About the Authors

Lucy A. Bates

Affiliation School of Psychology, University of St Andrews, St Andrews, United Kingdom

Rosie Handford

Affiliation School of Psychology, University of St Andrews, St Andrews, United Kingdom

Phyllis C. Lee

Affiliations Department of Psychology, University of Stirling, Stirling, United Kingdom, Amboseli Trust for Elephants, Nairobi, Kenya

Norah Njiraini

Affiliation Amboseli Trust for Elephants, Nairobi, Kenya

Joyce H. Poole

Affiliation Amboseli Trust for Elephants, Nairobi, Kenya

Katito Sayialel

Affiliation Amboseli Trust for Elephants, Nairobi, Kenya

Soila Sayialel

Affiliation Amboseli Trust for Elephants, Nairobi, Kenya

Cynthia J. Moss

Affiliation Amboseli Trust for Elephants, Nairobi, Kenya

Richard W. Byrne

rwb@st-andrews.ac.uk

Affiliation School of Psychology, University of St Andrews, St Andrews, United Kingdom

Competing Interests

The authors have declared that no competing interests exist.

Author Contributions

Conceived and designed the experiments: LAB RWB. Performed the experiments: PCL NN JP KS SS CJM. Analyzed the data: LAB RH. Wrote the paper: LAB RWB.