Seed production and seed predation in a patchy and time-varying environment. Dynamics of a milkweed ? tephritid fly system

Abstract
Vincetoxicum hirundinaria is a longlived perennial herb. Its pod production was measured during seven years in about 40 small isolated patches in two study areas in southeastern Sweden. Total pod production in the two areas varied synchronously and up to about 200 times between years. This variation was mainly due to drought conditions as determined by the interplay of weather (sunshine h) and site characteristics. Populations of a univoltine tephritid fly, euphranta connexa, whose larva feeds on V. hirundinaria seeds in the maturing pods, were monitored for five and seven years in the two study areas respectively. Larval fly populations in the two areas varied 30 and 50-fold between years, with highs in years when pods were abundant and lows when pods were scarce. The percentage of pods attacked by E. connexa, however, varied in the opposite direction, with very high attack rates (about 100%) when pods were scarce and low rates (down to 10–20%) when pods were common. Thus the temporal tracking of food resources by E. connexa was poor, resulting in yearly variations in the amount of unattacked pods (and seeds) being about 2,00-fold. Many host plant patches were small and totally without pods in some years, causing frequent local extinctions of E. connexa. Patches were, however, readily colonized in later years when pods appeared again. Low overall attack rates in certain years were thus only to a minor extent explainable by host plant patches being uncolonized by the fly. The efficient spatial tracking of resources by the fly population allows this population system to be analyzed largely in terms of its temporal dynamics. Although E. connexa populations often exploit only a minor part of their potential larval resources, the rate of population change was closely correlated with these resources expressed on a per capita basis. Natural enemies only have negligible effects on E. connexa population trends. In this population system the herbivore (seed predator) population is thus controlled by its food resources, but weather-imposed temporal variations in resource set are too large for the herbivore population to closely track its resource base. This temporal tracking inertia has important implications for the long-term production of healthy seeds.