About the Authors

Michael P. Muehlenbein

mpm1@indiana.edu

Affiliations Department of Anthropology, Indiana University, Bloomington, Indiana, United States of America, Sabah Wildlife Department, Kota Kinabalu, Sabah, Malaysia

Marc Ancrenaz

Affiliations Sabah Wildlife Department, Kota Kinabalu, Sabah, Malaysia, Kinabatangan Orangutan Conservation Programme, Sandakan, Sabah, Malaysia, North of England Zoological Society, Chester, United Kingdom

Rosman Sakong

Affiliation Red Ape Encounters, Sukau, Sabah, Malaysia

Laurentius Ambu

Affiliation Sabah Wildlife Department, Kota Kinabalu, Sabah, Malaysia

Sean Prall

Affiliation Department of Anthropology, Indiana University, Bloomington, Indiana, United States of America

Grace Fuller

Affiliations Cleveland Metroparks Zoo, Cleveland, Ohio, United States of America, Department of Biology, Case Western Reserve University, Cleveland, Ohio, United States of America

Mary Ann Raghanti

Affiliations Cleveland Metroparks Zoo, Cleveland, Ohio, United States of America, Department of Anthropology, School of Biomedical Sciences, Kent State University, Kent, Ohio, United States of America

Competing Interests

The authors have read the journal's policy and have the following conflicts: At the time of sample collection, Dr. Sakong was an employee of Red Ape Encounters (but is no longer). Dr. Ancrenaz was also a founding member of Red Ape Encounters. These previous affiliations have not influenced data analyses, interpretation of results, manuscript preparation or decision to publish. These previous affiliations do not alter the authors' adherence to all the PLoS ONE policies on sharing data and materials.

Author Contributions

Conceived and designed the experiments: MPM MA. Performed the experiments: MPM. Analyzed the data: MPM SP. Contributed reagents/materials/analysis tools: RS LA GF MAR. Wrote the paper: MPM. Collected and extracted the wild samples: RS. Collected and extracted the captive samples: GF MAR. Edited the manuscript: LA SP GF MAR.