In human societies, cultural norms arise when behaviours are transmitted through social networks via high-fidelity social learning1. However, a paucity of experimental studies has meant that there is no comparable understanding of the process by which socially transmitted behaviours might spread and persist in animal populations2,3. Here we show experimental evidence of the establishment of foraging traditions in a wild bird population. We introduced alternative novel foraging techniques into replicated wild sub-populations of great tits (Parus major) and used automated tracking to map the diffusion, establishment and long-term persistence of the seeded innovations. Furthermore, we used social network analysis to examine the social factors that influenced diffusion dynamics. From only two trained birds in each sub-population, the information spread rapidly through social network ties, to reach an average of 75% of individuals, with a total of 414 knowledgeable individuals performing 57,909 solutions over all replicates. The sub-populations were heavily biased towards using the technique that was originally introduced, resulting in established local traditions that were stable over two generations, despite a high population turnover. Finally, we demonstrate a strong effect of social conformity, with individuals disproportionately adopting the most frequent local variant when first acquiring an innovation, and continuing to favour social information over personal information. Cultural conformity is thought to be a key factor in the evolution of complex culture in humans4,5,6,7. In providing the first experimental demonstration of conformity in a wild non-primate, and of cultural norms in foraging techniques in any wild animal, our results suggest a much broader taxonomic occurrence of such an apparently complex cultural behaviour.
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This project was supported by grants from the BBSRC (BB/L006081/1) and the ERC (AdG 250164) to B.C.S., who was also supported by a visiting professorship at Uppsala University. L.M.A. was also supported by an Australian Postgraduate Award; and A.T., by a BBSRC David Phillips Fellowship (BB/H021817/1). The EGI social networks group, S. Lang and K. McMahon provided assistance in the field, and M. Whitaker produced electronic components for the puzzle boxes.
The authors declare no competing financial interests.
Extended data figures and tables
Extended Data Figure 1 Wytham Woods, UK, (51° 46′ N, 01° 20′ W), showing the location of replicates and puzzle boxes.
The total area of Wytham Woods is 385 ha; the location and size of the separate woodland areas within the woods are labelled on the map. The green points indicate the puzzle-box locations for the three control replicates, C1, C2 and C3: Broad Oak, Bean Wood and Singing Way, respectively. The blue points indicate the puzzle-box locations for the two option A replicates, T1 and T2: Common Piece and Brogden’s Belt, respectively. The red points indicate the puzzle-box locations for the three option B replicates, T3, T4 and T5: Great Wood, Marley Plantation and Pasticks, respectively. (d) indicates the locations where trained demonstrators were caught from and released to.
a, Schematic of a feeding station (shut), with sunflower-seed feeder, RFID antennae and data-logging hardware. The cage is to restrict access to small passerines only. b, Map of the study area showing the placement of 65 feeding stations. The stations are approximately 250 m apart and open simultaneously from dawn to dusk on Saturday and Sunday over winter. c, Grouping events were inferred from the temporal data stream gained from the feeding stations, with individuals assigned to grouping events in a bipartite network. d, Repeated co-occurrences were used to create social networks22.
The red nodes represent individuals that acquired the novel behaviour after 20 days of exposure. The black nodes represent naive individuals. The yellow nodes represent trained demonstrators. The networks are heavily thresholded to show only the links above the average edge strength for each replicate (T1–T5, 0.09, 0.05, 0.08, 0.07 and 0.07, respectively). a, Network for the T1 replicate (n = 123). b, Network for the T2 replicate (n = 137). c, Network for the T3 replicate (n = 154). d, Network for the T4 replicate (n = 95). e, Network for the T5 replicate (n = 110).
Only individuals that performed both options are included, and individuals that moved between replicates are excluded. The lines are running proportions of the seeded option for each individual over its last ten visits. a, T1 (option A), n = 30. b, T2 (option A), n = 10. c, T3 (option B), n = 19. d, T4 (option B), n = 15. e, T5 (option B), n = 4.
The birds were presented with a freely available mix of 40 mealworms, 40 peanut granules and 40 sunflower seeds for 1 h on 2 days over 1 week at 6 sites (3 sites for T2 and 3 sites for T4). The trials were conducted 2 weeks after the end of the main experiment, in March 2014. Food choice was identified from video camera footage, and the trial was halted when all of one prey item was taken. Only great tits were included, but the birds could not be individually identified. The birds clearly preferred the live mealworms to peanut granules or sunflower seeds.
This file contains Supplementary Text and Supplementary Table 1. (PDF 164 kb)
First shows great tit using option A, pushing the door to the left from the blue side. The bird extracts a reward and the puzzle-box automatically resets 1sec after its departure. Second shows great tit using option B, pushing the door to the right from the red side. Again the bird gains access to the feeder, and the puzzle-box resets back to the middle 1 sec after its departure. (MP4 27734 kb)
Footage shows great tits interacting at the puzzle-box over a 1 min period at a busy site, with multiple birds either solving the puzzle-box, scrounging from others, or visually observing the solves of others. Up to two scrounges in the 1 sec after a solve are permitted, and are registered as such at the puzzle-box before it shuts. (MP4 14203 kb)
Foraging social network for T3 replicate as an example. Nodes represent individuals; lines are edges indicating the strength of connection between individuals. Yellow nodes are trained demonstrators, and nodes turn red in the order in which they first performed the new behavior (whether option A/B). (MP4 8344 kb)
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Aplin, L., Farine, D., Morand-Ferron, J. et al. Experimentally induced innovations lead to persistent culture via conformity in wild birds. Nature 518, 538–541 (2015). https://doi.org/10.1038/nature13998
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