Insect Migration Research Articles

Dragonfly: A Master of Migration

Insect migration is vital for ecosystems, with trillions of individuals redistributing biomass and nutrients annually. Dragonflies, particularly Pantala flavescens are notable migratory insects capable of traveling over 6,000 kilometres across generations. Their migrations are influenced by resource availability, breeding needs, environmental conditions, and competition avoidance. This behaviour includes both seasonal movements and sporadic flights triggered by temperature and wind cues. Despite their well-known life cycle from aquatic nymphs to aerial adults, many aspects of their migration remain under-researched. Dragonflies exhibit diverse flight mechanisms, employing four distinct modes: counter-stroking for hovering, phased-stroking for cruising synchronized stroking for acceleration, and gliding for energy conservation. Migratory species show specific wing adaptations for improved endurance, such as elongated forewings and larger hindwings, along with sensory enhancements like enlarged thoracic setae. Specialized flight muscles allow for independent wing control, essential for complex movements. Additionally, physiological adaptations like fat accumulation and rapid development support their migratory behaviour. Recent studies also highlight nocturnal migrations, revealing unique flight patterns and adaptations. Together, these traits underscore the evolutionary success of dragonflies in diverse ecosystems.

Energy Reserve Allocation in the Trade-Off between Migration and Reproduction in Fall Armyworm.

Striking a trade-off between migration and reproduction becomes imperative during long-range migration to ensure proper energy allocation. However, the mechanisms involved in this trade-off remain poorly understood. Here, we used a takeoff assay to distinguish migratory from non-migratory individuals in the fall armyworm, which is a major migratory insect worldwide. Migratory females displayed delayed ovarian development and flew further and faster than non-migratory females during tethered flight. Transcriptome analyses demonstrated an enrichment of fatty acid genes across successive levels of ovarian development and different migratory behaviors. Additionally, genes with roles in phototransduction and carbohydrate digestion along with absorption function were enriched in migratory females. Consistent with this, we identified increased abdominal lipids in migratory females that were mobilized to supply energy to the flight muscles in the thorax. Our study reveals that the fall armyworm faces a trade-off in allocating abdominal triglycerides between migration and reproduction during flight. The findings provide valuable insights for future research on this trade-off and highlight the key energy components involved in this strategic balance.

Feeding and Growth Response of Fall Armyworm Spodoptera frugiperda (Lepidoptera: Noctuidae) towards Different Host Plants.

The fall armyworm, Spodoptera frugiperda, is a major migratory polyphagous insect pest of various crops. The essential nutrient and mineral profile of the host plants determines the feeding fitness of herbivorous insects. As a result, the growth and development of insects is affected. To determine the effect of the nutrient and mineral profile of different host plants (maize, castor bean, cotton, cabbage, okra, and sugarcane) on the growth and development of S. frugiperda, biological parameters like larval weight, pupal weight (male/female), and feeding and growth indices were calculated. The proximate compositions such as crude protein, crude fat, crude fibre, and ash and mineral contents of the tested host plants showed significant differences (p < 0.05). The feeding indices on these host plants also differed significantly (p < 0.05). The maximum relative growth rate (RGR), relative consumption rate (RCR), and consumption index (CI) were recorded in S. frugiperda larvae that fed on maize and castor bean leaves. The crude protein, dry matter, and ash contents in maize and castor bean were significantly higher and positively correlated with the RGR and RCR of S. frugiperda larvae. The larval, male and female pupal weights were the maximum in the larvae feeding on the castor bean host plant. These findings provide novel information based on nutritional ecology to develop sustainable integrated pest management strategies using selective crop rotation.

Weather Influence on Fall Armyworm Infestation in Maize: A Study in Perambalur District, Tamil Nadu, India

Maize, Zea mays L (Family: Poaceae), is a main cereal crop cultivated across diverse agro-climatic regions worldwide. The Fall Armyworm (FAW), Spodopterafrugiperda (J. E. Smith) (Lepidoptera: Noctuidae), migratory insect is notorious for its economic impact, inflicting severe damage on various crops including maize, rice, sorghum, millet, sugarcane, vegetables, and cotton, and has been documented to affect over 80 other plant species. A roving survey was carried out in four major maize growing blocks of Perambalur district viz., Perambalur, Veppanthattai, Alathur and Veppur during Kharif and Rabi season of 2023-2024. The experiment consisted of sixty standard weeks from 3rd July 2023 to 4th August 2024. The results indicated a positive correlation between the number of fall armyworm larvae/plant with Tmax, relative humidity (Evening), and the negative correlation was observed between the number of fall armyworm larvae/plant with Tmin, relative humidity (Morning), Sunshine and rainfall. The data concluded that the maximum temperature significantly influenced the emergence of the fall armyworm and its infestation on maize. The incidence of FAW started in 46th mean standard week MSW (13 larvae/plant) Alathur, (17.25 larvae/plant) Perambalur, (17.25 larvae/plant) Veppanthattai, (18 larvae/plant) Veppur and the population increased gradually and reached the peak in the 47th mean standard week MSW (26 larvae/plant) Alathur, (32.75 larvae/plant) Perambalur, (33 larvae/plant) Veppanthattai, (42 larvae/plant). Correlation analysis revealed that larval population showed a significant positive correlation with the maximum temperature and relative humidity. At the same time, minimum temperature, sunshine, and rainfall exhibited a negative association with larval incidence of S. frugiperda. Propose adaptation strategies for maize farmers in Perambalur to manage fall armyworm infestations in the context of changing weather patterns. This could include adjustments in planting dates, pest monitoring techniques, or integrated pest management approaches.

Application of Instance Segmentation to Identifying Insect Concentrations in Data from an Entomological Radar

Entomological radar is one of the most effective tools for monitoring insect migration, capable of detecting migratory insects concentrated in layers and facilitating the analysis of insect migration behavior. However, traditional entomological radar, with its low resolution, can only provide a rough observation of layer concentrations. The advent of High-Resolution Phased Array Radar (HPAR) has transformed this situation. With its high range resolution and high data update rate, HPAR can generate detailed concentration spatiotemporal distribution heatmaps. This technology facilitates the detection of changes in insect concentrations across different time periods and altitudes, thereby enabling the observation of large-scale take-off, landing, and layering phenomena. However, the lack of effective techniques for extracting insect concentration data of different phenomena from these heatmaps significantly limits detailed analyses of insect migration patterns. This paper is the first to apply instance segmentation technology to the extraction of insect data, proposing a method for segmenting and extracting insect concentration data from spatiotemporal distribution heatmaps at different phenomena. To address the characteristics of concentrations in spatiotemporal distributions, we developed the Heatmap Feature Fusion Network (HFF-Net). In HFF-Net, we incorporate the Global Context (GC) module to enhance feature extraction of concentration distributions, utilize the Atrous Spatial Pyramid Pooling with Depthwise Separable Convolution (SASPP) module to extend the receptive field for understanding various spatiotemporal distributions of concentrations, and refine segmentation masks with the Deformable Convolution Mask Fusion (DCMF) module to enhance segmentation detail. Experimental results show that our proposed network can effectively segment concentrations of different phenomena from heatmaps, providing technical support for detailed and systematic studies of insect migration behavior.

CRY1 is involved in the take-off behaviour of migratory Cnaphalocrocis medinalis individuals

BackgroundNumerous insect species undertake long-distance migrations on an enormous scale, with great implications for ecosystems. Given that take-off is the point where it all starts, whether and how the external light and internal circadian rhythm are involved in regulating the take-off behaviour remains largely unknown. Herein, we explore this issue in a migratory pest, Cnaphalocrocis medinalis, via behavioural observations and RNAi experiments.ResultsThe results showed that C. medinalis moths took off under conditions where the light intensity gradually weakened to 0.1 lx during the afternoon or evening, and the take-off proportions under full spectrum or blue light were significantly higher than that under red and green light. The ultraviolet-A/blue light–sensitive type 1 cryptochrome gene (Cmedcry1) was significantly higher in take-off moths than that of non-take-off moths. In contrast, the expression of the light-insensitive CRY2 (Cmedcry2) and circadian genes (Cmedtim and Cmedper) showed no significant differences. After silencing Cmedcry1, the take-off proportion significantly decreased. Thus, Cmedcry1 is involved in the decrease in light intensity induced take-off behaviour in C. medinalis.ConclusionsThis study can help further explain the molecular mechanisms behind insect migration, especially light perception and signal transmission during take-off phases.

Heat-stress memory enhances the acclimation of a migratory insect pest to global warming.

In the face of rising global temperatures, the mechanisms behind an organism's ability to acclimate to heat stress remain enigmatic. The rice leaf folder, Cnaphalocrocis medinalis, traditionally viewed as temperature-sensitive, paradoxically exhibits robust larval acclimation to heat stress. This study used the heat-acclimated strain HA39, developed through multigenerational exposure to 39°C during the larval stage, and the unacclimated strain HA27 reared at 27°C to unravel the transgenerational effects of heat acclimation and its regulatory mechanisms. Heat acclimation for larvae incurred a fitness cost in pupae when exposed to high temperature, yet a significant transgenerational effect surfaced, revealing heightened fitness benefit in pupae from HA39, even without additional heat exposure during larval recovery at 27°C. This transgenerational effect exhibited a short-term memory, diminishing after two recovery generations. Moreover, the effect correlated with increased superoxide dismutase (SOD) enzyme activity and expression levels of oxidoreductase genes, representing physiological and molecular foundations of heat acclimation. Heat-acclimated larvae displayed elevated DNA methylation levels, while pupae from HA39, in recovery generations, exhibited decreased methylation indicated by the upregulation of a demethylase gene and downregulation of two methyltransferase genes at high temperatures. In summary, heat acclimation induces DNA methylation, orchestrating heat-stress memory and influencing the expression levels of oxidoreductase genes and SOD activity. Heat-stress memory enhances the acclimation of the migratory insect pest to global warming.

A comparison of the large-scale gene expression patterns in summer and fall migratory Pantala flavescens (Fabricius) in northern China.

Pantala flavescens (Fabricius) is the most well-known seasonal migratory insect. This research focused on the molecular response of P. flavescens migration in summer and fall. A total of 17,810 assembled unigenes were obtained and 624 differentially expressed genes (DEGs) were identified in summer migration compared to fall migration. A number of DEGs, including cpr49Ae, itm2b, chitinase, cpr11B, laccase2, nd5, vtg2 and so on, had previously been reported to be involved in cold- and high-temperature resistance. Functional enrichment analysis showed three pathways 'that antibacterial humoral response, response to bacterial, and lipid transporter activity' were significantly enriched in summer migration while that six pathways 'structural constituent of cuticle, chitin binding, mitochondrion, propanoate metabolism, citrate cycle, hypertrophic cardiomyopathy' were significantly enriched in fall migration. These results will provide a valuable baseline for further understanding of the molecular mechanisms of insect adaptation to different climate migrations.

The most remarkable migrants-systematic analysis of the Western European insect flyway at a Pyrenean mountain pass.

In autumn 1950 David and Elizabeth Lack chanced upon a huge migration of insects and birds flying through the Pyrenean Pass of Bujaruelo, from France into Spain, later describing the spectacle as combining both grandeur and novelty. The intervening years have seen many changes to land use and climate, posing the question as to the current status of this migratory phenomenon. In addition, a lack of quantitative data has prevented insights into the ecological impact of this mass insect migration and the factors that may influence it. To address this, we revisited the site in autumn over a 4 year period and systematically monitored abundance and species composition of diurnal insect migrants. We estimate an annual mean of 17.1 million day-flying insect migrants from five orders (Diptera, Hymenoptera, Hemiptera, Lepidoptera and Odonata) moving south, with observations of southward 'mass migration' events associated with warmer temperatures, the presence of a headwind, sunlight, low windspeed and low rainfall. Diptera dominated the migratory assemblage, and annual numbers varied by more than fourfold. Numbers at this single site hint at the likely billions of insects crossing the entire Pyrenean mountain range each year, and we highlight the importance of this route for seasonal insect migrants.

Pollen metabarcoding reveals the origin and multigenerational migratory pathway of an intercontinental-scale butterfly outbreak

Migratory insects may move in large numbers, even surpassing migratory vertebrates in biomass. Long-distance migratory insects complete annual cycles through multiple generations, with each generation's reproductive success linked to the resources available at different breeding grounds. Climatic anomalies in these grounds are presumed to trigger rapid population outbreaks. Here, we infer the origin and track the multigenerational path of a remarkable outbreak of painted lady (Vanessa cardui) butterflies that took place at an intercontinental scale in Europe, the Middle East, and Africa from March 2019 to November 2019. Using metabarcoding, we identified pollen transported by 264 butterflies captured in 10 countries over 7months and modeled the distribution of the 398 plants detected. The analysis showed that swarms collected in Eastern Europe in early spring originated in Arabia and the Middle East, coinciding with a positive anomaly in vegetation growth in the region from November 2018 to April 2019. From there, the swarms advanced to Northern Europe during late spring, followed by an early reversal toward southwestern Europe in summer. The pollen-based evidence matched spatiotemporal abundance peaks revealed by citizen science, which also suggested an echo effect of the outbreak in West Africa during September-November. Our results show that population outbreaks in a part of species' migratory ranges may disseminate demographic effects across multiple generations in a wide geographic area. This study represents an unprecedented effort to track a continuous multigenerational insect migration on an intercontinental scale.

Jeannie N. Shinozuka, Biotic borders: Transpacific plant and insect migration and the rise of anti-Asian racism in America, 1890-1950, Chicago and London: The University of Chicago Press, 2022.

Jeannie N. Shinozuka, Biotic borders: Transpacific plant and insect migration and the rise of anti-Asian racism in America, 1890-1950, Chicago and London: The University of Chicago Press, 2022.

Milkweed and floral resource availability for monarch butterflies (Danaus plexippus) in the United States

The global decline of pollinators, particularly insects, underscores the importance of enhanced monitoring of their populations and habitats. However, monitoring some pollinator habitat is challenging due to widespread species distributions and shifts in habitat requirements through seasons and life stages. The monarch butterfly (Danaus plexippus), a migratory insect pollinator that breeds widely throughout North America, presents a unique case study for testing a sampling framework to overcome these challenges. Monarchs exhibit discrete resource needs across life stages (e.g., larval requirement for milkweed, adult requirement for floral nectar), utilizing many land use types across their extensive geographic range during breeding and migration seasons. The Integrated Monarch Monitoring Program (IMMP) uses a standardized protocol with a generalized random tessellation stratified (GRTS) sampling design to gather spatially balanced and ecologically representative information on monarch habitats within the United States. The IMMP is applicable to various land use types and habitats used by breeding monarchs and may be extended to sites outside of the GRTS design to collect data on non-random sites of interest, such as legacy or conservation sites. Additionally, the IMMP’s modular design and publicly available training allows for broad participation, including involvement from community scientists. Here, we summarize habitat metrics (milkweed and floral resources) across 1,233 sites covering much of the monarch’s breeding range. We examine variation in milkweed density and floral resource availability on probabilistic (random) and non-probabilistic (convenience) samples and among land use types (site types). Additionally, we assess resource availability within core geographies for monarch breeding and migration, specifically within the U.S. Fish and Wildlife Service’s Monarch Conservation Units (western, northern, and southern United States). Milkweed density, floral frequency, and floral richness were higher on non-random sites and in the North region. Among site types, milkweed density was highest on Rights-of-Way and Unclassified Grassland, while floral frequency was lowest on Rights-of-Way. The IMMP represents the first field-based habitat monitoring program of this scale for monarchs, yielding a robust dataset on monarchs and their habitats across their breeding range and offering a framework for surveying the habitat of insect species with diverse habitat requirements or widespread distributions.

Potential range shift of a long-distance migratory rice pest, Nilaparvata lugens, under climate change

The biogeographical range shift of insect pests is primarily governed by temperature. However, the range shift of seasonal long-distance migratory insects may be very different from that of sedentary insects. Nilaparvata lugens (BPH), a serious rice pest, can only overwinter in tropical-to-subtropical regions, and some populations migrate seasonally to temperate zones with the aid of low-level jet stream air currents. This study utilized the CLIMEX model to project the overwintering area under the climate change scenarios of RCP2.6 and RCP8.5, both in 2030s and 2080s. The overwintering boundary is predicted to expand poleward and new overwintering areas are predicted in the mid-latitude regions of central-to-eastern China and mid-to-southern Australia. With climate change, the habitable areas remained similar, but suitability decreased substantially, especially in the near-equatorial regions, owing to increasing heat stress. The range shift is similar between RCP2.6-2030s, RCP2.6-2080s, and RCP8.5-2030s, but extreme changes are projected under RCP8.5-2080s with marginal areas increasing from 27.2 to 38.8% and very favorable areas dropping from 27.5 to 3.6% compared to the current climate. These findings indicate that climate change will drive range shifts in BPH and alter regional risks differently. Therefore, international monitoring programs are needed to effectively manage these emerging challenges.

Continental-scale patterns in diel flight timing of high-altitude migratory insects.

Many insects depend on high-altitude, migratory movements during part of their life cycle. The daily timing of these migratory movements is not random, e.g. many insect species show peak migratory flight activity at dawn, noon or dusk. These insects provide essential ecosystem services such as pollination but also contribute to crop damage. Quantifying the diel timing of their migratory flight and its geographical and seasonal variation, are hence key towards effective conservation and pest management. Vertical-looking radars provide continuous and automated measurements of insect migration, but large-scale application has not been possible because of limited availability of suitable devices. Here, we quantify patterns in diel flight periodicity of migratory insects between 50 and 500 m above ground level during March-October 2021 using a network of 17 vertical-looking radars across Europe. Independent of the overall daily migratory movements and location, peak migratory movements occur around noon, during crepuscular evening and occasionally the morning. Relative daily proportions of insect migration intensity and traffic during the diel phases of crepuscular-morning, day, crepuscular-evening and night remain largely equal throughout May-September and across Europe. These findings highlight, extend, and generalize previous regional-scale findings on diel migratory insect movement patterns to the whole of temperate Europe. This article is part of the theme issue 'Towards a toolkit for global insect biodiversity monitoring'.

Effects of nocturnal celestial illumination on high-flying migrant insects.

Radar networks hold great promise for monitoring population trends of migrating insects. However, it is important to elucidate the nature of responses to environmental cues. We use data from a mini-network of vertical-looking entomological radars in the southern UK to investigate changes in nightly abundance, flight altitude and behaviour of insect migrants, in relation to meteorological and celestial conditions. Abundance of migrants showed positive relationships with air temperature, indicating that this is the single most important variable influencing the decision to initiate migration. In addition, there was a small but significant effect of moonlight illumination, with more insects migrating on full moon nights. While the effect of nocturnal illumination levels on abundance was relatively minor, there was a stronger effect on the insects' ability to orientate close to downwind: flight headings were more tightly clustered on nights when the moon was bright and when cloud cover was sparse. This indicates that nocturnal illumination is important for the navigational mechanisms used by nocturnal insect migrants. Further, our results clearly show that environmental conditions such as air temperature and light levels must be considered if long-term radar datasets are to be used to assess changing population trends of migrants. This article is part of the theme issue 'Towards a toolkit for global insect biodiversity monitoring'.

Monitoring insect numbers and biodiversity with a vertical-beam entomological radar.

Concerns about perceived widespread declines in insect numbers have led to recognition of a requirement for long-term monitoring of insect biodiversity. Here we examine whether an existing, radar-based, insect monitoring system developed for research on insect migration could be adapted to this role. The radar detects individual larger (greater than 10 mg) insects flying at heights of 150-2550 m and estimates their size and mass. It operates automatically and almost continuously through both day and night. Accumulation of data over a 'half-month' (approx. 15 days) averages out weather effects and broadens the source area of the wind-borne observation sample. Insect counts are scaled or interpolated to compensate for missed observations; adjustment for variation of detectability with range and insect size is also possible. Size distributions for individual days and nights exhibit distinct peaks, representing different insect types, and Simpson and Shannon-Wiener indices of biodiversity are calculated from these. Half-month count, biomass and index statistics exhibit variations associated with the annual cycle and year to year changes that can be attributed to drought and periods of high rainfall. While species-based biodiversity measures cannot be provided, the radar's capacity to estimate insect biomass over a wide area indicates utility for tracking insect population sizes. This article is part of the theme issue 'Towards a toolkit for global insect biodiversity monitoring'.

Abundance and Seasonal Migration Patterns of Green Lacewings (Neuroptera: Chrysopidae) across the Bohai Strait in Eastern Asia.

Many insects, including green lacewings, migrate seasonally to exploit suitable breeding and winter habitats. Green lacewings are important natural enemies of insect pests worldwide. Here, four dominant green lacewing species, Chrysoperla nipponensis (Okamoto), Chrysopa pallens (Rambur), Chrysoperla furcifera (Okamoto), and Chrysopa formosa Brauer, were investigated for their ability to migrate between northern and northeastern China across the Bohai Strait from late May to late October each year. Furthermore, there were significant interannual and seasonal differences in the number of migratory green lacewings collected. The number of green lacewings in spring was significantly lower than that in summer and autumn, and the highest average number of green lacewings occurred in June. In addition, there were differences in the sex ratio of migrating green lacewings between months, with a greater proportion of females than males. Finally, the seasonal migration trajectories simulated by the HYSPLIT model revealed that the green lacewings captured on Beihuang Island primarily originated from Shandong Province. Accordingly, these findings contribute to our understanding of green lacewing migration in eastern Asia and aid its incorporation within integrated pest management (IPM) packages for several crop pests. Furthermore, long-term tracking of migrant insect populations can reveal ecosystem services and trophic dynamic processes at the macroscale.

Massive seasonal high-altitude migrations of nocturnal insects above the agricultural plains of East China

Long-distance migrations of insects contribute to ecosystem functioning but also have important economic impacts when the migrants are pests or provide ecosystem services. We combined radar monitoring, aerial sampling, and searchlight trapping, to quantify the annual pattern of nocturnal insect migration above the densely populated agricultural lands of East China. A total of ~9.3 trillion nocturnal insect migrants (15,000 t of biomass), predominantly Lepidoptera, Hemiptera, and Diptera, including many crop pests and disease vectors, fly at heights up to 1 km above this 600 km-wide region every year. Larger migrants (>10 mg) exhibited seasonal reversal of movement directions, comprising northward expansion during spring and summer, followed by southward movements during fall. This north-south transfer was not balanced, however, with southward movement in fall 0.66× that of northward movement in spring and summer. Spring and summer migrations were strongest when the wind had a northward component, while in fall, stronger movements occurred on winds that allowed movement with a southward component; heading directions of larger insects were generally close to the track direction. These findings indicate adaptations leading to movement in seasonally favorable directions. We compare our results from China with similar studies in Europe and North America and conclude that ecological patterns and behavioral adaptations are similar across the Northern Hemisphere. The predominance of pests among these nocturnal migrants has severe implications for food security and grower prosperity throughout this heavily populated region, and knowledge of their migrations is potentially valuable for forecasting pest impacts and planning timely management actions.

Continental-scale migration patterns and origin of Helicoverpa zea (Lepidoptera: Noctuidae) based on a biogeochemical marker.

Insect migrations have ecological and economic impacts, particularly in agriculture. However, there is limited knowledge about the migratory movements of pests at the continental scale, which is an important factor influencing the spread of resistance genes. Understanding the migratory patterns of economic pests, like Helicoverpa zea (Boddie), is essential for improving Integrated Pest Management (IPM) and Insect Resistance Management (IRM) strategies. In this study, we used stable hydrogen isotopic ratios in wing tissue as a biogeochemical marker to examine migratory patterns and estimate the native origins of H. zea individuals collected across a wide latitudinal gradient in North America. Samples collected at higher latitudes (Ontario, Canada and Minnesota, USA) exhibited a greater proportion (60%-96%) of nonlocal individuals, with an increased probability of origin from the southeastern United States. Populations from mid-latitudes (Florida, North Carolina, and South Carolina) showed a blend of local and nonlocal (40%-60%) individuals. Finally, 15% of the southernmost population individuals (Puerto Rico) were classified as migratory, with some having a probability of origin at higher latitudes (>30°). Overall, our results provide evidence of a northward spring/summer migration of H. zea in North America and underscore the significance of the southeastern United States as a hub for genetic flow. In addition, based on stable hydrogen isotopic ratios, there is strong evidence of reverse (southward) migration of H. zea from the continental United States to Puerto Rico. Our study highlights the implications for IPM and IRM programs and the need for management strategies that account for both northward and southward migration patterns.

Biotic Borders: Transpacific Plant and Insect Migration and the Rise of Anti-Asian Racism in America, 1890–1950

Biotic Borders: Transpacific Plant and Insect Migration and the Rise of Anti-Asian Racism in America, 1890–1950