Related Experiment Video
Updated: Mar 16, 2026

A Modified Mirror Test as a Visual Guide for the Self-awareness Trait in Wild Antarctica Penguins, Pygoscelis adeliae
Published on: July 8, 2025
Husbandry and enclosure influences on penguin behavior and conservation breeding
Andrew R Marshall1,2, Nicolas J Deere3,4,5, Holly A Little3,4,6
1Department of Environment, Centre for Integrated Research, Conservation and Learning (CIRCLE), University of York, Wentworth Way, York, United Kingdom. andy.marshall@york.ac.uk.
Abstract:
Multi-zoo comparisons of animal welfare are rare, and yet vital for ensuring continued improvement of zoo enclosures and husbandry. Methods are not standardized for the development of zoo enclosures based on multiple indicators, and case study species are required. This study compares behavior and breeding success to various enclosure and husbandry parameters for the Humboldt penguin, Spheniscus humboldti, for the development of improved enclosure design. Behavioral sampling was completed at Flamingo Land over a period of 8 months. Further data on behavior, enclosure design, and breeding success were collected via questionnaires, visits to zoos, and literature review. Breeding success was primarily influenced by colony age and number of breeding pairs, suggesting an important social influence on reproduction. Across zoos, there was also significant variation in behavior. The proportion of time spent in water varied between zoos (2-23%) and was used as an indicator of physical activity and natural behavior. Regression models revealed that water-use was best predicted by total enclosure area per penguin, followed by land area, with some evidence for positive influence of pool surface area per penguin. Predominantly linear/curvilinear increases in our biological indicators with enclosure parameters suggest that optimal conditions for S. humboldti were not met among the selected zoos. We propose revised minimum conditions for S. humboldti enclosure design, which exceed those in the existing husbandry guidelines. We present a framework for the evaluation of zoo enclosures and suggest that a rigorous scientific protocol be established for the design of new enclosures, based on multivariate methods. Zoo Biol. 35:385-397, 2016. © Wiley Periodicals, Inc.
Related Concept Videos
Conservation of Declining Populations
What is Behavior?
Optimal Foraging
Conservation of Small Populations
Mate Choice
Evolutionary Psychology

