About the Authors

Janine L. Brown

Contributed equally to this work with: Janine L. Brown, Stephen Paris, Natalia A. Prado-Oviedo

brownjan@si.edu

Affiliation Center for Species Survival, Smithsonian Conservation Biology Institute, Front Royal, Virginia, United States of America

Stephen Paris

Contributed equally to this work with: Janine L. Brown, Stephen Paris, Natalia A. Prado-Oviedo

Affiliation Center for Species Survival, Smithsonian Conservation Biology Institute, Front Royal, Virginia, United States of America

Natalia A. Prado-Oviedo

Contributed equally to this work with: Janine L. Brown, Stephen Paris, Natalia A. Prado-Oviedo

Affiliation Center for Species Survival, Smithsonian Conservation Biology Institute, Front Royal, Virginia, United States of America

Cheryl L. Meehan

‡ These authors also contributed equally to this work.

Affiliation AWARE Institute, Portland, Oregon, United States of America

Jennifer N. Hogan

‡ These authors also contributed equally to this work.

Affiliation AWARE Institute, Portland, Oregon, United States of America

Kari A. Morfeld

‡ These authors also contributed equally to this work.

Affiliations Center for Species Survival, Smithsonian Conservation Biology Institute, Front Royal, Virginia, United States of America, Lincoln Children’s Zoo, Lincoln, Nebraska, United States of America

Kathy Carlstead

‡ These authors also contributed equally to this work.

Affiliation Portland, OR, United States of America

Competing Interests

JLB, SP, NP-O and KM are employed by Smithsonian National Zoological Park (an AZA-accredited zoo). Author KM was employed by Lincoln Children’s Zoo (an AZA-accredited zoo) at the time of the study. KC was employed by Honolulu Zoo (an AZA-accredited zoo) at the time of the study. CM and JH are employed by AWARE Institute. AWARE Institute is a commercial entity founded by CM in 2014 and provides animal welfare assessment, research and education services to zoos and aquariums. These interests do not alter the authors' adherence to all the PLOS ONE policies on sharing data and materials.

Author Contributions

Conceived and designed the experiments: JLB KC. Performed the experiments: JLB SP NP-O KM. Analyzed the data: CM JNH KM. Contributed reagents/materials/analysis tools: JLB SP NP-O. Wrote the paper: JLB CM KC.