Migratory connectivity of the monarch butterfly (Danaus plexippus): patterns of spring re-colonization in eastern North America

Each year, millions of monarch butterflies (Danaus plexippus) migrate up to 3000 km from their overwintering grounds in central Mexico to breed in eastern North America. Malcolm et al. (1993) articulated two non-mutually exclusive hypotheses to explain how Monarchs re-colonize North America each spring. The 'successive brood' hypothesis proposes that monarchs migrate from Mexico to the Gulf Coast, lay eggs and die, leaving northern re-colonization of the breeding range to subsequent generations. The 'single sweep' hypothesis proposes that overwintering monarchs continue to migrate northward after arriving on the Gulf coast and may reach the northern portion of the breeding range, laying eggs along the way. To examine these hypotheses, we sampled monarchs throughout the northern breeding range and combined stable-hydrogen isotopes (δD) to estimate natal origin with wing wear scores to differentiate between individuals born in the current vs. previous year. Similar to Malcolm et al. (1993), we found that the majority of the northern breeding range was re-colonized by the first generation of monarchs (90%). We also estimated that a small number of individuals (10%) originated directly from Mexico and, therefore adopted a sweep strategy. Contrary to Malcolm et al. (1993), we found that 62% of monarchs sampled in the Great Lakes originated from the Central U.S., suggesting that this region is important for sustaining production in the northern breeding areas. Our results provide new evidence of re-colonization patterns in monarchs and contribute important information towards identifying productive breeding regions of this unique migratory insect.     

Linking the continental migratory cycle of the monarch butterfly to understand its population decline

Threats to several of the world's great animal migrations necessitate a research agenda focused on identifying drivers of their population dynamics. The monarch butterfly is an iconic species whose continental migratory population in eastern North America has been declining precipitously. Recent analyses have linked the monarch decline to reduced abundance of milkweed host plants in the USA caused by increased use of genetically modified herbicide‐resistant crops. To identify the most sensitive stages in the monarch's annual multi‐generational migration, and to test the milkweed limitation hypothesis, we analyzed 22 years of citizen science records from four monitoring programs across North America. We analyzed the relationships between butterfly population indices at successive stages of the annual migratory cycle to assess demographic connections and to address the roles of migrant population size versus temporal trends that reflect changes in habitat or resource quality. We find a sharp annual population decline in the first breeding generation in the southern USA, driven by the progressively smaller numbers of spring migrants from the overwintering grounds in Mexico. Monarch populations then build regionally during the summer generations. Contrary to the milkweed limitation hypothesis, we did not find statistically significant temporal trends in stage‐to‐stage population relationships in the mid‐western or northeastern USA. In contrast, there are statistically significant negative temporal trends at the overwintering grounds in Mexico, suggesting that monarch success during the fall migration and re‐establishment strongly contributes to the butterfly decline. Lack of milkweed, the only host plant for monarch butterfly caterpillars, is unlikely to be driving the monarch's population decline. Conservation efforts therefore require additional focus on the later phases in the monarch's annual migratory cycle. We hypothesize that lack of nectar sources, habitat fragmentation, continued degradation at the overwintering sites, or other threats to successful fall migration are critical limiting factors for declining monarchs.     

Lack of genetic differentiation between monarch butterflies with divergent migration destinations

Monarch butterflies are best known for their spectacular annual migration from eastern North America to Mexico. Monarchs also occur in the North American states west of the Rocky Mountains, from where they fly shorter distances to the California Coast. Whether eastern and western North American monarchs form one genetic population or are genetically differentiated remains hotly debated, and resolution of this debate is essential to understand monarch migration patterns and to protect this iconic insect species. We studied the genetic structure of North American migratory monarch populations, as well as nonmigratory populations in Hawaii and New Zealand. Our results show that eastern and western migratory monarchs form one admixed population and that monarchs from Hawaii and New Zealand have genetically diverged from North American butterflies. These findings suggest that eastern and western monarch butterflies maintain their divergent migrations despite genetic mixing. The finding that eastern and western monarchs form one genetic population also suggests that the conservation of overwintering sites in Mexico is crucial for the protection of monarchs in both eastern and western North America.     

Ecological niches in sequential generations of eastern North American monarch butterflies (Lepidoptera: Danaidae): the ecology of migration and likely climate change implications

Eastern North American monarch butterflies (Danaus plexippus L.) show a series of range shifts during their breeding season. Using ecological niche modeling, we studied the environmental context of these shifts by identifying the ecological conditions that monarchs use in successive summer months. Monarchs use a consistent ecological regimen through the summer, but these conditions contrast strikingly with those used during the winter. Hence, monarchs exhibit niche-following among sequential breeding generations but niche-switching between the breeding and overwintering stages of their annual cycle. We projected their breeding ecological niche onto monthly future climate scenarios, which indicated northward shifts, particularly at the northern extreme of their summer movements, over the next 50 yrs; if both monarchs and their milkweed host plants cannot track these changing climates, monarchs could lose distributional area during critical breeding months.     

Associations between host migration and the prevalence of a protozoan parasite in natural populations of adult monarch butterflies

1. Monarch butterflies Danaus plexippus (L.) (Lepidoptera: Nymphalidae) are susceptible to infection by the obligate protozoan parasite Ophryocystis elektroscirrha (McLaughlin and Myers) (Apicomplexa: Neogregarinida). Because monarchs form resident and migratory populations in different parts of the world, this host–parasite system provides the opportunity to examine how variation in parasite prevalence relates to host movement patterns. 2. Parasite prevalence was evaluated using 14 790 adult monarchs captured between 1968 and 1997. Comparison of three populations in North America indicated that parasite prevalence is associated negatively with host dispersal distances. A continuously breeding, nonmigratory population in southern Florida showed high prevalence (over 70% heavily infected). The western population migrates moderate distances to overwintering sites on the Pacific Coast and has intermediate prevalence (30% heavily infected). The eastern migratory population, which travels the longest distance to Mexican overwintering sites, has exhibited less than 8% infection throughout the past 30 years. 3. Variation in parasite loads within North American migratory populations was investigated to determine whether the prevalence of heavy infection and average parasite loads declined during migration or overwintering. Average parasite loads of summer‐breeding adults in western North America decreased with increasing distance from overwintering sites. This suggests that heavily infected monarchs are less likely to remigrate long distances in spring. No differences in the frequency of heavily infected adults were found among eastern or western North American monarchs throughout the overwintering period, however, suggesting that this parasite does not affect overwintering mortality. 4. Changes in the prevalence of monarchs with low parasite loads demonstrate that spore transfer occurs during migration and overwintering, possibly when adult butterflies contact each other as a result of their clustering behaviour. 5. This study of geographical and temporal variation in O. elektroscirrha among populations of D. plexippus demonstrates the potential role of seasonal migration in mediating interactions between hosts and parasites, and suggests several mechanisms through which migratory behaviour may influence parasite prevalence.     

Biological Observations of Monarch Butterfly Behavior at a Migratory Stopover Site: Results from a Long-term Tagging Study in Coastal South Carolina

Like most migratory species, monarch butterflies (Danaus plexippus) must stop frequently during their long southward migration to rest and refuel, and the places where they stop are important for the success of the migration. The behavior of monarch butterflies at migratory stopover sites has never been examined in detail. Here we present results of a long-term study of monarchs at one stopover site in coastal South Carolina where over 12,000 monarchs have been captured, measured and tagged (with numbered stickers to track recovery rates) over 13 years. Only 3 monarchs (0.023%) were recovered at the monarchs’ overwintering sites in Mexico, which is consistent with other tagging studies on the eastern coast. The migration season was longer at this site than at inland locations and monarchs continued to be captured in November and December, when most monarchs had already arrived at the overwintering areas in Mexico. In addition, there were 94 monarchs captured between Jan 1 and Mar 15, indicating that some monarchs overwinter at this site. Of all monarchs captured during the migration season, 80% were captured while nectaring and 10% while roosting. Others were basking, resting, flying and even mating. The sex ratio was male biased by three to one in all behavior categories except those captured mating. Roosting and nectaring monarchs had fresher wings than those in other behavior categories, suggesting that these are younger individuals. There were 13 observations of females ovipositing on non-native Asclepias curassavica during the fall months, which speaks to the potential for this plant to pull monarchs out of the migratory pool. Aside from these insights, this study also serves as an example of the potential that monarch tagging studies have to advance scientific understanding of monarch migration.     

Colonization and usage of eight milkweed (<Emphasis Type="Italic">Asclepias</Emphasis>) species by monarch butterflies and bees in urban garden settings

Planting milkweeds on public and private lands has emerged as a central conservation strategy for restoring declining North American migratory populations of the monarch butterfly (Danaus plexippus). Nearly all actionable science on this issue has focused on restoring common milkweed (Asclepias syriaca L.) in rural land types. The aim of this study was to develop recommendations for the best milkweeds for managed gardens intended to support both monarch butterflies and bees. Eight milkweed (Asclepias) species varying in height, form, and leaf shape were grown in a common-garden experiment at a public arboretum. We measured milkweed growth, tillering, and bloom periods, conducted bi-weekly counts of eggs and larvae to assess colonization by wild monarchs, and evaluated suitability for growth of monarch larvae. We also quantified bee visitation and compared the bee assemblages associated with six of the eight species, augmented with additional collections from other sites. Monarchs rapidly colonized the gardens, but did not equally use all of the milkweed species. More eggs and larvae were found on taller, broad-leaved milkweeds, but there was relatively little difference in larval performance, suggesting ovipositional preference for more apparent plants. Asclepias tuberosa and A. fascicularis attracted the greatest number of bees, whereas bee genus diversity was greatest on A. verticillata, A. fascicularis, and A. tuberosa. Milkweeds that do not spread extensively by tillering may be best suited for managed gardens. Combining milkweeds that are preferred by ovipositing monarchs with ones that are particularly attractive to bees may enhance conservation value of small urban gardens.     

Genomic evidence for gene flow between monarchs with divergent migratory phenotypes and flight performance

Monarch butterflies are known for their spectacular annual migration in eastern North America, with millions of monarchs flying up to 4,500 km to overwintering sites in central Mexico. Monarchs also live west of the Rocky Mountains, where they travel shorter distances to overwinter along the Pacific Coast. It is often assumed that eastern and western monarchs form distinct evolutionary units, but genomic studies to support this notion are lacking. We used a tethered flight mill to show that migratory eastern monarchs have greater flight performance than western monarchs, consistent with their greater migratory distances. However, analysing more than 20 million SNPs in 43 monarch genomes, we found no evidence for genomic differentiation between eastern and western monarchs. Genomic analysis also showed identical and low levels of genetic diversity, and demographic analyses indicated similar effective population sizes and ongoing gene flow between eastern and western monarchs. Gene expression analysis of a subset of candidate genes during active flight revealed differential gene expression related to nonmuscular motor activity. Our results demonstrate that eastern and western monarchs maintain migratory differences despite ongoing gene flow, and suggest that migratory differences between eastern and western monarchs are not driven by select major‐effects alleles. Instead, variation in migratory distance and destination may be driven by environmentally induced differential gene expression or by many alleles of small effect.     

Regional climate on the breeding grounds predicts variation in the natal origin of monarch butterflies overwintering in Mexico over 38 years

Addressing population declines of migratory insects requires linking populations across different portions of the annual cycle and understanding the effects of variation in weather and climate on productivity, recruitment, and patterns of long‐distance movement. We used stable H and C isotopes and geospatial modeling to estimate the natal origin of monarch butterflies (Danaus plexippus) in eastern North America using over 1000 monarchs collected over almost four decades at Mexican overwintering colonies. Multinomial regression was used to ascertain which climate‐related factors best‐predicted temporal variation in natal origin across six breeding regions. The region producing the largest proportion of overwintering monarchs was the US Midwest (mean annual proportion = 0.38; 95% CI: 0.36–0.41) followed by the north‐central (0.17; 0.14–0.18), northeast (0.15; 0.11–0.16), northwest (0.12; 0.12–0.16), southwest (0.11; 0.08–0.12), and southeast (0.08; 0.07–0.11) regions. There was no evidence of directional shifts in the relative contributions of different natal regions over time, which suggests these regions are comprising the same relative proportion of the overwintering population in recent years as in the mid‐1970s. Instead, interannual variation in the proportion of monarchs from each region covaried with climate, as measured by the Southern Oscillation Index and regional‐specific daily maximum temperature and precipitation, which together likely dictate larval development rates and food plant condition. Our results provide the first robust long‐term analysis of predictors of the natal origins of monarchs overwintering in Mexico. Conservation efforts on the breeding grounds focused on the Midwest region will likely have the greatest benefit to eastern North American migratory monarchs, but the population will likely remain sensitive to regional and stochastic weather patterns.     

Stable isotopes (δD and δ13C) are geographic indicators of natal origins of monarch butterflies in eastern North America

Wing membranes of laboratory and field-reared monarch butterflies (Danaus plexippus) were analyzed for their stable-hydrogen (δD) and carbon (δ¹³C) isotope ratios to determine whether this technique could be used to identify their natal origins. We hypothesized that the hydrogen isotopic composition of monarch butterfly wing keratin would reflect the hydrogen isotope patterns of rainfall in areas of natal origin where wings were formed. Monarchs were reared in the laboratory on milkweed plants (Asclepias sp.) grown with water of known deuterium content, and, with the assistance of volunteers, on native milkweeds throughout eastern North America. The results show that the stable hydrogen isotopic composition of monarch butterflies is highly correlated with the isotopic composition of the milkweed host plants, which in turn corresponds closely with the long-term geographic patterns of deuterium in rainfall. Stable-carbon isotope values in milkweed host plants were similarly correlated with those values in monarch butterflies and showed a general pattern of enrichment along a southwest to northeast gradient bisecting the Great Lakes. These findings indicate that natal origins of migratory and wintering monarchs in Mexico can be inferred from the combined δD and δ¹³C isotopic signatures in their wings. This relationship establishes that analysis of hydrogen and carbon isotopes can be used to answer questions concerning the biology of migratory monarch butterflies and provides a new approach to tracking similar migratory movements of other organisms. Received: 1 July 1998 / Accepted: 11 November 1998     

Sex ratios and Ophryocystis elektroscirrha infection levels of Danaus plexippus during spring migration through Oklahoma,USA

Monarch butterflies, Danaus plexippus L. (Lepidoptera: Nymphalidae), have a multiple brood migration in the spring as they move between their overwintering grounds and summer breeding grounds. In Oklahoma, USA, monarchs produce at least one generation in the spring, which develops and continues the northward migration, leaving Oklahoma without a breeding population during the hot summer months. Female monarchs leave the overwintering grounds prior to males, but it is not clear whether females re‐colonize areas along the migration route prior to, or at the same time as males. Male‐to‐female ratios are 1:1 at emergence, but studies have identified a male‐biased sex ratio in the field. Both males and females are susceptible to infection by the obligate protozoan parasite, Ophryocystis elektroscirrha McLaughlin & Myers (OE), which reduces flight abilities and life spans of infected individuals. We examine sex ratios during the spring migration through Oklahoma and whether sex ratios or OE infection estimates vary with capture technique (active or passive). Our data suggest populations are male‐biased during the 1st week of spring migration in Oklahoma, but shift to female‐biased by the 3rd week in both cool and warm springs. Therefore, males may leave southern areas prior to females or migrate longer distances per day. Active sampling (i.e., netting) did not bias sex compared to passive sampling (i.e., sticky traps). Significantly fewer OE‐carrying monarchs (with two or more spores) were captured via netting than by sticky traps which may be caused by sticky trap glue affecting tape sampling effectiveness, but there was no difference in the number of heavily infected individuals (more than 100 spores). Therefore, data from netted monarchs may underestimate OE infection rates within populations.     

Monarch butterflies cross the Appalachians from the west to recolonize the east coast of North America

Each spring, millions of monarch butterflies (Danaus plexippus) migrate from overwintering sites in Mexico to recolonize eastern North America. However, few monarchs are found along the east coast of the USA until mid-summer. Brower (Brower, L. P. 1996 J. Exp. Biol. 199, 93–103.) proposed that east coast recolonization is accomplished by individuals migrating from the west over the Appalachians, but to date no evidence exists to support this hypothesis. We used hydrogen (δD) and carbon (δ¹³C) stable isotope measurements to estimate natal origins of 90 monarchs sampled from 17 sites along the eastern United States coast. We found the majority of monarchs (88%) originated in the mid-west and Great Lakes regions, providing, to our knowledge, the first direct evidence that second generation monarchs born in June complete a (trans-) longitudinal migration across the Appalachian mountains. The remaining individuals (12%) originated from parents that migrated directly from the Gulf coast during early spring. Our results provide evidence of a west to east longitudinal migration and provide additional rationale for conserving east coast populations by identifying breeding sources.     

Postglacial northward expansion and genetic differentiation between migratory and sedentary populations of the broad‐tailed hummingbird (Selasphorus platycercus)

Unlike other migratory hummingbirds in North America, the broad‐tailed hummingbird (Selasphorus platycercus) exhibits both long‐distance migratory behaviour in the USA and sedentary behaviour in Mexico and Guatemala. We examined the evolution of migration linked to its northward expansion using a multiperspective approach. We analysed variation in morphology, mitochondrial and nuclear DNA, estimated migration rates between migratory and sedentary populations, compared divergence times with the occurrence of Quaternary climate events and constructed species distribution models to predict where migratory and sedentary populations resided during the Last Glacial Maximum (LGM) and Last Interglacial (LIG) events. Our results are consistent with a recent northward population expansion driven by migration from southern sedentary populations. Phylogeographical analyses and population genetics methods revealed that migratory populations in the USA and sedentary populations in Mexico of the platycercus subspecies form one admixed population, and that sedentary populations from southern Mexico and Guatemala (guatemalae) undertook independent evolutionary trajectories. Species distribution modelling revealed that the species is a niche tracker and that the climate conditions associated with modern obligate migrants in the USA were not present during the LIG, which provides indirect evidence for recent migratory behaviour in broad‐tailed hummingbirds on the temporal scale of glacial cycles. The finding that platycercus hummingbirds form one genetic population and that suitable habitat for migratory populations was observed in eastern Mexico during the LIG also suggests that the conservation of overwintering sites is crucial for obligate migratory populations currently facing climate change effects.     

Parasites hinder monarch butterfly flight: implications for disease spread in migratory hosts

Monarch butterflies (Danaus plexippus) are parasitized by the protozoan Ophryocystis elektroscirrha throughout their geographical range. Monarchs inhabiting seasonally fluctuating environments migrate annually, and parasite prevalence is lower among migratory relative to non‐migratory populations. One explanation for this pattern is that long‐distance migration weeds out infected animals, thus reducing parasite prevalence and transmission between generations. In this study we experimentally infected monarchs from a migratory population and recorded their long‐distance flight performance using a tethered flight mill. Results showed that parasitized butterflies exhibited shorter flight distances, slower flight speeds, and lost proportionately more body mass per km flown. Differences between parasitized and unparasitized monarchs were generally not explained by individual variation in wing size, shape, or wing loading, suggesting that poorer flight performance among parasitized hosts was not directly caused by morphological constraints. Effects of parasite infection on powered flight support a role for long‐distance migration in dramatically reducing parasite prevalence in this and other host–pathogen systems.     

Do Healthy Monarchs Migrate Farther? Tracking Natal Origins of Parasitized vs. Uninfected Monarch Butterflies Overwintering in Mexico

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 (δ ²H) measurements as an indicator of latitude of origin at the start of fall migration. On average, uninfected monarchs had lower δ ²H 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.     

Can roadside habitat lead monarchs on a route to recovery?

The Eastern North American monarch butterfly population has severely declined over the past decade. The decreasing availability of larval host plants (milkweeds) due to the use of herbicide-tolerant crops has been implicated in this decline. Roadsides could provide additional habitat for monarchs. In this study we document the occurrence of milkweed and monarchs on roadsides, and discuss whether roadsides are appropriate targets for monarch habitat restoration. We sampled roadside rights-of-way in the Upper Midwestern U.S. during the summer of 2015 to estimate the abundance, distribution, and diversity of milkweeds and the extent to which monarchs use these milkweeds. We then compared monarch densities in roadsides to other habitat types and modelled immature monarch densities based on several site characteristics. Our findings suggest that roadsides have conservation potential for monarchs, especially when other habitat is scarce and if wildlife-friendly management practices are enacted. Milkweeds were found on ~60% of roadside transects. Asclepias syriaca was the most common of the seven species encountered, occurring on 97% of transects with milkweed. Immature monarchs were observed in roadsides, but in lower densities than other habitats during the same time period. At lower milkweed densities, immature monarch density per unit area is positively correlated with milkweed density. However, milkweed density weakens as a predictor of immature monarch density over ~0.6 plants per m², possibly indicating a saturation effect.     

Fueling the fall migration of the monarch butterfly

Monarch butterflies in eastern North America accumulate lipidsduring their fall migration to central Mexico, and use themas their energy source during a 5 month overwintering period.When and where along their migratory journey the butterfliesaccumulate these lipids has implications for the importanceof fall nectar sources in North America. We analyzed the lipidcontent of 765 summer breeding and fall migrant monarch butterfliescollected at 1 nectaring site in central Virginia over 4 years(1998–2001), and compared them with 16 additional publishedand unpublished datasets from other sites, dating back to 1941.Virginia migrants store significantly more lipid than summerbutterflies, and show significant intraseason and between-yearvariation. None of the Virginia samples, and none of the historicalsamples, with one exception, had lipid levels comparable withthose found in migrants that had reached Texas and northernMexico. This evidence suggests that upon reaching Texas, thebutterflies undergo a behavioral shift and spend more time nectaring.The one exceptional sample led us to the discovery that monarchsthat form roosts along their migratory routes have higher lipidcontents than monarchs collected while nectaring at flowers.We propose that for much of their journey monarchs are opportunisticmigrants, and the variation within and between samples reflectsbutterflies' individual experiences. The stored lipids appearto be of less importance as fuel for the butterflies' migrationthan for their survival during their overwintering period, inpart because soaring on favorable winds reduces the energeticcost of flying. The conservation of nectar plants in Texas andnorthern Mexico is crucial to sustaining the monarch's migratoryspectacle, and nectar abundance throughout eastern North Americais also important. As generalists in their selection of nectarsources and nectaring habitats, monarchs are unlikely to beaffected by small changes in plant communities. Agriculturaltransformations of natural communities in the eastern UnitedStates and Great Plains, however, and especially the extensiveplanting of genetically modified herbicide-resistant soybeansand corn, may be changing the availability of nectar for monarchsand other pollinators. This new technology is eliminating virtuallyall forbs in and surrounding agricultural fields, includingthe monarch's larval hostplants (milkweeds) and native and nonnativenectar sources. To evaluate whether changes in nectar availabilityare altering the butterflies' ability to accumulate energy,we recommend that monarchs' lipid contents be assayed annuallyat sites throughout eastern North America.     

Patterns of parasitism in monarch butterflies during the breeding season in eastern North America

1. Migratory behaviour can result in reduced prevalence of pathogens in host populations. Two hypotheses have been proposed to explain this relationship: (i) ‘migratory escape’, where migrants benefit from escaping pathogen accumulation in contaminated environments; and (ii) ‘migratory culling’, where the selective removal of infected individuals occurs during migration. 2. In the host–parasite system between the monarch butterfly (Danaus plexippus Linn.) and its obligate protozoan parasite Ophryocystis elektroscirrha (OE), there is evidence to support both hypotheses, particularly during the monarchs' autumn migration. However, these processes can operate simultaneously and could vary throughout the monarchs' annual migratory cycle. Assessing the relative strength for each hypothesis has not previously been done. 3. To evaluate both hypotheses, parasite infection prevalence was examined in monarchs sampled in eastern North America during April–September, and stable isotopes (δ²H, δ¹³C) were used to estimate natal origin and infer migration distance. There was stronger support for the migratory escape hypothesis, wherein infection prevalence increased over the breeding season and was higher at southern latitudes, where the breeding season tends to be longer compared with northern latitudes. Little support was found for the migratory culling hypothesis, as infection prevalence was similar whether monarchs travelled shorter or longer distances. 4. These results suggest that migration allows individuals to escape parasites not only during the autumn, as shown in previous work, but during the monarchs' spring and summer movements when they recolonise the breeding range. These results imply a potential fitness advantage to monarchs that migrate further north to exploit parasite‐free habitats.     

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.     

Tracking climate impacts on the migratory monarch butterfly

Understanding the impacts of climate on migratory species is complicated by the fact that these species travel through several climates that may be changing in diverse ways throughout their complete migratory cycle. Most studies are not designed to tease out the direct and indirect effects of climate at various stages along the migration route. We assess the impacts of spring and summer climate conditions on breeding monarch butterflies, a species that completes its annual migration cycle over several generations. No single, broad‐scale climate metric can explain summer breeding phenology or the substantial year‐to‐year fluctuations observed in population abundances. As such, we built a Poisson regression model to help explain annual arrival times and abundances in the Midwestern United States. We incorporated the climate conditions experienced both during a spring migration/breeding phase in Texas as well as during subsequent arrival and breeding during the main recruitment period in Ohio. Using data from a state‐wide butterfly monitoring network in Ohio, our results suggest that climate acts in conflicting ways during the spring and summer seasons. High spring precipitation in Texas is associated with the largest annual population growth in Ohio and the earliest arrival to the summer breeding ground, as are intermediate spring temperatures in Texas. On the other hand, the timing of monarch arrivals to the summer breeding grounds is not affected by climate conditions within Ohio. Once in Ohio for summer breeding, precipitation has minimal impacts on overall abundances, whereas warmer summer temperatures are generally associated with the highest expected abundances, yet this effect is mitigated by the average seasonal temperature of each location in that the warmest sites receive no benefit of above average summer temperatures. Our results highlight the complex relationship between climate and performance for a migrating species and suggest that attempts to understand how monarchs will be affected by future climate conditions will be challenging.