Wild chimpanzees select tool material based on efficiency and knowledge.

Some animals have basic culture, but to date there is not much evidence that cultural traits evolve as part of a cumulative process as seen in humans. This may be due to limits in animal physical cognition, such as an inability to compare the efficiency of a novel behavioural innovation with an already existing tradition. We investigated this possibility with a study on a natural tool innovation in wild chimpanzees: moss-sponging, which recently emerged in some individuals to extract mineral-rich liquids at a natural clay-pit. The behaviour probably arose as a variant of leaf-sponging, a tool technique seen in all studied chimpanzee communities. We found that moss-sponges not only absorbed more liquid but were manufactured and used more rapidly than leaf-sponges, suggesting a functional improvement. To investigate whether chimpanzees understood the advantage of moss- over leaf-sponges, we experimentally offered small amounts of rainwater in an artificial cavity of a portable log, together with both sponge materials, moss and leaves. We found that established moss-spongers (having used both leaves and moss to make sponges) preferred moss to prepare a sponge to access the rainwater, whereas leaf-spongers (never observed using moss) preferred leaves. Survey data finally demonstrated that moss was common in forest areas near clay-pits but nearly absent in other forest areas, suggesting that natural moss-sponging was at least partly constrained by ecology. Together, these results suggest that chimpanzees perceive functional improvements in tool quality, a crucial prerequisite for cumulative culture.

Kin-based cultural transmission of tool use in wild chimpanzees.

Current research on animal culture has focused strongly on cataloging the diversity of socially transmitted behaviors and on the social learning mechanisms that sustain their spread. Comparably less is known about the persistence of cultural behavior following innovation in groups of wild animals. We present observational data and a field experiment designed to address this question in a wild chimpanzee community, capitalizing on a novel tool behavior, moss-sponging, which appeared naturally in the community in 2011. We found that, 3 years later, moss-sponging was still present in the individuals that acquired the behavior shortly after its emergence and that it had spread further, to other community members. Our field experiment suggests that this secondary radiation and consolidation of moss-sponging is the result of transmission through matrilines, in contrast to the previously documented association-based spread among the initial cohort. We conclude that the spread of cultural behavior in wild chimpanzees follows a sequential structure of initial proximity-based horizontal transmission followed by kin-based vertical transmission.

Mineral Acquisition from Clay by Budongo Forest Chimpanzees.

Chimpanzees of the Sonso community, Budongo Forest, Uganda were observed eating clay and drinking clay-water from waterholes. We show that clay, clay-rich water, and clay obtained with leaf sponges, provide a range of minerals in different concentrations. The presence of aluminium in the clay consumed indicates that it takes the form of kaolinite. We discuss the contribution of clay geophagy to the mineral intake of the Sonso chimpanzees and show that clay eaten using leaf sponges is particularly rich in minerals. We show that termite mound soil, also regularly consumed, is rich in minerals. We discuss the frequency of clay and termite soil geophagy in the context of the disappearance from Budongo Forest of a formerly rich source of minerals, the decaying pith of Raphia farinifera palms.