[1] Univ St Andrews, Sch Psychol, Ctr Social Learning & Cognit Evolut, St Andrews KY16 9JP, Fife, Scotland
[2] Univ St Andrews, Sch Psychol, Evolut & Scottish Primate Res Grp, St Andrews KY16 9JP, Fife, Scotland
[3] Max Planck Inst Evolutionary Anthropol, Dept Primatol, D-04103 Leipzig, Germany
[4] Congo Program, Wildlife Conservat Soc, Mbeli Bai Study, Brazzaville, Rep Congo