to ask if individuals could: (1) recombine socially-learned traits and (2) socially transmit a two-step trait. Our results show birds could recombine skills into more complex foraging behaviours, and naïve birds across both generations could learn the two-step trait. However, closer interrogation revealed that acquisition was not achieved entirely through social learning-rather, birds socially learned components before reconstructing full solutions asocially. As a consequence, singular cultural traditions failed to emerge, although subpopulations of birds shared preferences for a subset of behavioural variants. Our results show that while tits can socially learn complex foraging behaviours, these may need to be scaffolded by rewarding each component. This article is part of a discussion meeting issue 'The emergence of collective knowledge and cumulative culture in animals, humans and machines'.

Stable producer-scrounger dynamics in wild birds: sociability and learning speed covary with scrounging behaviour.

There has been extensive game-theoretic modelling of conditions leading to equilibria of producer-scrounger dichotomies in groups. However there is a surprising paucity of experimental evidence in wild populations. Here, we examine producer-scrounger games in five subpopulations of birds feeding at a socially learnt foraging task. Over four weeks, a bimodal distribution of producers and scroungers emerged in all areas, with pronounced and consistent individual tactic specialization persisting over 3 years. Tactics were unrelated to exploratory personality, but correlated with latency to contact and learn the foraging task, with the late arrivers and slower learners more likely to adopt the scrounging role. Additionally, the social environment was also important: at the broad scale, larger subpopulations with a higher social density contained proportionally more scroungers, while within subpopulations scroungers tended to be central in the social network and be observed in larger foraging flocks. This study thus provides a rare example of a stable, dimorphic distribution of producer-scrounger tactics in a wild population. It further gives support across multiple scales for a major prediction of social foraging theory; that the frequency of scroungers increases with group size.

Consistent individual differences in the social phenotypes of wild great tits, Parus major.

Despite growing interest in animal social networks, surprisingly little is known about whether individuals are consistent in their social network characteristics. Networks are rarely repeatedly sampled; yet an assumption of individual consistency in social behaviour is often made when drawing conclusions about the consequences of social processes and structure. A characterization of such social phenotypes is therefore vital to understanding the significance of social network structure for individual fitness outcomes, and for understanding the evolution and ecology of individual variation in social behaviour more broadly. Here, we measured foraging associations over three winters in a large PIT-tagged population of great tits, and used a range of social network metrics to quantify individual variation in social behaviour. We then examined repeatability in social behaviour over both short (week to week) and long (year to year) timescales, and investigated variation in repeatability across age and sex classes. Social behaviours were significantly repeatable across all timescales, with the highest repeatability observed in group size choice and unweighted degree, a measure of gregariousness. By conducting randomizations to control for the spatial and temporal distribution of individuals, we further show that differences in social phenotypes were not solely explained by within-population variation in local densities, but also reflected fine-scale variation in social decision making. Our results provide rare evidence of stable social phenotypes in a wild population of animals. Such stable social phenotypes can be targets of selection and may have important fitness consequences, both for individuals and for their social-foraging associates.

Individual personalities predict social behaviour in wild networks of great tits (Parus major).

Social environments have an important effect on a range of ecological processes, and form a crucial component of selection. However, little is known of the link between personality, social behaviour and population structure. We combine a well-understood personality trait with large-scale social networks in wild songbirds, and show that personality underpins multiple aspects of social organisation. First, we demonstrate a relationship between network centrality and personality with 'proactive' (fast-exploring) individuals associating weakly with greater numbers of conspecifics and moving between flocks. Second, temporal stability of associations relates to personality: 'reactive' (slow-exploring) birds form synergistically stable relationships. Finally, we show that personality influences social structure, with males non-randomly distributed across groups. These results provide strong evidence that songbirds follow alternative social strategies related to personality. This has implications not only for the causes of social network structure but also for the strength and direction of selection on personality in natural populations.

Social networks predict patch discovery in a wild population of songbirds.

Animals use social information in a wide variety of contexts. Its extensive use by individuals to locate food patches has been documented in a number of species, and various mechanisms of discovery have been identified. However, less is known about whether individuals differ in their access to, and use of, social information to find food. We measured the social network of a wild population of three sympatric tit species (family Paridae) and then recorded individual discovery of novel food patches. By using recently developed methods for network-based diffusion analysis, we show that order of arrival at new food patches was predicted by social associations. Models based only on group searching did not explain this relationship. Furthermore, network position was correlated with likelihood of patch discovery, with central individuals more likely to locate and use novel foraging patches than those with limited social connections. These results demonstrate the utility of social network analysis as a method to investigate social information use, and suggest that the greater probability of receiving social information about new foraging patches confers a benefit on more socially connected individuals.