Nov. 3, 2005 — Lions had more than a symbolic association with early English monarchs, according to a study published in the current issue of the International Journal of Osteoarchaeology.

Appearing on the English monarchy's coat of arms since the reign of Henry II (1133-1189), lions were one of the main attractions at the Tower of London, the massive stone fortress on the Thames that has witnessed the drama of English history for over eight centuries.

"The lions were potent symbols of the monarchy at the time of the Hundred Years War and the Wars of the Roses, and may have been among the first seen in Britain since they became extinct some 10,000 years before," Hannah O'Regan at Liverpool John Moores University's Research Centre in Evolutionary Anthropology and Palaeoecology, told Animal Planet News.

O'Regan analysed and radiocarbon-dated animal remains excavated at the Tower in 1937. The bones had never been fully examined, and remained stored at the Natural History Museum for the last 70 years.

The bones comprised two lion skulls, fragments of a leopard skull and the skulls of 19 dogs.

The best-preserved lion skull was radiocarbon dated to between 1280 and 1385, a period covering the reigns of Edward I, II and III. The dating makes the lion the earliest Medieval big cat known in Britain, said O'Regan.

While the badly damaged leopard skull was dated to between 1440 and 1625, which covers the Plantagenet reign, the Tudors and Stuarts, the second lion skull was dated to 1420-1480.

During those 60 years, the Chronicles of London report that all the lions in the Tower died.

It is thought the lions were housed in the Tower's "Royal Menagerie", or zoo. The animal collection was established by King John, who reigned in England from 1199-1216, and was closed in 1835, on the orders of the Duke of Wellington.

Lions, bears, elephants and other exotic species — all gifts presented by England's allies — were among the most popular attractions.

The last sight for beheaded queens, slaughtered princes and tortured prisoners, the Tower of London was a place of cruelty also for animals.

Lions were confined to cages measuring just six metres square (65 square feet), appalling conditions for animals that could reach 2.5 metres (eight feet) in length.

"Although exotic and difficult to procure, the cats were baited at times. James I in particular enjoyed baiting, and had the viewing gallery rebuilt so that there was more room to watch the matches," O'Regan said.

The presence of so many dogs at the Tower suggest that they were also used for animal baiting purposes.

Indeed, two puncture wounds found on a dog skull indicates dog fighting, according to O'Regan.

Despite their royal status, the lions were unceremoniously dumped, unskinned, in the Tower's moat when they died.

The researcher also found an abnormality in the later lion skull still seen today in captive lions — a bump of bone at the large hole at the base of the skull which allows passage of the spinal cord.

"It is a swelling of bone in the foramen magnum and it has been also noted in captive cat skulls from the early 20th century, indicating that it is a condition with a long history," O'Regan said.

"We don't know what causes it, or if it is found in wild lions as well as captive ones," O'Regan said.

Jeremy Ashbee, Inspector of Ancient Monuments and Historic Buildings at English Heritage and former curator at the Tower of London, described the research of "great significance for the history of the Tower, of the monarchy, of zoological collections and natural history in general."

According to Richard Sabin, Curator of Mammals at the Natural History Museum, the study will pave the way to new areas of research, providing "more information about the origins and diets of these historically very important animals."

"It's the physical remains, particularly the bones, that really tell the animals' stories," Sabin said.