Generation cycles in experimental populations of a multivoltine insect.

Some animals exhibit large-amplitude fluctuations with periods close to their generation length. These fluctuations can be caused by seasonal environmental influences, scheduled pesticide applications, or ecological factors such as intraspecific regulation and consumer-resource interactions. While theory provides various mechanisms for how environmental and ecological factors might generate generational fluctuations, there has never been a field experiment testing the relative contributions of seasonal demographic synchronisation and intraspecific regulation to generation cycles in natural populations. The smaller tea tortrix, Adoxophyes honmai, is a serious pest of tea plants and a temperate multivoltine insect that undergoes 3-5 large-amplitude generation cycles each year under natural conditions in Japan. Theory suggests that these fluctuations may represent limit cycles driven by asymmetric intraspecific interference, where older larvae directly affect younger ones. However, in the field, these populations also experience strong seasonality and periodic insecticide applications that are thought to either generate or modify these fluctuations. We conducted a replicated field-cage experiment on the tortrix populations to manipulate the initial degree of stage synchrony and the timing of introduction. The experiment included four treatments contrasting pulsed versus continuous age structures at the onset of spring, along with two introduction timings separated by 20 days (approximately half a generation time). We minimised artificial interventions, such as harvesting and insecticide application, as well as the effect of natural enemies while allowing meteorological influences during the season. To compare the field-cage experiment results with theoretical predictions, we constructed an age-structured population model featuring asymmetric larval interference. We executed simulations using the same introduction scenarios as in the field-cage experiment. We observed the emergence of clear generational cycles in all treatments of the field-cage even in the absence of any initial demographic synchrony. This suggests an internal mechanism regulating population cycles, possibly intraspecific interference. However, the generational cycles in the field-cage were synchronised across treatments and with outer field populations. The results from the field-cage experiment and simulation analyses indicate that external environmental factors, such as precipitation, acted as a pacemaker for the generational cycles created by the internal regulatory mechanism.