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Among cyclic populations of herbivores, inter-specific temporal synchrony has been attributed to both climatic factors and trophic interactions. In northern Europe, winter and autumnal moths undergo regular 9–11 year population cycles. The winter moth cycle has typically been phase-locked with that of the autumnal moth, but with a 1–3- year phase lag. We examined potential effects of natural enemies on this phase lag using field experiments and observational data. We found that larval parasitism was significantly higher in autumnal than in winter moths. Conversely, pupal predation by generalist invertebrates was clearly greater in winter than in autumnal moths. The difference in parasitism rates may contribute to the earlier collapse of the autumnal moth cycle. In addition, the phase lag may be strengthened by higher pupal mortality in winter moths in the early increase phase of the cycles. As a consequence, we put forward a hypothesis on reversed effects of natural enemies, providing a potential explanation for phase-lagged population cycles of these moth species.
1. Generalist natural enemies are usually not considered as being capable of causing population cycles in forest insects, but they may influence the population dynamics of their prey in the low density cycle phase when specialist enemies are largely absent. 2. In the present field study, the total response of the generalist invertebrate predator community to experimentally established pupal densities of the closely related autumnal (Epirrita autumnata) and winter moths (Operophtera brumata) was analysed. 3. Due to the high amount of variation in the dataset, the exact shape of the response curve could not be convincingly estimated. Nevertheless, two important conclusions can be drawn from the analyses. 4. Firstly, the natural invertebrate predator community seems to become saturated at rather low densities of both autumnal and winter moth pupae. Secondly, the predator community seems to become saturated at much lower densities of autumnal than of winter moth pupae. 5. Furthermore, pupal mass was significantly negatively correlated with invertebrate predation probability in autumnal moth pupae. 6. These results indicate that differences in the predator assemblage being able to consume pupae of the two moth species, as well as different handling times, could be responsible for the substantially higher predation rates in winter than in autumnal moth pupae. 7. As a consequence, the population dynamics of autumnal moths might be less affected by generalist invertebrate predators than those of winter moths, as autumnal moths seem able to escape from the regulating influence of generalist predators at much lower population densities than winter moths.
1. Both direct and indirect competition can have profound effects on species abundance and expansion rates, especially for a species trying to strengthen a foothold in new areas, such as the winter moth (Operophtera brumata) currently in northernmost Finland. There, winter moths have overlapping outbreak ranges with autumnal moths (Epirrita autumnata), who also share the same host, the mountain birch (Betula pubescens ssp. czerepanovii). Competitive interactions are also possible, but so far unstudied, are explanations for the observed 1–3 years phase lag between the population cycles of the two moth species. 2. In two field experiments, we studied host plant-mediated indirect inter-specific competition and direct interference/exploitation competition between autumnal and winter moths. The experimental larvae were grown either with the competing species or with the same number of conspecifics until pupation. Inter-specific competition was judged from differences in pupal mass (reflecting lifespan fecundity), larval development time and larval survival. 3. Larval performance measurements suggested that neither direct nor indirect interspecific competition with the autumnal moth reduce the growth rate of winter moth populations. Winter moths even had a higher probability of survival when reared together with autumnal moths. 4. Thus, we conclude that neither direct nor indirect inter-specific competition is capable of suppressing the spread of the winter moth outbreak range and that both are also an unlikely cause for the phase lag between the phase-locked population cycles of the two moth species.