Professor Frans de Waal, a primate behaviourist at Emory University, Atlanta, Georgia, said: “I don’t believe animals are moral in the sense we humans are – with well developed and reasoned sense of right and wrong – rather that human morality incorporates a set of psychological tendencies and capacities such as empathy, reciprocity, a desire for co-operation and harmony that are older than our species.

“Human morality was not formed from scratch, but grew out of our primate psychology. Primate psychology has ancient roots, and I agree that other animals show many of the same tendencies and have an intense sociality.”

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