Adult Monarchs Research Articles

Adult monarch butterflies show high tolerance to neonicotinoid insecticides

Abstract Numerous studies have documented the negative effects of neonicotinoids on bees; it remains crucial to examine how neonicotinoids affect other non‐target nectar‐feeding insects, such as the monarch butterfly, Danaus plexippus. Wildflowers growing near agricultural areas can be contaminated with neonicotinoids that affect survival or cause sublethal changes to behaviours of nectar‐feeding insects. Nectar residues of imidacloprid and clothianidin found in milkweeds and wildflowers adjacent to agricultural field range from 0 to 72.8 ng/mL. At field‐relevant doses, two neonicotinoids (imidacloprid and clothianidin) were studied for their effects on adult monarch survival, reproduction, flight and behaviour. First, we fed adult monarchs artificial nectar solutions ranging from 15 to 386 ng/mL of imidacloprid and 19 to 531 ng/mL of clothianidin. Neonicotinoid ingestion slightly reduced monarch reproduction but had no significant effects on survival, weight change, or activity levels. Second, we fed monarchs higher clothianidin doses (909 and 4030 ng/mL), that exceed field‐relevant levels by 22 and 99 times. These higher doses reduced monarch nectar consumption, survival, flight performance and reaction time in response to a drop test. Results show that adult monarchs tolerate field‐relevant doses as high as 54 ng/mL for imidacloprid and 75 ng/mL for clothianidin, with minimal lethal or sub‐lethal effects until much higher doses are supplied. We conclude that adult monarchs are more tolerant of ingested clothianidin and imidacloprid than indicated by previous research.

Isotopic (δ2H and δ13C) tracing the provenance and fate of individual fatty acids fueling migrating animals: A case study of the monarch butterfly (Danaus plexippus)

IntroductionAmong long-distance migratory insects, the monarch butterfly (Danaus plexippus) is one of the most iconic, whose journey is fueled by nectar from flowering plants along the migratory route which may involve up to 3,500 km. Understanding how and where monarchs obtain their dietary resources to fuel migratory flight and ensure overwintering stores would provide new insights into the migratory strategy of this species and subsequently help focus conservation efforts.MethodsThis pilot study was designed as a first attempt to assess the composition, dynamics, and isotopic (δ2H, δ13C) composition of essential and non-essential fatty acids (FA) acquired or manufactured de novo from larval host milkweed (Asclepias spp.) by monarch butterflies and from adult emergence to overwintering.ResultsData from controlled laboratory isotopic tracer tests suggested that adult monarchs convert their dietary energy mainly into 16:0 and 18:1 fatty acids and store them as neutral lipids in their abdomen. FA isotopic composition reflects not only dietary sources but also subsequent isotopic fractionation from metabolism. On the other hand, δ2H values of essential omega-3 fatty acid alpha-linolenic acid (ALA) correlated with δ2HWing, as markers of an individual’s geographic origin and indicated the importance of larval diet. Additionally, in wild-type females, high isotopic fractionation in δ13CALA between neutral and polar lipids might indicate increased bioconversion activity during gravidity. Finally, δ2HLIN showed positive H isotope fractionation from larval dietary sources, indicating that catabolic processes were involved in their manufacture. Furthermore, δ2HLIN showed a negative correlation with δ2HWing values, which could potentially be useful when investigating individual life-history traits, such as migratory catabolic efforts or periods of fasting.DiscussionThis interpretation was supported by significant larger variations in δ2HLIN and δ2HLIN overwintering monarchs compared to other FA. Altogether, our results provide the first evidence that the H isotopic analysis of individual fatty acids in migrating and overwintering monarchs can be used to infer the nutritional history of individuals including the provenance of nectaring sites used to fuel key life history events.

First Population Study on Winter Breeding Monarch Butterflies, Danaus plexippus (Lepidoptera: Nymphalidae) in the Urban South Bay of San Francisco, California.

Simple SummaryPopulations of western monarch butterflies, which formerly populated coastal overwintering sites in California in numbers exceeding one million, dwindled to less than 2000 in 2020/21. In the same winter, breeding populations of monarchs occurred commonly in the San Francisco Bay urban area for the first time. The urgent conservation need to understand this possible shift in overwintering strategy prompted this first study of the viability and ecology of monarch breeding populations in the South Bay urban area of San Francisco during January–June 2021. Adult monarchs along with eggs and larvae were common during winter and most of spring, utilizing non-native ornamental milkweed and multiple nectar sources from native and ornamental plants. Evidence from weekly counts and tagged butterflies indicated increased dispersal to the north and east during late April-mid-May, possibly representing spring migration to the Pacific Northwest and eastern California. Infection of reared adult monarchs with a protozoan parasite was high. Winter breeding of monarchs in the San Francisco urban area as an alternative and sustainable overwintering strategy for the western US population will likely depend on the continued presence of ornamental milkweeds, sustainable co-existence of monarchs and protozoan parasites, and successful migration to the greater western US during spring.The western North American monarch butterfly population assessed by counts of non-reproductive overwintering butterflies at coastal sites in California declined to less than 2000 in 2020/21. Simultaneously, reports of reproductive monarchs increased in San Francisco urban areas, perhaps representing a shift in overwintering strategy. To better understand monarch winter breeding in the Bay area, we studied adult and immature populations in Santa Clara County during January–June 2021. Adult monarchs were common with numbers ranging from 0.23–1.54/min during ~30 min weekly surveys at one site, with lowest numbers late April to mid-May. Eggs and larvae, primarily on ornamental milkweeds, were found on nearly all survey dates with lowest numbers mid-late April to mid-May. Levels of infection of adults by the parasite Ophryocystis elektroscirrha were consistently high during the study (69.3–77.5%). From 499 monarchs tagged post-eclosion, recovery rates of 19.2–23.6% occurred from releases in January-February and May-June but only 11.9–13.0% from March-April releases. Although distances were small, butterflies tagged in April were recovered from greater distances than other months. Tagged monarchs flew primarily north or east. There were reduced numbers of adult monarchs during late April-mid-May with some evidence of northerly and easterly emigration at the same time from tagged butterflies, suggesting some movement out of the South Bay area, perhaps representing spring migration. We conclude that monarchs can successfully breed and maintain populations on ornamental milkweeds during winter at urban sites in the South Bay of San Francisco and may still migrate during spring to remain part of the wider western population.

The puzzle of monarch butterflies (Danaus plexippus) and their association with plants containing pyrrolizidine alkaloids

1. The chemical defences of monarch butterflies involve two kinds of secondary plant metabolites: cardiac glycosides (CGs) that are obtained from larval hostplants, and pyrrolizidine alkaloids (PAs) that are gathered by adults usually independent from feeding behaviour. While monarchs and CGs have received great attention, monarchs and PAs appear largely ignored in science as well as by the public.2. We review and discuss PA‐pharmacophagy specifically for Danaus plexippus and its closest relatives and aim to encourage research and future consideration of this peculiar type of insect‐plant relationship, which represents an intrinsic but surprisingly understudied aspect of the lives of adult monarchs.

Evaluating Cardiac Reactions of Monarch Butterflies to Human Handling across Three Life Stages

Humans have an especially close connection with the monarch butterfly, Danaus plexippus, and this connection often involves direct handling of larvae, pupae or adults. Monarchs are frequently handled in citizen science programs, education and outreach activities, classroom demonstrations, and when rearing larvae in captivity. A pertinent question is whether monarchs perceive this human handling in the same way as being attacked by a predator, i.e. does it trigger a physiological fight-or-flight reaction? Here, I investigated if brief periods of handling monarch larvae, pupae, or adults elevates their heart rate (a central component of the insect fight-or-flight response), using non-destructive methods: either microscopy or an electronic heart monitor designed for invertebrates. Results are presented from separate investigations on each life stage. Three minutes of gentle handling caused a significant elevation (20%) in heart rates of 30 fifth-instar larvae. Mild physical disturbance for 20 seconds to a collection of 57 pupae elicited brief bursts of heart contractions (∼88 beats/min), a rate which is three times higher than typical contraction periods at this stage. Adult monarchs captured during fall migration (n=72) showed minor heart rate elevations (6%) after 3 minutes of gentle handling for tagging purposes. The differences in reactions across life stages found here is helpful for elucidating the relative sensitivity of each stage to handling. More importantly, these results elucidate the unseen physiological impact of direct human contact on developing monarchs (larvae and pupae); elevation in heart rate is an integral part of the fight-or-flight reaction in animals and a universal sign that an animal perceives an imminent threat. In this sense then, developing monarchs appear to perceive human handling as a threat, which may alter our (human) perception of this practice.

Monarch butterflies reared under autumn‐like conditions have more efficient flight and lower post‐flight metabolism

1. Many migratory animals undergo physiological and behavioural changes to prepare for and sustain long‐distance movements. Because insect migrations are common and diverse, studies that examine how migratory insects meet the energetic demands of long‐distance movements are badly needed.2. Monarch butterflies (Danaus plexippus) migrate up to 4000 km annually from eastern North America to wintering sites in central Mexico. Autumn generation monarchs undergo physiological and behavioural changes in response to environmental cues to initiate migration. In particular, exposure to cooler temperatures and shorter day lengths in early autumn causes monarchs to enter the hormonally induced state of reproductive diapause.3. This study examined differences in flight‐associated metabolic rate (MR) and flight performance metrics for monarchs experimentally reared under autumn‐like conditions (typically experienced before the southward migration) relative to monarchs reared under summer‐like conditions.4. Adult monarchs reared under autumn‐like conditions showed lower post‐flight MRs, greater flight efficiency, and lower measures of reproductive activity relative to monarchs reared under summer‐like conditions. Increases in post‐flight metabolism were associated with monarch body weight, age, and flight velocity.5. These findings suggest that a trans‐generational shift in flight energetics is an important component of the monarch's complex migratory syndrome, and that physiological changes that accompany reproductive diapause facilitate energy conservation during flight.

Incorporating field behaviors into monarch surveys to promote informed conservation actions

Monarch butterflies (Danaus plexippus) are an iconic species across North America, but ongoing winter population declines have made them a species of conservation concern. Incorporating behaviors into monitoring could connect landscape and behavioral ecology to benefit monarch conservation. Yet, behavioral field observations for breeding adult monarchs are seldom the focus of research, despite their potential benefits to targeted conservation. Our objectives were to quantify monarch activity budgets and compare behaviors of individuals at different sites with varying local vegetation and landscape characteristics in the Northern Great Plains. We opportunistically made 15-minute field observation surveys and recorded behaviors including basking, flying, mating, nectaring, ovipositing, and resting of wild monarchs. We collected plant community data including flowering ramet density and plant species composition. Over two years, we made behavioral observations on 51 monarchs. Collectively, monarchs spent most of their time flying and nectaring during observations. Contrary to our hypothesis, ovipositing and nectaring did not peak at sites with the highest milkweed (Asclepias spp.) cover. Instead, we observed more nectaring as milkweed decreased and plant species diversity increased. Individually, the proportion of time spent ovipositing and the abundance of adults increased at sites with both milkweed cover and higher plant species diversity. Our findings suggest we can improve conservation efforts for monarchs by promoting nectar sources and milkweed across landscapes, providing resources for all life stages. Moreover, continuing behavioral observations to understand how monarchs utilize sites with different vegetation and landscape characteristics can provide complementary information to abundance surveys to better inform conservation actions.

Balancing sampling intensity against spatial coverage for a community science monitoring programme

Abstract Community science is an increasingly integral part of biodiversity research and monitoring, often achieving broad spatial and temporal coverage but lower sampling intensity than studies conducted by professional scientists. When designing a community‐science monitoring programme, careful assessment of sampling designs that could be both feasible and successful at meeting programme goals is essential. Monarch butterflies (Danaus plexippus) are the focus of several successful community‐science projects in the U.S., but broader coverage is needed to monitor breeding areas and explain population declines observed in overwintering areas. The U.S. Monarch Conservation Science Partnership's Integrated Monarch Monitoring Program (IMMP) will representatively monitor monarchs and milkweed across North America. We performed a simulation‐based power analysis to predict trade‐offs between sampling breadth (number of sites and years) and sampling intensity (number of visits or subplots per site and year) for the IMMP. We evaluated whether each sampling design would produce sufficient statistical power to detect population trends and differences among land‐use sectors in densities of milkweed, monarch eggs, and adult monarchs. Sampling breadth had much stronger effects than sampling intensity on statistical power for all three monitoring targets. Depending on land‐use sector, monitoring 400–800 sites over 10–15 years would detect trends in densities of milkweed and adult monarchs, but no scenarios were successful for monarch eggs. Sampling 400–800 sites would also detect small (for adult monarchs) to large (for milkweed) differences among land‐use sectors in density of all three monitoring targets within the first 2–5 years. As more data become available from the IMMP, the sampling goals can be updated. Synthesis and applications. Careful sample design is an essential step in developing a successful monitoring programme. For monarchs and milkweed, we found that sampling breadth (number of sites and years) had a much stronger effect on statistical power than sampling intensity (number of visits or subsamples per site), suggesting field protocols could be tailored to maximize recruitment and retention of volunteers by minimizing the effort required to monitor each site. Many long‐term monitoring programmes might similarly benefit from evaluating the statistical trade‐offs between sampling breadth and intensity in their sampling designs.

A Neonicotinoid Insecticide at a Rate Found in Nectar Reduces Longevity but Not Oogenesis in Monarch Butterflies, Danaus plexippus (L.). (Lepidoptera: Nymphalidae).

The monarch butterfly in North America has suffered a serious population decline since the mid-1990s. The introduction and widespread use of neonicotinoid insecticides during the same time period has been suggested as a potential driver of this decline but no studies have looked at the impact of these insecticides on adult monarchs. A brief laboratory study assessed the impact of Imidacloprid, the most commonly used neonicotinoid, on western monarch butterfly longevity and oogenesis. Imidacloprid at 23.5 ppb, a field-realistic rate reported from wild nectar and pollen, was fed ad libitum to newly-eclosed monarchs in a sugar-based diet for 22 days. Treated monarchs showed reduced longevity, suffering 78.8% mortality by day 22, compared to 20% in untreated monarchs. Prior to death, butterflies exhibited signs of poisoning including uncoordinated flapping of wings and uncontrolled vibrating of wings and body. Imidacloprid did not reduce egg production. Shortened adult longevity has serious consequences for monarch population development, migration and overwintering. The potential widespread impact of imidacloprid-contaminated crop and wild flower nectar, may be a significant driver of monarch population decline. More research on the impact of neonicotinoid insecticides on the monarch and other butterflies should be viewed as a serious priority.

Mowing Frequency Influences Number of Flowering Stems but not Population Age Structure of Asclepias viridis, an Important Monarch Host Plant

Milkweeds (Asclepias spp.) are host plants for monarch butterflies (Danaus plexippus), which are currently being considered for listing under the Endangered Species Act. Previous research has documented declines in milkweed availability in the Midwest, but less research has focused on the southern Great Plains. Furthermore, the influence of management practices on the structure of milkweed populations has not been evaluated. We assess the age structure of Asclepias viridis, an important milkweed species for monarchs in the southern Great Plains, in two land uses that differ in the frequency and timing of mowing. Plant age may influence resource availability for larval and adult monarchs by affecting foliar biomass and flower production, and flower production and subsequent seed production may influence the recruitment of new individuals to the population. Using herb-chronology, we found age structure did not differ between roadsides mowed several times during the growing season and grasslands mowed less frequently. However, we did find younger individuals in less frequently mowed grasslands had more stems and more flowering stems than those in more frequently mowed roadsides. Though plant age did not differ among the land uses and mowing regimes included in this study, our results suggest frequent mowing can negatively affect foliar and reproductive biomass of younger A. viridis plants. Limiting mowing during the early summer flowering season for A. viridis could support efforts to maximize host plant and nectar availability for monarchs and other pollinators.

Design Implications for Surveys to Monitor Monarch Butterfly Population Trends

The Iowa Multiple Species Inventory and Monitoring (MSIM) Program includes a protocol for monitoring butterfly density on conservation lands using transects. Most data are collected from sites chosen randomly; additional sites are chosen non-randomly for other reasons. We analyzed a 12-year dataset for monarchs to address how density (per 50 m2 transect section) responded to site selection (random versus non-random), latitude, and measures of the amount of milkweed and canopy cover on survey transects. Between 2006 and 2017, we conducted 2,328 surveys on 420 sites and detected a total of 2,757 adult monarchs. Monarch densities peaked in 2010 for random sites and 2012 for non-random sites, but densities were lowest in 2013 for both site types. The density of monarchs at non-random transects (0.047, 95% CI = 0.031, 0.623)) was higher than that at random transects (0.029, 95% CI = 0.019, 0.044) and the temporal trends of density for random and non-random sites were significantly different. Monarch density was positively correlated with UTM northing, suggesting that monarch density increases from south to north in Iowa. The percent of plots containing milkweed was positively correlated with monarch density whereas percent tree canopy cover was negatively correlated with monarch density. Our results show that non-random transects had more monarchs, which may be a concern when interpreting findings from some citizen science efforts that used non-probabilistic sampling designs. Collectively, the MSIM program data provide a comprehensive assessment of monarch densities statewide as well as the first empirically-derived density estimates for monarchs on the breeding grounds and may prove helpful when refining future monitoring efforts.

Sequential Partial Migration Across Monarch Generations in Michigan

Abstract Running title: Monarch alternative migration: We collected 434 adult monarchs and surveyed milkweeds for immature monarchs in southwest Michigan, USA in order to test the hypothesis that monarchs are temporally variable, sequential partial migrants rather than partial migrants that may be spatially separated. Adult size, wing wear, female egg counts, fat content and sequestered chemical defenses were measured in monarchs across an entire season from spring migrant arrival, through breeding, until autumn migrant departure. We predicted that a population characterized by starting from all migrants and no residents, through breeding residents, to all migrants and no residents should show life history measures consistent with changes in these proportions. Results show that female monarch spring migrants arrive with chorionated eggs and high wing loads in both intact and fat-extracted adults. Wing loads of both males and females decrease during the summer and increase again immediately before autumn departure, when the fat content of all adults increases markedly. The high fat content of spring arrivals is also characteristic of migrants. Cardenolide content of adults showed a similar pattern of high content in spring arrivals, a decrease in the summer and then an accumulation of cardenolide defenses in adults in late summer just before migratory departure. We conclude that these results are consistent with temporally variable, sequential partial migration in a short-lived insect that contrasts with spatially variable partial migration in longer-lived vertebrates.

Climate change and an invasive, tropical milkweed: an ecological trap for monarch butterflies.

While it is well established that climate change affects species distributions and abundances, the impacts of climate change on species interactions has not been extensively studied. This is particularly important for specialists whose interactions are tightly linked, such as between the monarch butterfly (Danaus plexippus) and the plant genus Asclepias, on which it depends. We used open-top chambers (OTCs) to increase temperatures in experimental plots and placed either nonnative Asclepias curassavica or native A.incarnata in each plot along with monarch larvae. We found, under current climatic conditions, adult monarchs had higher survival and mass when feeding on A.curassavica. However, under future conditions, monarchs fared much worse on A.curassavica. The decrease in adult survival and mass was associated with increasing cardenolide concentrations under warmer temperatures. Increased temperatures alone reduced monarch forewing length. Cardenolide concentrations in A.curassavica may have transitioned from beneficial to detrimental as temperature increased. Thus, the increasing cardenolide concentrations may have pushed the larvae over a tipping point into an ecological trap; whereby past environmental cues associated with increased fitness give misleading information. Given the ubiquity of specialist plant-herbivore interactions, the potential for such ecological traps to emerge as temperatures increase may have far-reaching consequences.

Local and cross‐seasonal associations of climate and land use with abundance of monarch butterflies Danaus plexippus

Quantifying how climate and land use factors drive population dynamics at regional scales is complex because it depends on the extent of spatial and temporal synchrony among local populations, and the integration of population processes throughout a species’ annual cycle. We modeled weekly, site‐specific summer abundance (1994–2013) of monarch butterflies Danaus plexippus at sites across Illinois, USA to assess relative associations of monarch abundance with climate and land use variables during the winter, spring, and summer stages of their annual cycle. We developed negative binomial regression models to estimate monarch abundance during recruitment in Illinois as a function of local climate, site‐specific crop cover, and county‐level herbicide (glyphosate) application. We also incorporated cross‐seasonal covariates, including annual abundance of wintering monarchs in Mexico and climate conditions during spring migration and breeding in Texas, USA. We provide the first empirical evidence of a negative association between county‐level glyphosate application and local abundance of adult monarchs, particularly in areas of concentrated agriculture. However, this association was only evident during the initial years of the adoption of herbicide‐resistant crops (1994–2003). We also found that wetter and, to a lesser degree, cooler springs in Texas were associated with higher summer abundances in Illinois, as were relatively cool local summer temperatures in Illinois. Site‐specific abundance of monarchs averaged approximately one fewer per site from 2004–2013 than during the previous decade, suggesting a recent decline in local abundance of monarch butterflies on their summer breeding grounds in Illinois. Our results demonstrate that seasonal climate and land use are associated with trends in adult monarch abundance, and our approach highlights the value of considering fine‐resolution temporal fluctuations in population‐level responses to environmental conditions when inferring the dynamics of migratory species.

Long-Term Trends in Eastern North American Monarch Butterflies: A Collection of Studies Focusing on Spring, Summer, and Fall Dynamics:

The monarch butterfly, Danaus plexippus (Nymphalidae, Danainae), is one of the most familiar and appreciated insects in North America and is an iconic symbol of insect conservation (Gustafsson et al. 2015); attention on this charismatic species has increased since declines of their Mexican overwintering colonies were reported (Brower et al. 2012). These declines are alarming, yet they may or may not be representative of the patterns for other phases of this species’ ontogeny or its phenology. The collection of articles in this special feature was produced by researchers associated with a variety of nation-wide citizen science projects that have been tracking monarchs for many years, with the overarching goal of identifying long-term trends in their abundance or distribution outside of the wintering period. The articles in this feature include: 1) a study that reports patterns of spring recolonization within the breeding range (Howard and Davis); 2) two articles that examined long-term changes in reproductive output (Stenoien et al., Nail et al.); 3) a paper that analyzes censuses of summer adult monarchs from two butterfly monitoring programs (Ries et al.); 4) two studies focusing on fall migration censuses (Badgett and Davis, Crewe and McCracken); and 5) a paper examining 18 yr of migration tagging and recovery data from one site (Steffy). The collective results of these studies should have a significant impact on our current understanding of eastern monarch population dynamics. For example, despite dramatic reductions in overwintering size, there has been no detectable change in the recolonization of the overall breeding range of monarchs in the past 18 yr (Howard and Davis), and at least one model suggests …

Trans-generational parasite protection associated with paternal diet.

Multiple generations of hosts are often exposed to the same pathogens, favouring the evolution of trans-generational defences. Because females have more opportunities to transfer protective molecules to offspring, many studies have focused on maternally derived protection. However, males of many species can transfer compounds along with sperm, including chemicals that could provide protection. Here, we assess maternally and paternally derived protection in a monarch butterfly-protozoan parasite system where parasite resistance is heavily influenced by secondary plant chemicals, known as cardenolides, present in the larval diet of milkweed plants. We reared monarch butterflies on medicinal and non-medicinal milkweed species and then measured resistance of their offspring to infection. We also measured cardenolide content in adult monarchs reared on the two species, and in the eggs that they produced. We found that offspring were more resistant to infection when their fathers were reared on medicinal milkweed, while maternal diet had less of an effect. We also found that eggs contained the highest levels of cardenolides when both parents were reared on the medicinal species. Moreover, females reared on non-medicinal milkweed produced eggs with significantly higher levels of cardenolides if they mated with males reared on the medicinal milkweed species. However, we found an equivocal relationship between the cardenolides present in eggs and parasite resistance in the offspring. Our results demonstrate that males reared on medicinal plants can transfer protection to their offspring, but the exact mechanism remains unresolved. This suggests that paternal protection from parasitism might be important, particularly when there are environmental sources of parasite resistance and when males transfer spermatophores during mating.

Richard III’s Baby Teeth

Richard III’s Baby Teeth

Genetic Factors and Host Traits Predict Spore Morphology for a Butterfly Pathogen.

Monarch butterflies (Danaus plexippus) throughout the world are commonly infected by the specialist pathogen Ophryocystis elektroscirrha (OE). This protozoan is transmitted when larvae ingest infectious stages (spores) scattered onto host plant leaves by infected adults. Parasites replicate internally during larval and pupal stages, and adult monarchs emerge covered with millions of dormant spores on the outsides of their bodies. Across multiple monarch populations, OE varies in prevalence and virulence. Here, we examined geographic and genetic variation in OE spore morphology using clonal parasite lineages derived from each of four host populations (eastern and western North America, South Florida and Hawaii). Spores were harvested from experimentally inoculated, captive-reared adult monarchs. Using light microscopy and digital image analysis, we measured the size, shape and color of 30 replicate spores per host. Analyses examined predictors of spore morphology, including parasite source population and clone, parasite load, and the following host traits: family line, sex, wing area, and wing color (orange and black pigmentation). Results showed significant differences in spore size and shape among parasite clones, suggesting genetic determinants of morphological variation. Spore size also increased with monarch wing size, and monarchs with larger and darker orange wings tended to have darker colored spores, consistent with the idea that parasite development depends on variation in host quality and resources. We found no evidence for effects of source population on variation in spore morphology. Collectively, these results provide support for heritable variation in spore morphology and a role for host traits in affecting parasite development.

Fire creates host plant patches for monarch butterflies

Monarch butterflies (Danaus plexippus) depend on the presence of host plants (Asclepias spp.) within their breeding range for reproduction. In the southern Great Plains, Asclepias viridis is a perennial that flowers in May and June, and starts to senesce by August. It is locally abundant and readily used by monarchs as a host plant. We evaluated the effects of summer prescribed fire on A. viridis and the use of A. viridis by monarch butterflies. Summer prescribed fire generated a newly emergent population of A. viridis that was absent in other areas. Pre-migrant monarch butterflies laid eggs on A. viridis in summer burned plots in late August and September, allowing adequate time for a new generation of adult monarchs to emerge and migrate south to their overwintering grounds. Thus, summer prescribed fire may provide host plant patches and/or corridors for pre-migrant monarchs during a time when host plant availability may be limited in other areas.

Monarch butterfly migration and parasite transmission in eastern North America

Seasonal migration occurs in many animal systems and is likely to influence interactions between animals and their parasites. Here, we focus on monarch butterflies (Danaus plexippus) and a protozoan parasite (Ophryocystis elektroscirrha) to investigate how host migration affects infectious disease processes. Previous work showed that parasite prevalence was lower among migratory than nonmigratory monarch populations; two explanations for this pattern are that (1) migration allows animals to periodically escape contaminated habitats (i.e., migratory escape), and (2) long‐distance migration weeds out infected animals (i.e., migratory culling). We combined field‐sampling and analysis of citizen science data to examine spatiotemporal trends of parasite prevalence and evaluate evidence for these two mechanisms. Analysis of within‐breeding‐season variation in eastern North America showed that parasite prevalence increased from early to late in the breeding season, consistent with the hypothesis of migratory escape. Prevalence was also positively related to monarch breeding activity, as indexed by larval density. Among adult monarchs captured at different points along the east coast fall migratory flyway, parasite prevalence declined as monarchs progressed southward, consistent with the hypothesis of migratory culling. Parasite prevalence was also lower among monarchs sampled at two overwintering sites in Mexico than among monarchs sampled during the summer breeding period. Collectively, these results indicate that seasonal migration can affect parasite transmission in wild animal populations, with implications for predicting disease risks for species with threatened migrations.