Animal Cognition

, Volume 22, Issue 2, pp 265–275 | Cite as

Tortoises develop and overcome position biases in a reversal learning task

  • Justin M. Bridgeman
  • Glenn J. TattersallEmail author
Original Paper


The capability of animals to alter their behaviour in response to novel or familiar stimuli, or behavioural flexibility, is strongly associated with their ability to learn in novel environments. Reptiles are capable of learning complex tasks and offer a unique opportunity to study the relationship between visual proficiency and behavioural flexibility. The focus of this study was to investigate the behavioural flexibility of red-footed tortoises and their ability to perform reversal learning. Reversal learning involves learning a particular discrimination task, after which the previously rewarded cue is reversed and then subjects perform the task with new reward contingencies. Red-footed tortoises were required to learn to recognise and approach visual cues within a Y-maze. Once subjects learned the visual discrimination, tortoises were required to successfully learn four reversals. Tortoises required significantly more trials to reach criterion (80% correct) in the first reversal, indicating the difficulty of unlearning the positive stimulus presented during training. Nevertheless, subsequent reversals required a similar number of sessions to the training stage, demonstrating that reversal learning improved up to a point. All subjects tested developed a position bias within the Y-maze that was absent prior to training, but most were able to exhibit reversal learning. Red-footed tortoises primarily adopted a win-stay choice strategy while learning the discrimination without much evidence for a lose-shift choice strategy, which may explain limits to their behavioural flexibility. However, improving performance across reversals while simultaneously overcoming a position bias provides insights into the cognitive abilities of tortoises.


Reptile Reversal learning Lateralisation Position bias Tortoise 



We would like to thank Tom Eles for the assistance with animal care, Dr. Cheryl McCormick for the scientific discussions during the course of this study, and Dr. Miriam Richards and Brock University’s Animal Behaviour class of 2014–2015 for assistance with pilot data collection on tortoise behaviour. This research was supported by a Natural Sciences and Engineering Research Council of Canada Discovery Grant to GJT (RGPIN-2014-05814). Data are made available at the Brock University repository (


The research program supporting this study was funded by an NSERC Discovery Grant to GJT (RGPIN-2014-05814).

Compliance with Ethical Standards

Conflict of interest

Justin Bridgeman and Glenn Tattersall declare that they have no conflict of interest.

Ethical approval

All applicable international, national, and/or institutional guidelines for the care and use of animals were followed.

Supplementary material

10071_2019_1243_MOESM1_ESM.pdf (216 kb)
Supplementary material 1 (PDF 216 KB)

Supplementary material 2 (M4V 8233 KB)

Supplementary material 3 (M4V 5709 KB)

Supplementary material 4 (M4V 5760 KB)

Supplementary material 5 (M4V 5561 KB)


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Copyright information

© Springer-Verlag GmbH Germany, part of Springer Nature 2019

Authors and Affiliations

  1. 1.Department of Biological SciencesBrock UniversitySt. CatharinesCanada

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