Animal cultures: how we've only seen the tip of the iceberg.

For humans we implicitly assume that the way we do things is the product of social learning and thus cultural. For animals, this conclusion requires proof. Here, we first review the most commonly used procedure for documenting animal culture: the method of exclusion, which charts geographic behavioral variation between populations as evidence for culture. Using published data, we show that, whereas it is an adequate proof of principle, the method of exclusion has major deficiencies when capturing cultural diversity and complexity. Therefore, we propose a new method, namely the direct counting of socially learned skills, which we apply to previously collected data on wild orangutans. This method reveals a far greater cultural repertoire among orangutans, and a different distribution of cultural elements among behavioral domains than found by the method of exclusion, as well as clear ecological correlates for most cultural elements. The widespread occurrence of social learning ability throughout the animal kingdom suggests that these conclusions also apply to many other species. Culture is most likely more widespread and pervasive than commonly thought and an important avenue to local adaptation. The complex and normative dimensions of culture seem unique to our species, but were most likely built upon a very broad, pre-existing cultural capacity that we inherited from our ancestors.