Raptor predation and population limitation in red grouse

Summary 1.  We assessed the impact of predation by hen harriers and peregrine falcons on a red grouse population in southern Scotland during 1992–98. Grouse density in April, July and October declined during this time, coincident with an increase in the numbers of breeding harriers and peregrines. 2.  Winter losses of grouse between October and April averaged 33% and were density‐dependent. Raptors were the cause of about 70% of winter mortality and they killed about 30% of the grouse present in October. We were unable to determine whether winter mortality in raptors was additive to other losses. 3.  Summer losses of adult grouse between April and July averaged 30% and were density‐dependent. Raptors were the cause of more than 90% of the early summer mortality of adult grouse. Summer losses of grouse chicks between May and July averaged 45% and were not density‐dependent. Harriers killed about 28% of grouse chicks by late July and about 37% by the end of August. Summer raptor predation on adult grouse and chicks appeared to be largely additive to other losses and we estimated that it reduced autumn grouse densities by about 50%. 4.  A model combining the estimated reduction in autumn grouse density caused by raptors with the observed density dependence in winter loss predicted that, in the absence of raptors for 2 years, grouse density in spring would be 1·9 times greater, and grouse density in autumn 3·9 times greater, than in the presence of raptors. The model suggested that raptor predation prevented the grouse population from increasing and was thus a limiting factor.