Multiscale seasonal factors drive the size of winter monarch colonies.

Monarch butterflies in eastern North America have declined by 84% on Mexican wintering grounds since the observed peak in 1996. However, coarse-scale population indices from northern US breeding grounds do not show a consistent downward trend. This discrepancy has led to speculation that autumn migration may be a critical limiting period. We address this hypothesis by examining the role of multiscale processes impacting monarchs during autumn, assessed using arrival abundances at all known winter colony sites over a 12-y period (2004-2015). We quantified effects of continental-scale (climate, landscape greenness, and disease) and local-scale (colony habitat quality) drivers of spatiotemporal trends in winter colony sizes. We also included effects of peak summer and migratory population indices. Our results demonstrate that higher summer abundance on northern breeding grounds led to larger winter colonies as did greener autumns, a proxy for increased nectar availability in southern US floral corridors. Colony sizes were also positively correlated with the amount of local dense forest cover and whether they were located within the Monarch Butterfly Biosphere Reserve, but were not influenced by disease rates. Although we demonstrate a demographic link between summer and fine-scale winter population sizes, we also reveal that conditions experienced during, and at the culmination of, autumn migration impact annual dynamics. Monarchs face a growing threat if floral resources and winter habitat availability diminish under climate change. Our study tackles a long-standing gap in the monarch's annual cycle and highlights the importance of evaluating migratory conditions to understand mechanisms governing long-term population trends.

Population Genetics of Overwintering Monarch Butterflies, Danaus plexippus (Linnaeus), from Central Mexico Inferred from Mitochondrial DNA and Microsatellite Markers.

Population genetic variation and demographic history in Danaus plexippus (L.), from Mexico were assessed based on analyses of mitochondrial cytochrome c oxidase subunit I (COI; 658 bp) and subunit II (COII; 503 bp) gene segments and 7 microsatellite loci. The sample of 133 individuals included both migratory monarchs, mainly from 4 overwintering sites within the Monarch Butterfly Biosphere Reserve (MBBR) in central Mexico (states of Michoacán and México), and a nonmigratory population from Irapuato, Guanajuato. Haplotype (h) and nucleotide (π) diversities were relatively low, averaging 0.466 and 0.00073, respectively, for COI, and 0.629 and 0.00245 for COII. Analysis of molecular variance of the COI data set, which included additional GenBank sequences from a nonmigratory Costa Rican population, showed significant population structure between Mexican migratory monarchs and nonmigratory monarchs from both Mexico and Costa Rica, suggesting limited gene flow between the 2 behaviorally distinct groups. Interestingly, while the COI haplotype frequencies of the nonmigratory populations differed from the migratory, they were similar to each other, despite the great physical distance between them. Microsatellite analyses, however, suggested a lack of structure between the 2 groups, possibly owing to the number of significant deviations from Hardy-Weinberg equilibrium resulting from heterzoygote deficiencies found for most of the loci. Estimates of demographic history of the combined migratory MBBR monarch population, based on the mismatch distribution and Bayesian skyline analyses of the concatenated COI and COII data set (n = 89) suggested a population expansion dating to the late Pleistocene (~35000-40000 years before present) followed by a stable effective female population size (Nef) of about 6 million over the last 10000 years.

Trends in deforestation and forest degradation after a decade of monitoring in the Monarch Butterfly Biosphere Reserve in Mexico.

We used aerial photographs, satellite images, and field surveys to monitor forest cover in the core zones of the Monarch Butterfly Biosphere Reserve in Mexico from 2001 to 2012. We used our data to assess the effectiveness of conservation actions that involved local, state, and federal authorities and community members (e.g., local landowners and private and civil organizations) in one of the world's most iconic protected areas. From 2001 through 2012, 1254 ha were deforested (i.e., cleared areas had <10% canopy cover), 925 ha were degraded (i.e., areas for which canopy forest decreased), and 122 ha were affected by climatic conditions. Of the total 2179 ha of affected area, 2057 ha were affected by illegal logging: 1503 ha by large-scale logging and 554 ha by small-scale logging. Mexican authorities effectively enforced efforts to protect the monarch reserve, particularly from 2007 to 2012. Those efforts, together with the decade-long financial support from Mexican and international philanthropists and businesses to create local alternative-income generation and employment, resulted in the decrease of large-scale illegal logging from 731 ha affected in 2005-2007 to none affected in 2012, although small-scale logging is of growing concern. However, dire regional social and economic problems remain, and they must be addressed to ensure the reserve's long-term conservation. The monarch butterfly (Danaus plexippus) overwintering colonies in Mexico-which engage in one of the longest known insect migrations-are threatened by deforestation, and a multistakeholder, regional, sustainable-development strategy is needed to protect the reserve.

The monarch butterfly in Mexico: a conservation model.

Each fall, millions of monarch butterflies (Danaus plexippus L.) travel from Canada and the United States to overwinter in Mexico and California. In 2022, the IUCN listed migratory monarchs as endangered because of their population decline. The main accepted drivers are widespread use of herbicides, effects of climate, and land use change that causes habitat loss. We analyzed the main actions taken to officially protect the overwintering sites and the migration phenomenon with the establishment of the Monarch Butterfly Biosphere Reserve in 2000. The conservation of the monarch overwintering sites in Mexico is an example of continuous work from their discovery to the present. We highlight the importance of monitoring the areas covered by overwintering monarchs in Mexico. These colonies represent the largest concentrations of monarch populations in the world. In the last 10 years, the average area covered by monarchs was 2.72 ( ± 0.47 SE) hectares.

Changes in climate drive recent monarch butterfly dynamics.

Declines in the abundance and diversity of insects pose a substantial threat to terrestrial ecosystems worldwide. Yet, identifying the causes of these declines has proved difficult, even for well-studied species like monarch butterflies, whose eastern North American population has decreased markedly over the last three decades. Three hypotheses have been proposed to explain the changes observed in the eastern monarch population: loss of milkweed host plants from increased herbicide use, mortality during autumn migration and/or early-winter resettlement and changes in breeding-season climate. Here, we use a hierarchical modelling approach, combining data from >18,000 systematic surveys to evaluate support for each of these hypotheses over a 25-yr period. Between 2004 and 2018, breeding-season weather was nearly seven times more important than other factors in explaining variation in summer population size, which was positively associated with the size of the subsequent overwintering population. Although data limitations prevent definitive evaluation of the factors governing population size between 1994 and 2003 (the period of the steepest monarch decline coinciding with a widespread increase in herbicide use), breeding-season weather was similarly identified as an important driver of monarch population size. If observed changes in spring and summer climate continue, portions of the current breeding range may become inhospitable for monarchs. Our results highlight the increasingly important contribution of a changing climate to insect declines.

Are female monarch butterflies declining in eastern North America? Evidence of a 30-year change in sex ratios at Mexican overwintering sites.

Every autumn the entire eastern North American population of monarch butterflies (Danaus plexippus) undergoes a spectacular migration to overwintering sites in the mountains of central Mexico, where they form massive clusters and can number in the millions. Since their discovery, these sites have been extensively studied, and in many of these studies, monarchs were captured and sexes recorded. In a recent effort to compile the sex ratio data from these published records, a surprising trend was found, which appears to show a gradual decline in proportion of females over time. Sex ratio data from 14 collections of monarchs, all spanning 30 years and totaling 69 113 individuals, showed a significant negative correlation between proportion of females and year (r = -0.69, p = 0.007). Between 1976 and 1985, 53 per cent of overwintering monarchs were female, whereas in the last decade, 43 per cent were female. The relationship was significant with and without weighting the analyses by sampling effort. Moreover, analysis of a recent three-year dataset of sex ratios revealed no variation among nine separate colonies, so differences in sampling location did not influence the trend. Additional evidence from autumn migration collections appears to confirm that proportions of females are declining, and also suggests the sex ratio is shifting on breeding grounds. While breeding monarchs face a number of threats, one possibility is an increase in prevalence of the protozoan parasite, Ophryocystis elektroscirrha, which recent evidence shows affects females more so than males. Further study will be needed to determine the exact cause of this trend, but for now it should be monitored closely.