The lion Panthera leo (Linnaeus, 1758) is a charismatic large cat that has been imported into Europe since early historic times. Lions were amongst the many exotic animals that were imported to Rome during the early Imperial Period for the gladiatorial games, although towards the end of the Roman Empire the increased scarcity of these animals in the wild forced combat shows to be largely replaced by exhibitions (Baratay and Hardouin-Fugier, 2002). Exotic wild animals do not appear to have been kept regularly in western Europe until the 13th century, when they were rediscovered through Western contact with the Byzantine and Muslim worlds (Baratay and Hardouin-Fugier, 2002). In England, the Royal Menagerie was established in 12th-13th centuries in Wood-stock near Oxford, and slightly later was relocated to the Tower of London, where the first residents were three leopards sent to Henry III by the Holy Roman Emperor Frederick II in 1235 (Hahn, 2003).
Although the Royal Menagerie and its animals are known from documentary records, few physical remains survive (O’Regan et al., 2005). Amongst the rare exceptions are two lion skulls that were recovered from the moat of the Tower of London during excavations in 1936-1937. These skulls were recently radiocarbon-dated to AD1280-1385 and AD1420-1480, making them the earliest confirmed lion remains in the British Isles since the extinction of the Pleistocene cave lion (P. l. spelaea) (O’Regan et al., 2005). The discovery of these first English lions attracted significant media attention (BBC, 2005). However, the geographical origin of these animals has not yet been investigated. Such knowledge would provide novel insights not only into the history of the Royal Menagerie, but also into patterns of animal trafficking during the Medieval period. Direct animal trade between Europe and sub-Saharan Africa was not well developed until the 18th century (Anonymus, 1876). Therefore, it may be reasonable to presume that the Tower lions were unlikely to have originated from sub-Saharan regions. Nevertheless, there is an undeniable possibility that sub-Saharan lions reached Europe as they could have reached shipping ports in North Africa and the Middle East through trans-Saharan trade routes that were well established by the early Medieval period (Yamaguchi, 2000b). Apart from a tiny population in northwest India, lions had been practically exterminated outside sub-Saharan Africa by the turn of the 20th century (Yamaguchi and Haddane, 2002; Patterson, 2004). In this context, if the foregoing first hypothesis turned out to be the case, the Tower lion skulls would possess significant value for the history of the lion, as well as the history of Medieval England.
Recent advances in ancient DNA (aDNA) techniques (e.g. Shapiro et al., 2004), in association with the available data concerning genetic profiles of the lion across its natural range (Dubach et al., 2005; Barnett et al., 2006a, 2006b) have made it possible to identify the origins of unprovenanced lion specimens, such as the Tower lions (Barnett et al., 2007).
In this paper we use aDNA techniques to extract and amplify mitochondrial DNA (mtDNA) from the two Tower lions, and compare the results with those of Barnett et al. (2006a, 2006b). We also conduct a cranio-morphometric analysis to investigate which lion population the Tower lions are morphologically similar to. Then, by combining both molecular and morphological results, we will try to determine the geographic origin of the first lions in England.