28Several species of non-human apes have been suggested to rely on copying to acquire some of 29 their behavioural forms. One of the most cited examples -and UN-protected -is nut-cracking 30 in chimpanzees. However, copying might not be the most parsimonious explanation for nut-31 cracking, considering the lack of evidence for spontaneous copying in this species. The zone 32 of latent solutions (ZLS) hypothesis argues instead that the behavioural form of nut-cracking 33 is individually learnt, whilst non-copying social learning fosters frequency differences across 34 populations. In order to differentiate between the copying and the ZLS hypothesis, four nut-35 cracking-naïve orangutans (Mage=16; age range=10-19; 4F; at time of testing) were provided 36 with nuts and hammers but were not demonstrated the behaviour. Whilst the adults in the 37 group were able to open nuts with their teeth, one juvenile spontaneously expressed nut-38 cracking with a wooden hammer. We therefore show that the behavioural form of nut-39 cracking does not necessarily rely on copying in orangutans. 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56Overall, then, we may surmise that wild apes, alongside untrained/unenculturated captive 89 apes, most likely lack the ability to copy novel actions.
90Despite these data, action copying is still often cited as the main mechanism behind ape, and 91 especially chimpanzee, behavioural forms. Some have even further claimed that (certain) ape 92 behaviours (such as tool-use behaviours) depend on copying social learning to be acquired by 93