Louping ill virus in the UK: a review of the hosts, transmission and ecological consequences of control

Louping ill virus (LIV) is a tick-borne flavivirus that is part of the tick-borne encephalitis complex of viruses (TBEV) and has economic and welfare importance by causing illness and death in livestock, especially sheep, Ovies aries, and red grouse, Lagopus lagopus scoticus, an economically valuable gamebird. Unlike Western TBEV which is found primarily in woodlands and is reservoired by small rodents, LIV is not generally transmitted by small rodents but instead by sheep, red grouse and mountain hares and, therefore, is associated with upland heather moorland and rough grazing land. Red grouse are a particularly interesting transmission host because they may acquire most of their LIV infections through eating ticks rather than being bitten by ticks. Furthermore, the main incentive for the application of LIV control methods is not to protect sheep, but to protect red grouse, which is an economically important gamebird. The widespread intensive culling of mountain hares which has been adopted in several areas of Scotland to try to control ticks and LIV has become an important issue in Scotland in recent years. This review outlines the reservoir hosts and transmission cycles of LIV in the UK, then describes the various control methods that have been tried or modelled, with far-reaching implications for conservation and public opinion.