Beech tree masting explains the inter-annual variation in the fall and spring peaks of Ixodes ricinus ticks with different time lags

Abstract

BackgroundThe tick Ixodes ricinus is an important vector of tick-borne diseases including Lyme borreliosis. In continental Europe, the nymphal stage of I. ricinus often has a bimodal phenology with a large spring/early summer peak and a smaller fall peak. While there is consensus about the origin of the spring nymphal peak, there are two alternative hypotheses for the fall nymphal peak, direct development versus delayed diapause. These two hypotheses make different predictions about the time lags of the correlations between the spring peak, the fall peak, and seed production (masting) by deciduous trees. MethodsTo determine which hypothesis is most important for explaining the fall peak, we used data from a long-term surveillance study (15 years) on the density of I. ricinus nymphal ticks at 4 different elevation sites in an area of Switzerland that is endemic for Lyme borreliosis, and long-term data on the mast of the European beech tree from the literature. ResultsI. ricinus nymphs had a bimodal phenology at the three lower elevation sites, but a unimodal phenology at the top elevation site. At the lower elevation sites, the density of nymphs (DON) in the fall was strongly correlated with the DON in the spring of the following year. The inter-annual variation in the densities of I. ricinus nymphs in the fall and spring were best explained by a 1-year versus a 2-year time lag with the beech tree masting index. Fall nymphs had higher fat content and are younger than spring nymphs. All of these observations are consistent with the direct development hypothesis for the fall peak of I. ricinus nymphs at our study site. Our study provides new insight into the complex bimodal phenology of this important disease vector. ConclusionsPublic health officials in Europe should be aware that following a strong mast year, the DON will increase 1 year later in the fall and 2 years later in the spring and summer. Population ecology studies of I. ricinus should consider that the spring and fall peak in the same calendar year represent different generations of ticks.