Age structure of a population of Ixodes ricinus (Acari: Ixodidae) in relation to its seasonal questing.

Three sites in south west Scotland, each having one deciduous and one coniferous woodland with populations of roe deer, were sampled over three years for Ixodes ricinus Linnaeus ticks using cloth drags. Nymphs and adult ticks were age graded by dissection of gut and Malpighian tubules and staining for lipid with Sudan red. Temperature and relative humidity were recorded in developmental and questing microclimates. Two annual cohorts of both nymphs and adults were found. For both instars one cohort became apparent in early spring and the other in autumn. The timing of the cohorts of nymphs corresponded to peaks of numbers questing, but neither females nor males had any clear seasonal pattern of questing. The maximum life of questing nymphs and adults was three to four months. Nymphs and adults were found questing at all months of the year and at temperatures ranging from 3.5 degrees C to 29.3 degrees C. Questing of larvae in summer peaks of numbers was positively correlated with temperature and negatively correlated with relative humidity. These results are explained by a descriptive model which invokes behavioural and morphogenetic diapause. Tests of the model are discussed and it is proposed that the main adaptive advantage of the complex life cycle is to permit moulting and oviposition to occur at favourable summer temperatures in ticks that have engorged at any time of the year.