JOHN M. PLEASANTS1 and KAREN S. OBERHAUSER2 1 Department of Ecology, Evolution and Organismal Biology, Iowa State University, Ames, IA, USA and 2 Department of Fisheries, Wildlife and Conservation Biology, University of Minnesota, St Paul, MN, USA

Abstract. 1. The size of the Mexican overwintering population of monarch butter- flies has decreased over the last decade. Approximately half of these butterflies come from the U.S. Midwest where larvae feed on common milkweed. There has been a large decline in milkweed in agricultural fields in the Midwest over the last decade. This loss is coincident with the increased use of glyphosate herbicide in conjunction with increased planting of genetically modified (GM) glyphosate-tolerant corn (maize) and soybeans (soya). 2. We investigate whether the decline in the size of the overwintering population can be attributed to a decline in monarch production owing to a loss of milkweeds in agricultural fields in the Midwest. We estimate Midwest annual monarch production using data on the number of monarch eggs per milkweed plant for milkweeds in different habitats, the density of milkweeds in different habitats, and the area occupied by those habitats on the landscape. 3. We estimate that there has been a 58% decline in milkweeds on the Midwest landscape and an 81% decline in monarch production in the Midwest from 1999 to 2010. Monarch production in the Midwest each year was positively correlated with the size of the subsequent overwintering population in Mexico. Taken together, these results strongly suggest that a loss of agricultural milkweeds is a major contributor to the decline in the monarch population. 4. The smaller monarch population size that has become the norm will make the species more vulnerable to other conservation threats.

Key words. Glyphosate, GMO, milkweed, monarch butterfly Introduction Monarch butterflies (Danaus plexippus L. Lepidoptera: Danainae) in the Eastern North American migratory population undergo a multi-generation annual cycle that includes wintering in central Mexico. In the spring, adults that have overwintered migrate north and reproduce in Texas and states to the north and east. Their offspring move farther north into much of the eastern half of the United States and southern Canada, and two to three more generations are produced (Cockrell et al., 1993; Malcolm et al., 1993; Prysby & Oberhauser, 2004). Most adults that emerge after mid-August are in a state of reproductive diapause (Herman, 1985; Goehring & Oberhauser, 2002) and…