Female Monarch depositing egg on Common Milkweed (Asclepias syriaca)

Evaluating the Migration Mortality Hypothesis Using Monarch Tagging Data

Frontiers in Ecology and Evolution

August 7, 2020


Orley Talor, Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence, KS, United States

John M. Pleastant, Department of Ecology, Evolution and Organismal Biology, Iowa State University, Ames, IA, United States

Ralph Grundel, U.S. Geological Survey, Great Lakes Science Center, Chesterton, IN, United States

Samulel D. Pecoraro, U.S. Geological Survey, Great Lakes Science Center, Chesterton, IN, United States

James P. Lovett, Monarch Watch, Kansas Biological Survey, University of Kansas, Lawrence, KS, United States

Ann Ryan, Monarch Watch, Kansas Biological Survey, University of Kansas, Lawrence, KS, United States

The decline in the eastern North American population of the monarch butterfly population since the late 1990s has been attributed to the loss of milkweed during the summer breeding season and the consequent reduction in the size of the summer population that migrates to central Mexico to overwinter (milkweed limitation hypothesis). However, in some studies the size of the summer population was not found to decline and was not correlated with the size of the overwintering population. The authors of these studies concluded that milkweed limitation could not explain the overwintering population decline. They hypothesized that increased mortality during fall migration was responsible (migration mortality hypothesis). We used data from the long-term monarch tagging program, managed by Monarch Watch, to examine three predictions of the migration mortality hypothesis: (1) that the summer population size is not correlated with the overwintering population size, (2) that migration success is the main determinant of overwintering population size, and (3) that migration success has declined over the last two decades. As an index of the summer population size, we used the number of wild-caught migrating individuals tagged in the U.S. Midwest from 1998 to 2015. As an index of migration success we used the recovery rate of Midwest tagged individuals in Mexico. With regard to the three predictions: (1) the number of tagged individuals in the Midwest, explained 74% of the variation in the size of the overwintering population. Other measures of summer population size were also correlated with overwintering population size. Thus, there is no disconnection between late summer and winter population sizes. (2) Migration success was not significantly correlated with overwintering population size, and (3) migration success did not decrease during this period. Migration success was correlated with the level of greenness of the area in the southern U.S. used for nectar by migrating butterflies. Thus, the main determinant of yearly variation in overwintering population size is summer population size with migration success being a minor determinant. Consequently, increasing milkweed habitat, which has the potential of increasing the summer monarch population, is the conservation measure that will have the greatest impact.


Since the late 1990s, the monarch butterfly, Danaus plexippus, population has declined significantly based on measurements made at the Mexican overwintering grounds (Brower et al., 2011; Semmens et al., 2016). Identifying the cause or causes of the decline is important in order to focus conservation measures appropriately. Two explanations for the decline in the size of the overwintering population dominate the literature. The first, known as the “milkweed limitation” hypothesis, posits that the decline in the number of milkweed host plants in the major summer breeding area in the Upper Midwest of the U.S. (Figure 1) has led to a reduction in the size of the migratory population (Pleasants et al., 2017). The second, known as the “migration mortality” hypothesis, posits that the resources and conditions during the fall migration have declined resulting in an increase in mortality during the migration and a decline in the overwintering population (Agrawal and Inamine, 2018).

Figure 1. All wild-caught butterflies tagged from north of 40° latitude and east of 100° longitude were included in the study. This area includes the region we are calling the Midwest, encompassing the area from 40 to 50° latitude and 80 to 100° longitude (outlined in red) and the region we are calling the Northeast, encompassing the area from 40 to 50° latitude and 65 to 80° longitude (outlined in blue). What we are calling the Total Area is the Midwest and Northeast combined. The NDVI values (Saunders et al., 2019) come from the region that encompasses the area from 30 to 40° latitude and 90 to 105° longitude (outlined in green). The dark blue square indicates the location of the overwintering colonies. Butterflies were tagged in other sectors besides the Midwest and Northeast but those data are not included in this study.

The milkweed limitation hypothesis is supported by data showing that in the early 2000s the majority of monarch production came from common milkweed, Asclepias syriaca, in corn and soybean fields in the Midwest (Oberhauser et al., 2001) and that the abundance of those milkweeds declined precipitously due to glyphosate herbicide use in those fields (Pleasants and Oberhauser, 2013; Flockhart et al., 2015; Pleasants et al., 2017; Thogmartin et al., 2017a; Saunders et al., 2018). The loss of the milkweeds from corn and soybean fields began in the late 1990s with the adoption of glyphosate-tolerant crops. Milkweeds had been nearly eliminated from these fields by 2006 (Pleasants, 2017). During this period, an estimated 71% of the monarch production potential of milkweeds on the Midwest landscape was eliminated, amounting to 25 million hectares of agricultural habitat that no longer had milkweeds (Pleasants, 2017). The subsequent decrease in the availability of milkweed is thought to have limited the size of the summer breeding population. Support for this hypothesis comes from the pattern of decline in milkweed availability that parallels the decline in the size of the overwintering population (Pleasants et al., 2017). Further support comes from the strong correlation between yearly late summer Midwest monarch egg production and yearly overwintering population size (Pleasants and Oberhauser, 2013; Pleasants et al., 2017).


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