The underestimated giants: operant conditioning, visual discrimination and long-term memory in giant tortoises

  • Tamar GutnickEmail author
  • Anton Weissenbacher
  • Michael J. KubaEmail author
Original Paper


Relatively little is known about cognition in turtles, and most studies have focused on aquatic animals. Almost nothing is known about the giant land tortoises. These are visual animals that travel large distances in the wild, interact with each other and with their environment, and live extremely long lives. Here, we show that Galapagos and Seychelle tortoises, housed in a zoo environment, readily underwent operant conditioning and we provide evidence that they learned faster when trained in the presence of a group rather than individually. The animals readily learned to distinguish colors in a two-choice discrimination task. However, since each animal was assigned its own individual colour for this task, the presence of the group had no obvious effect on the speed of learning. When tested 95 days after the initial training, all animals remembered the operant task. When tested in the discrimination task, most animals relearned the task up to three times faster than naïve animals. Remarkably, animals that were tested 9 years after the initial training still retained the operant conditioning. As animals remembered the operant task, but needed to relearn the discrimination task constitutes the first evidence for a differentiation between implicit and explicit memory in tortoises. Our study is a first step towards a wider appreciation of the cognitive abilities of these unique animals.


Operant learning Visual discrimination learning Long-term memory Aldabra tortoise Galapagos tortoise 



We want to thank the Vienna Zoo animal keeper team in the Aquarium house who supported and enriched this project with their help and ideas and the Elephant house and keepers for housing the tortoises and us for part of the experiment. The curators and the keepers at the Zürich zoo were essential for the completion of the work at their institution.

Compliance with ethical standards

Conflict of interest

T Gutnick has received an honorarium from Tiergarten Schönbrunn supported by donation from H.H. Lederer.

Ethical approval

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

Supplementary material

Supplementary Video SV1: Male Aldabra Tortoise training for Stage 2 - go to ball (MOV 18376 kb)

Supplementary Video SV2: Stage 3 - Color discrimination: Female Galapagos tortoise performing correct trial. Note the visual barrier in the background (MOV 21878 kb)


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Copyright information

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

Authors and Affiliations

  1. 1.Okinawa Institute of Science and TechnologyGraduate UniversityOkinawaJapan
  2. 2.Department of NeurobiologyInstitute of Life Sciences, The Hebrew UniversityJerusalemIsrael
  3. 3.Tiergarten SchönbrunnViennaAustria

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