Abstract
Many populations of forest Lepidoptera exhibit 10-year cycles in densities, with impressive outbreaks across large regions. Delayed density-dependent interactions with natural enemies are recognized as key factors driving these cyclic population dynamics, but emphasis has typically been on the larval stages. Eggs, pupae and adults also suffer mortality from predators, parasitoids and pathogens, but little is known about possible density relationships between mortality factors and these non-feeding life stages. In a long-term field study, we experimentally deployed autumnal moth (Epirrita autumnata) eggs and pupae to their natural enemies yearly throughout the 10-year population cycle in northern Norway. The abundance of another geometrid, the winter moth (Operophtera brumata), increased in the study area, permitting comparisons between the two moth species in predation and parasitism. Survival of autumnal moth eggs and pupae was related to the moth abundance in an inverse and delayed manner. Egg and pupal parasitoids dominated as density-dependent mortality factors and predicted the subsequent growth rate of the host population size. In contrast, effects of egg and pupal predators were weakly density dependent, and generally predation remained low. Parasitism rates did not differ between the autumnal and winter moth pupae, whereas predators preferred winter moth pupae over those of the autumnal moth. We conclude that parasitism of the autumnal moth by egg and pupal parasitoids can be related to the changes of the moth density in a delayed density-dependent manner. Furthermore, egg and pupal parasitoids cannot be overlooked as causal factors for the population cycles of forest Lepidoptera in general.
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Acknowledgments
We thank all field assistants for their great help with the fieldwork. Ph.D. students and post doc researchers of the earlier autumnal moth projects, Tea Ammunét, Netta Klemola and Annette Scheiner, are especially acknowledged. Reijo Jussila, Ilari E. Sääksjärvi and Veli Vikberg are gratefully thanked for their help in parasitoid identification. We also thank Thomas Hoffmeister, Judith H. Myers, Seppo Neuvonen, Jens Roland and an anonymous referee for their valuable comments on the manuscript, the staff of the Kevo Subarctic Research Station for excellent working facilities and the staff of the Statskog for permissions to conduct our research on public lands in Norway. The research was supported financially by the Academy of Finland (projects 111195, 129143 and 204190 to T. K. and 7650, 34509 and 48697 to K. R.), by the Turku University Foundation (grants to T. K. and K. R.), and by the Maj and Tor Nessling Foundation (grants to T. K.).
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Communicated by Thomas S. Hoffmeister.
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Online Resource 1 Map of northern Europe showing outbreak range of the autumnal and winter moth and main study locations of the moths in northern Fennoscandia (DOCX 124 kb)
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Online Resource 2 Likelihood-based model comparisons and parameter estimates of GLMMs on the survival, predation, and parasitism of autumnal moth eggs and pupae (DOCX 32 kb)
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Online Resource 4 Information on parasitoids and potential predators of the pupae and eggs of autumnal and winter moths (DOCX 21 kb)
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Klemola, T., Andersson, T. & Ruohomäki, K. Delayed density-dependent parasitism of eggs and pupae as a contributor to the cyclic population dynamics of the autumnal moth. Oecologia 175, 1211–1225 (2014). https://doi.org/10.1007/s00442-014-2984-9
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DOI: https://doi.org/10.1007/s00442-014-2984-9