The lions that terrified and delighted royalty, nobles and minions in the Tower of London were part of a subspecies that is now extinct.

Analysis of two skulls recovered from the moat at the Tower revealed that they belonged to Barbary lions. They dated from the 14th and 15th centuries and would have formed part of the collection that made up the Royal Menagerie.

Richard Sabin, curator of mammals at the Natural History Museum, said: “It’s an extinct subspecies. They are representatives of a form of lion from the Barbary Coast that no longer exists in the wild.”

The lions, which were caged for the pleasure of kings and queens of England from medieval times, were the first in Britain since the last Ice Age. Tower staff charged with caring for the big cats became so adept at getting them to breed that the animals became known as “English lions”.

Tigers, leopards, elephants and bears were also kept at the Tower, where they provided an exotic form of entertainment.

“Lions have been imported into Europe for various purposes since early historic times,” Mr Sabin said. “We’ve not known, however, until now the exact geographical origin of the animals found in London. Direct animal trade between Europe and sub-Saharan Africa was not developed until the 18th century so our results provide new insights into the patterns of historic animal-trafficking.”

Genetic tests were carried out on the skulls to reveal the species type, and researchers plan further tests to analyse isotopes in the bone to reveal where they were born. Details of isotopes that the lions absorbed from water and food as cubs should allow scientists to determine whether the pair were trapped in North Africa and transported to Britain or bred in Europe from African stock, perhaps at the Tower itself.

Mr Sabin said that the Royal Menagerie had been an important institution. “It was set up by King John. He had been given animals by other noblemen – it was a type of gift-giving that went on between important people of Europe in that period. Transporting the lions must have been quite a feat.”

The findings by the team from the Natural History Museum in London and the University of Oxford were backed up by comparing the skulls to the remains of Barbary and Asiatic lions. The findings were reported in the journal Contributions in Zoology.

The skulls were excavated in the 1930s and transferred to the museum, where they were put into storage. Radiocarbon tests dated one of the skulls between 1280 and 1385 and the other between 1420 and 1480, making them the earliest confirmed lion remains in the British Isles since the extinction of the Pleistocene cave lion.

The Royal Menagerie was closed in 1835 on the orders of the Duke of Wellington and the animals transferred to London Zoo. Barbary lions were driven to extinction by hunters in the early 20th century.

Nobuyuki Yamaguchi, of the University of Oxford, said: “According to historic records, a contiguous lion population could be found from North Africa and through the Middle East to India, until almost 4,000 years ago. Western North Africa was the nearest region to Europe to sustain lion populations until the early 20th century, making it an obvious and practical source for medieval merchants.”

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