EXPERTS from Liverpool John Moores University have dated two lion skulls found at the Tower of London to the 13th and 15th centuries.

The discovery adds to evidence that the "king of the beasts" had more than a symbolic association with early English monarchs.

Lions have appeared on the coat of arms of English kings and queens since the reign of Henry II, who ruled from 1154 to 1189.

The animals are thought to have been housed in the Tower's Royal Menagerie, established by King John in the 13th century.

Despite their royal status, it seems that, like other animals, the lions were unceremon- iously dumped in the moat when they died.

They would have been kept in appalling conditions by today's standards, confined to cages measuring just six metres square radiating off from a central courtyard.

The later lion skull bears signs of an abnormality still seen today in captive lions.

Dr Hannah O'Regan, from Liverpool JMU, who led the study, said: "In addition to contributing to our knowledge of historical menageries, zoo archaeology also has a lot to offer in the study of modern zoo animals.

"Our research has highlighted a condition in captive lions that has been seen in specimens kept in different locations and different countries 500 years apart. Such studies of past zoo or captive populations have the potential to inform not only archaeology but also conservation strategies."

Although the site of the early Royal Menagerie is unknown, Edward I had a semi-circular structure later known as the "Lion Tower" built in the south-west corner of the castle in 1276.

By the 16th century, this structure definitely housed the Royal Menagerie, in which bears, leopards and other wild animals were also kept.

Jeremy Ashbee, Inspector of Ancient Monuments and Historic Buildings at English Heritage, and former curator at the Tower of London, said: "The menagerie seems to have been a private collection for the king, a sign that he enjoyed good relations with foreign monarchs, who presented him with animals.

"Lions were particularly prized as the living emblems of the royal arms of England, much like modern mascots."

The lion skulls were recovered in the 1930s from the moat adjacent to the Middle and Lion towers, suggesting that the animals were kept in this area.

The best-preserved lion skull was dated to between 1280 and 1385, making it Britain's earliest medieval big cat.