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Elucidating the adaptations that promote flight in animals can aid the understanding of evolution and species divergence, and/or provide inspiration for aerospace engineering and the design of better aerial vehicles. The famed long-distance migration of monarch butterflies in North America still holds many questions and opportunities for inspiration. For example, there is little research on whether the monarch's primary wing colors themselves (black, orange, or white) have any aerodynamic or migration function. Dark colors on wings of other animals have recently been shown to aid flight by enhancing solar absorption, which reduces drag forces. However, too much black surface could be problematic for monarchs, which are exposed to increasing amounts of solar energy along their flightpath. This paper describes the results of two related investigations that attempt to elucidate the importance of wing color to the monarch migration. By measuring the color proportions of nearly 400 monarch wings collected at different stages of their journey, we found, surprisingly, that successful migrants tended to have less black on their wings (about 3% less), but also more white pigment (about 3% more); monarchs have a band of light-colored marginal wing spots. Second, image analysis of museum specimens revealed migratory monarchs had significantly larger white spots, proportional to the wing area, than most non-migratory, New World Danaid butterflies, which argues spot size has evolved along with migratory behavior. Combined, these findings strongly suggest that the long-distance migration itself selects for larger white spots every fall, so that only those individuals with large spots will survive to pass on their genes. Further experimental work is needed to elucidate how the spots aid the migration, but it is possible that they enhance aerodynamic efficiency; other work by the authors demonstrates how alternating white and black pigment on wings can reduce drag. These results will serve as a useful starting point for such endeavors, which should improve understanding of one of the world's most fascinating animal migrations, and also provide practical knowledge for the field of aerospace engineering., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2023 Davis et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Monarch butterflies in North America have an exceptionally large breeding distribution, occupying regions west of the Rocky Mountains and throughout the eastern seaboard. An experimental study conducted 17 years ago and published in this journal appeared to show that western monarch larvae tend to have smaller black stripes than those from eastern parents, which at the time was thought to be an adaptation to higher solar exposure in California. Here, we revisit this question by measuring melanism of eastern and western larvae from online photographs submitted to iNaturalist by members of the public. We downloaded over 500 photos of larvae from 10 different states (4 in the west, 6 in the east) and used image analysis to quantify the size of each larva's black stripes (% of surface covered by black). We compared average melanism level between east and west, while also accounting for temperature (at the photo location), seasonal effects, and latitude. Results showed larvae tended to be darker with lower development temperatures, and later in the season, though there was no significant difference between eastern and western larvae in their degree of melanism. It is not yet clear why findings here were different from the prior experimental study. Also, the wild larvae appeared to be notably lighter in color than the captive-reared larvae of the original study, suggesting that the captive environment does not truly replicate the developmental experience of wild larvae, at least in terms of cuticular development. Thus, if there truly are innate tendencies for western larvae to differ (morphologically) from eastern, they are not observable in nature., Competing Interests: Declaration of competing interest The authors declare there are no competing interests., (Copyright © 2022. Published by Elsevier Ltd.)

Many insects are in clear decline, with monarch butterflies (Danaus plexippus) drawing particular attention as a flagship species. It is well documented that, among migratory populations, numbers of overwintering monarchs have been falling across several decades, but trends among breeding monarchs are less clear. Here, we compile >135,000 monarch observations between 1993 and 2018 from the North American Butterfly Association's annual butterfly count to examine spatiotemporal patterns and potential drivers of adult monarch relative abundance trends across the entire breeding range in eastern and western North America. While the data revealed declines at some sites, particularly the US Northeast and parts of the Midwest, numbers in other areas, notably the US Southeast and Northwest, were unchanged or increasing, yielding a slightly positive overall trend across the species range. Negative impacts of agricultural glyphosate use appeared to be counterbalanced by positive effects of annual temperature, particularly in the US Midwest. Overall, our results suggest that population growth in summer is compensating for losses during the winter and that changing environmental variables have offsetting effects on mortality and/or reproduction. We suggest that density-dependent reproductive compensation when lower numbers arrive each spring is currently able to maintain relatively stable breeding monarch numbers. However, we caution against complacency since accelerating climate change may bring growing threats. In addition, increases of summer monarchs in some regions, especially in California and in the south, may reflect replacement of migratory with resident populations. Nonetheless, it is perhaps reassuring that ubiquitous downward trends in summer monarch abundance are not evident., (© 2022 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

Insect-pathogen dynamics can show seasonal and inter-annual variations that covary with fluctuations in insect abundance and climate. Long-term analyses are especially needed to track parasite dynamics in migratory insects, in part because their vast habitat ranges and high mobility might dampen local effects of density and climate on infection prevalence. Monarch butterflies Danaus plexippus are commonly infected with the protozoan Ophryocystis elektroscirrha (OE). Because this parasite lowers monarch survival and flight performance, and because migratory monarchs have experienced declines in recent decades, it is important to understand the patterns and drivers of infection. Here we compiled data on OE infection spanning 50 years, from wild monarchs sampled in the United States, Canada and Mexico during summer breeding, fall migrating and overwintering periods. We examined eastern versus western North American monarchs separately, to ask how abundance estimates, resource availability, climate and breeding season length impact infection trends. We further assessed the intensity of migratory culling, which occurs when infected individuals are removed from the population during migration. Average infection prevalence was four times higher in western compared to eastern subpopulations. In eastern North America, the proportion of infected monarchs increased threefold since the mid-2000s. In the western region, the proportion of infected monarchs declined sharply from 2000 to 2015, and increased thereafter. For both eastern and western subpopulations, years with greater summer adult abundance predicted greater infection prevalence, indicating that transmission increases with host breeding density. Environmental variables (temperature and NDVI) were not associated with changes in the proportion of infected adults. We found evidence for migratory culling of infected butterflies, based on declines in parasitism during fall migration. We estimated that tens of millions fewer monarchs reach overwintering sites in Mexico as a result of OE, highlighting the need to consider the parasite as a potential threat to the monarch population. Increases in infection among eastern North American monarchs post-2002 suggest that changes to the host's ecology or environment have intensified parasite transmission. Further work is needed to examine the degree to which human practices, such as mass caterpillar rearing and the widespread planting of exotic milkweed, have contributed to this trend., (© 2022 British Ecological Society.)

For many animals and insects that are experiencing dramatic population declines, the only recourse for conservationists is captive rearing. To ensure success, reared individuals should be biologically indistinct from those in the wild. We tested if this is true with monarch butterflies, Danaus plexippus , which are increasingly being reared for release by citizens and commercial breeders. Since late-summer monarchs should be as migration capable as possible for surviving the arduous long-distance migration, we evaluated four migration-relevant traits across two groups of captive-reared monarchs ( n = 41 and 42) and one group of wild-caught migrants ( n = 41). Monarchs (descendants of wild individuals) were reared from eggs to adulthood either in a warm indoor room next to a window, or in an incubator that mimicked late-summer conditions. Using an apparatus consisting of a perch mounted to an electronic force gauge, we assessed 'grip strength' of all groups, then used image analysis to measure forewing size, pigmentation and elongation. In three of the four traits, reared monarchs underperformed compared to wild ones, even those reared under conditions that should have produced migration-ready individuals. The average strength of reared monarchs combined was 56% less than the wild group, even when accounting for size. Their orange wing colour was paler (an indicator of poor condition and flight ability) and their forewings were less elongated (elongation is associated with migration propensity) than wild monarchs. The reason(s) behind these effects is unknown but could stem from the frequent disturbance and/or handling of reared monarchs, or the fact that rearing removes the element of natural selection from all stages. Regardless, these results explain prior tagging studies that showed reared monarchs have lower migratory success compared to wild.

Developed countries around the world are criss-crossed with vast networks of roadways. Conservationists have recently focused attention on roadsides as possible locations for establishing pollinator habitat, with the monarch butterfly ( Danaus plexippus ) featuring prominently in such discussions. However, roadsides are inherently loud, which could negatively affect developing larvae. We conducted a series of experiments testing if simulated highway noise stresses monarch larvae, which we gauged by non-destructive monitoring of heart rates. In two replicated experiments, larvae exposed for 2 h experienced a significant increase in heart rate (16 and 17% elevation), indicating they perceive traffic noise as a stressor. Meanwhile, experiments exposing larvae for either 7 or 12 days to continuous traffic noise both showed no heart rate elevation at the end of larval development, suggesting chronic noise exposure leads to habituation or desensitization. Habituation to stress as larvae may impair reactions to real-world stressors as adults, which could be problematic for a butterfly that undertakes an annual two-month migration that is fraught with dangers. More generally, these results could have far-reaching implications for the billions of insects worldwide that develop near roadways, and argue that further study is needed before promoting roadside habitat for butterfly conservation., (© 2018 The Author(s).)

Long-distance migration can lower parasite prevalence if strenuous journeys remove infected animals from wild populations. We examined wild monarch butterflies (Danaus plexippus) to investigate the potential costs of the protozoan Ophryocystis elektroscirrha on migratory success. We collected monarchs from two wintering sites in central Mexico to compare infection status with hydrogen isotope (δ2H) measurements as an indicator of latitude of origin at the start of fall migration. On average, uninfected monarchs had lower δ2H values than parasitized butterflies, indicating that uninfected butterflies originated from more northerly latitudes and travelled farther distances to reach Mexico. Within the infected class, monarchs with higher quantitative spore loads originated from more southerly latitudes, indicating that heavily infected monarchs originating from farther north are less likely to reach Mexico. We ruled out the alternative explanation that lower latitudes give rise to more infected monarchs prior to the onset of migration using citizen science data to examine regional differences in parasite prevalence during the summer breeding season. We also found a positive association between monarch wing area and estimated distance flown. Collectively, these results emphasize that seasonal migrations can help lower infection levels in wild animal populations. Our findings, combined with recent declines in the numbers of migratory monarchs wintering in Mexico and observations of sedentary (winter breeding) monarch populations in the southern U.S., suggest that shifts from migratory to sedentary behavior will likely lead to greater infection prevalence for North American monarchs.

In animals with complex life cycles, all resources needed to form adult tissues are procured at the larval stage. For butterflies, the proper development of wings involves synthesizing tissue during metamorphosis based on the raw materials obtained by larvae. Similarly, manufacture of pigment for wing scales also requires resources acquired by larvae. We conducted an experiment to test the effects of food deprivation in the larval stage on multiple measures of adult wing morphology and coloration of monarch butterflies (Danaus plexippus), a species in which long-distance migration makes flight efficiency critical. In a captive setting, we restricted food (milkweed) from late-stage larvae for either 24 hrs or 48 hrs, then after metamorphosis we used image analysis methods to measure forewing surface area and elongation (length/width), which are both important for migration. We also measured the brightness of orange pigment and the intensity of black on the wing. There were correlations between several wing features, including an unexpected association between wing elongation and melanism, which will require further study to fully understand. The clearest effect of food restriction was a reduction in adult wing size in the high stress group (by approximately 2%). Patterns observed for other wing traits were ambiguous: monarchs in the low stress group (but not the high) had less elongated and paler orange pigmentation. There was no effect on wing melanism. Although some patterns obtained in this study were unclear, our results concerning wing size have direct bearing on the monarch migration. We show that if milkweed is limited for monarch larvae, their wings become stunted, which could ultimately result in lower migration success.

The distinctive orange and black wings of monarchs (Danaus plexippus) have long been known to advertise their bitter taste and toxicity to potential predators. Recent work also showed that both the orange and black coloration of this species can vary in response to individual-level and environmental factors. Here we examine the relationship between wing color and flight performance in captive-reared monarchs using a tethered flight mill apparatus to quantify butterfly flight speed, duration and distance. In three different experiments (totaling 121 individuals) we used image analysis to measure body size and four wing traits among newly-emerged butterflies prior to flight trials: wing area, aspect ratio (length/width), melanism, and orange hue. Results showed that monarchs with darker orange (approaching red) wings flew longer distances than those with lighter orange wings in analyses that controlled for sex and other morphometric traits. This finding is consistent with past work showing that among wild monarchs, those sampled during the fall migration are darker in hue (redder) than non-migratory monarchs. Together, these results suggest that pigment deposition onto wing scales during metamorphosis could be linked with traits that influence flight, such as thorax muscle size, energy storage or metabolism. Our results reinforce an association between wing color and flight performance in insects that is suggested by past studies of wing melansim and seasonal polyphenism, and provide an important starting point for work focused on mechanistic links between insect movement and color.

The demands of long-distance flight represent an important evolutionary force operating on the traits of migratory species. Monarchs are widespread butterflies known for their annual migrations in North America. We examined divergence in wing morphology among migratory monarchs from eastern and western N. America, and nonmigratory monarchs in S. Florida, Puerto Rico, Costa Rica, and Hawaii. For the three N. American populations, we also examined monarchs reared in four common environment experiments. We used image analysis to measure multiple traits including forewing area and aspect ratio; for laboratory-reared monarchs we also quantified body area and wing loading. Results showed wild monarchs from all nonmigratory populations were smaller than those from migratory populations. Wild and captive-reared eastern monarchs had the largest and most elongated forewings, whereas monarchs from Puerto Rico and Costa Rica had the smallest and roundest forewings. Eastern monarchs also had the largest bodies and high measures of wing loading, whereas western and S. Florida monarchs had less elongated forewings and smaller bodies. Among captive-reared butterflies, family-level effects provided evidence that genetic factors contributed to variation in wing traits. Collectively, these results support evolutionary responses to long-distance flight in monarchs, with implications for the conservation of phenotypically distinct wild populations.

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.

Abstract: Pollinator declines have motivated efforts to plant nectar and host plants for butterflies and other pollinators, but whether gardens promote pollinator conservation requires further investigation. We established garden plots to determine whether plant type (native vs. exotic) and weed maintenance (low or high) influence adult butterfly abundance and species richness, and the abundance and survival of immature stages of four species (monarch, queen, black swallowtail, and gulf fritillary). To assess how predator activity and mortality of immature stages might differ inside gardens compared to other suitable habitat patches, we compared caterpillar survival on sentinel host plants placed within and outside of plots. The abundance and species richness of adult butterflies (all taxa) increased with the number of flowering plants per plot, but did not depend on plant type or weeding treatment. Exotic plots had greater adult monarch and gulf fritillary abundances, greater monarch and queen egg counts, and lower black swallowtail caterpillar counts relative to native plots. The survival of immature stages (egg to instar 5) ranged from 2 to 13% and was similar across plots, except for gulf fritillary larvae, which had higher survival in exotic plots. Monarch caterpillar mortality risk was higher on sentinel plants placed inside plots, relative to those outside of plots. This study suggests that garden plots attracted a diversity of adult butterflies and supported the reproduction of focal species. Given lower immature monarch survival within versus outside of plots, further work is needed to examine natural enemy pressure within pollinator gardens. [ABSTRACT FROM AUTHOR]

Declines in overwintering colonies of monarch butterflies (Danaus plexxipus) in Mexico raise questions about other life cycle phases, such as spring migration, where monarchs recolonize their breeding range in the United States and Canada with sequential generations. We used data from a longterm citizen science program, "Journey North" (now with 18 yr), to identify possible changes to the recolonization. This program asks people to report the date and location when they see the first adult monarch annually, and this database now contains >11,000 records. We examined sighting dates and migration range size, the latter based on the number of 2-degree latitude-longitude grid squares with monarch sightings, to look for evidence of change in either of these two parameters over the 18yr. Our analyses used regression models that accounted for increasing volunteer participation over the years. We found monarchs are being sighted later at a rate of 1 d later every 4 yr. This does not appear to be related to later emergence of milkweed, based on examination of milkweed reports. Later sightings could be interpreted as a sign of reductions in monarch abundance (it takes longer to see the first monarch of the year). We also found a potential decline in the geographic range of the initial spring migration wave (a decline of 9% over 18 yr). However we detected no change in the continental area encompassed at the end of recolonization, indicating monarchs are still successfully f illing their traditional breeding range in eastern North America. [ABSTRACT FROM AUTHOR]

. 1. The status of the eastern North American monarch butterfly population is a highly sensitive issue, given that winter and breeding habitats are being lost at an alarming rate each year, and because of this, most believe the population to be declining, although there has been little empirical data to support this idea. In a recent forum article of this journal, Brower et al. (2011) report a statistically significant decline in winter colony size over a 17-year period and suggest that this is the first sign of impending collapse. 2. I conducted an analysis of numbers of migrating monarchs from two fall monitoring stations in the United States (Cape May, NJ and Peninsula Point, MI), which span 15 and 19 years, respectively, and at both locations there was no significant linear trend in average monarch numbers counted over time, which is in marked contrast to the conclusion drawn by Brower et al. (2011). 3. Although I identify several possible reasons for the discrepancy between the fall census counts and size of the overwintering areas, these differing patterns argue for a more balanced perspective regarding the status of this population, and certainly for considering more than one phase of the life cycle. Even though it is difficult to imagine how monarchs will fare in the future with so many threats to their population, the data presented here suggest that the population remains stable for now, probably because of the high fecundity of the species and its ability to rebound from small winter numbers. [ABSTRACT FROM AUTHOR]

Heart rates of insects reflect the current level of activity and stress individuals are experiencing, and therefore this information can be useful from a research perspective. In Lepidoptera, as with most insects, the 'heart' consists of a transverse longitudinal tube that runs along the abdomen, just under the abdominal tergites, which pumps or contracts rhythmically to distribute hemolymph. Here, we describe a simple method we developed to observe heart contractions in adult monarch butterflies, Danaus plexippus. The procedure involves stabilizing the live specimen in a pre-prepared plastic bag, while positioning the abdomen in such a way so that the beating heart can be seen (with magnification) through the intersegmental membranes. With this procedure, no harm comes to the specimen, making the technique useful in both lab and field studies. The technique also requires little equipment, except for a dissecting microscope (or other magnifier). Moreover, the procedure should be useful for monitoring other similarly-sized Lepidoptera, and we confirmed this with a Papilio glaucus specimen. Using this method on 10 male and 10 female monarch specimens from captive-reared stock, we found the resting heart rate was 63 beats/min on average (range: 35-86). This information will be useful for comparative purposes, or as a reference point for future studies of monarch physiology. [ABSTRACT FROM AUTHOR]

Abstract: [1.] As ectotherms, insects often experience varying temperatures throughout their life cycle, and some respond by becoming more or less melanistic (dark coloring) during development to increase or decrease thermal energy absorption as larvae or adults. [2.] Monarch butterflies (Danaus plexippus) breed in temperate and tropical environments worldwide and are exposed to different average and extreme temperatures in different parts of their geographic range. In this study, we compared variation in thermally induced melanism among monarch butterflies from eastern and western North America and from South Florida. [3.] We raised the progeny of wild-captured adult butterflies from these populations in a common garden experiment, rearing individuals in cold (19°C), moderate (26°C), and hot (32°C) temperatures to examine population variation in larval and adult pigmentation. [4.] Across all populations, monarch larvae developed the darkest coloration in the cold treatment and were lightest when reared in hot temperatures. Similar results were observed for measures of adult wing melanism, with the exception of adult females, which developed darker colored wings in warmer temperatures. [5.] Significant population-level differences in average measures of melanism among larvae and adult butterflies were observed. Larvae from the eastern population became substantially darker in colder temperatures than S. Florida or western larvae. Western larvae were lightest overall, which might be adaptive to high temperatures experienced throughout portions of their summer breeding range. S. Florida larvae showed a lower response to cold temperatures relative to monarchs from either migratory population. [6.] Population level differences were also observed for thermal responses in wing melanism, particularly among adult females. Moreover, we found significant family level effects for each measure of larval and adult melanism, pointing to a genetic basis or strong maternal effects influencing these traits in monarch butterflies. [Copyright &y& Elsevier]

Since 1996, the numbers of migrating monarch butterflies stopping over at Peninsula Point, Michigan, have been monitored by volunteers during the fall migration with standardized daily counts. In this study, we describe this project and examine: 1) general patterns of migration and stopover of monarchs at this site, and 2) how environmental conditions influence monarch stopover frequency. We tested for yearly, seasonal, and diurnal variation in monarch counts within each season. We further combined these data with basic weather information recorded at the time of each count to explore the effects of wind direction and speed, temperature, and cloud cover on monarch stopover abundance. A total of 22,539 monarchs was counted over 7 yr, with yearly totals ranging from 757 in 1998 to 6,638 in 1997. Over the 7-yr period, an average of 29 monarchs was recorded per count at Peninsula Point. Interestingly, in the migration season immediately following a major population decline at overwintering sites in Mexico, the total number of monarchs counted at Peninsula Point was not significantly different from long-term average counts. The timing of the peak of migration was not consistent from year to year, and there were few consistent temporal trends within seasons. More monarchs were counted with walking transects during the day than with a roost count in the early morning. Furthermore, more monarchs were counted earlier in the season than later. Of the environmental variables we examined, wind direction had a significant influence on the number of monarchs recorded on each count with higher counts during north winds. Cloud cover also influenced monarch counts, so that the number of monarchs observed increased with temperature and decreased with cloud cover. Based on the large numbers of monarchs that stop there each fall, we suggest that Peninsula Point represents an important monarch stopover site, and thus has the potential to increase our knowledge of monarch migration and stopover ecology greatly. [ABSTRACT FROM AUTHOR]