The migratory locust (Locusta migratoria) is the sole species within the genus Locusta, a short-horned grasshopper belonging to the family Acrididae, distinguished by its phase polyphenism that enables transitions between a solitary, non-swarming phase and a gregarious phase characterized by aggregation, behavioral changes, and long-distance migration in dense hopper bands and adult swarms.[1][2] This phenotypic plasticity, triggered primarily by population density, results in morphological, physiological, and behavioral differences between phases, with gregarious individuals displaying bolder coloration, larger wing muscles, and heightened locomotion.[3][4]Native to the Old World, L. migratoria occupies diverse habitats from sea level to over 4,000 meters in elevation, favoring wetlands such as river deltas, lake shores, and reed beds dominated by plants like Phragmites species, across regions spanning Africa, Asia, southern Europe, and Australia.[1][5] Adults typically measure 35–50 mm in length for males and 45–55 mm for females, with swarms capable of covering thousands of square kilometers and consuming up to 2,000 times their body weight in vegetation daily during outbreaks.[6][7] These plagues have inflicted substantial economic losses on agriculture throughout history, driving international monitoring programs and control strategies focused on early intervention in breeding areas.[7][8]
Taxonomy and classification
Etymology and nomenclature
The scientific name Locusta migratoria was established by Carl Linnaeus in the tenth edition of Systema Naturae published on October 1, 1758, under the binomial nomenclature system.[9] The genus Locusta originates from the Latin locusta, denoting a locust or grasshopper, a term used in classical texts to describe such orthopterans.[10] The specific epithet migratoria is a Latin adjective derived from migratorius, meaning "wandering" or "migratory," directly referencing the species' characteristic long-distance swarming flights and invasions.[9]In nomenclature, L. migratoria is the type and sole species of the monotypic genus Locusta within the family Acrididae.[1] It has accumulated numerous synonyms over time, exceeding 20 historical designations, often reflecting regional variants or misclassifications as distinct species or subspecies prior to modern taxonomic revisions.[1] The common English name "migratory locust" emphasizes its propensity for forming dense, mobile swarms capable of traversing continents, distinguishing it from non-migratory grasshopper phases. Subspecies nomenclature includes forms like L. m. migratorioides for African populations, highlighting geographic variation in swarming potential.[11]
Scientific classification
The migratory locust is formally described as Locusta migratoria Linnaeus, 1758, the type and sole species in the genus Locusta.[1][12] It belongs to the family Acrididae, which encompasses most short-horned grasshoppers and locusts, and the subfamily Oedipodinae, characterized by banded hind legs and habitat preferences in open grasslands.[1][13]
This classification reflects the organism's placement among hemimetabolous insects with incomplete metamorphosis, adapted for jumping via enlarged hind legs.[11][12] The binomial authority traces to Carl Linnaeus's Systema Naturae (10th edition, 1758), where it was initially described from European specimens, though the species exhibits cosmopolitan distribution across Africa, Asia, Australia, and parts of Europe.[1] Subspecies distinctions, such as L. m. migratorioides in Africa, are recognized based on morphological and geographic variation but do not alter the core species-level taxonomy.[14][11]
Subspecies and geographic variation
The migratory locust (Locusta migratoria) is classified into several subspecies, primarily distinguished by morphological, genetic, and distributional differences across its vast Old World range. The nominal subspecies L. m. migratoria (Linnaeus, 1758), often referred to as the Eurasian or Asian migratory locust, predominates in temperate and subtropical regions of Europe, Central Asia, and eastern China, where it exhibits adaptations such as embryonic diapause to overwinter in colder climates.[15][6] In contrast, L. m. migratorioides (Reiche & Fairmaire, 1849), the African migratory locust, is adapted to tropical and semi-arid environments in sub-Saharan Africa, with populations showing enhanced migratory capabilities and less pronounced diapause due to milder seasonal fluctuations.[16][17] The Oriental subspecies L. m. manilensis (Meyen, 1835) occupies southeastern Asia, including the Philippines and parts of Indonesia, featuring smaller body sizes and higher densities in rice paddy habitats compared to northern conspecifics.[18][19]These subspecies reflect clinal variations tied to latitude and climate, with northern populations (L. m. migratoria) demonstrating greater cold hardiness in eggs, enabling survival at temperatures as low as -30°C through supercooling and cryoprotectant accumulation, whereas central and southern forms prioritize faster development over winter tolerance.[20] Photoperiodic responses for diapause induction also vary geographically: higher-latitude strains enter diapause earlier under short-day conditions to align with seasonal breeding, as evidenced by experiments showing critical day-length thresholds decreasing from 13-14 hours in northern Chinese populations to shorter durations in equatorial ones.[21] Body size exhibits non-linear patterns, increasing southward in northern ranges before declining in central arid zones and expanding again in southern tropics, correlated with resource availability and predation pressures rather than Bergmann's rule alone.[22] RNAi sensitivity, a proxy for genetic differentiation, further differs inter- and intra-populationally, with African strains showing reduced efficacy compared to Asian ones, suggesting adaptive divergence in generegulation.[23] Such variations underscore L. migratoria's phenotypic plasticity, enabling persistence from sea level to over 4,000 m elevation across Africa, Asia, and into Australia.[1]
Morphology and physical characteristics
Adult morphology
The adult migratory locust (Locusta migratoria) possesses a body structure characteristic of the family Acrididae, comprising a distinct head, thorax, and abdomen, with fully developed wings enabling long-distance flight. Body length varies by phase and sex: in the solitary phase, females measure 50-70 mm and males 40-55 mm, while in the gregarious phase, females are 40-45 mm and males 35-45 mm long.[1] The head features large compound eyes, three ocelli, and filiform antennae shorter than the body length; mouthparts are mandibulate for chewingvegetation. The pronotum on the thorax is raised with a median carina, and the underside is often hairy; hind legs are enlarged for jumping, bearing spines on the tibiae and featuring a tympanal organ for sound detection.[24]Forewings, or tegmina, are leathery and cover the abdomen, displaying dark coloration with thin pale longitudinal lines in both phases; hindwings are hyaline and transparent, sometimes faintly smoky at the apex, folded fan-like beneath the tegmina when at rest. Coloration exhibits polyphenic variation: solitary adults are typically green or brown for crypsis, adapting to humidity and environment, whereas gregarious adults develop bright yellow hues with black markings on the head, thorax, and forewings, and males may intensify to distinct yellow upon maturation. Hind femora are brownish to bluish-black internally, with tibiae yellowish, beige, or red.[25][1] The abdomen is elongated with 11 segments, terminating in cerci; females possess a robust ovipositor for egg-laying.[8]
Nymphal stages
The nymphs of the migratory locust, Locusta migratoria, hatch from egg pods buried in soil and progress through five instars over approximately 20-40 days, depending on temperature and nutrition, with molting occurring between each stage to allow for growth in size and morphological development.[26][27] First instar nymphs measure about 3-5 mm in length, lack wing pads, and possess a soft exoskeleton with simple antennae and mouthparts adapted for herbivory.[28] Their legs are structured for jumping, enabling rapid movement despite the absence of wings.[29]In the second instar, small wing pads become visible as external thoracic protrusions, marking the onset of imaginal disc development for future adult wings.[30] Nymphs increase in size to around 8-10 mm, with enhanced jumping capability due to proportional leg elongation. By the third instar, wing pads enlarge and orient backwards in a spade-like shape, while body length reaches 15-20 mm; coloration in gregarious individuals features contrasting black patterns on a yellowish or orange background, aiding in phase-specific camouflage or signaling.[31][32]Fourth instar nymphs exhibit further wing pad expansion, with pads positioned more laterally and body size approaching 30 mm, accompanied by denser setae for sensory function.[31] The final, fifth instar is the largest at 40-50 mm, with wing pads shifting forward and darkening as they prepare to unfold into functional wings post-molt; reproductive structures begin differentiating, though full maturity occurs in adults.[28][1] Throughout these stages, gregarious nymphs form cohesive hopper bands, marching in unison, which influences physical wear on the exoskeleton from collective movement but is primarily behavioral.[33] Rarely, six or seven instars occur under suboptimal conditions, extending development time.[1]
Sexual dimorphism
Adult females of the migratory locust (Locusta migratoria) exhibit pronounced sexual size dimorphism, typically measuring 54–72 mm in body length, while males range from 42–55 mm.[34] This size difference aligns with broader patterns in acridid grasshoppers, where larger female body size correlates with increased ovariole number and egg production capacity, enabling females from larger populations to produce egg pods with more eggs.[22] The degree of sexual size dimorphism tends to increase as mean population body sizes grow, deviating from Rensch's rule observed in many other taxa.[22]Morphometric ratios, such as the hind femur length to head width (F/C ratio), differ between sexes; from the fourth instar onward, females maintain a lower F/C ratio than males of the same instar and phase, reflecting subtle proportional variances in leg and head structures.[35] Females possess highly extensible abdominal intersegmental membranes that are thicker, with a lamella epicuticle and extensive folding, adaptations that accommodate egg-laying by allowing significant abdominal expansion absent in males.[36]In males, reproductive morphology includes a single testicular organ, unique among many orthopterans that typically have paired testes, with follicular cells organized in a distinctive manner supporting spermatogenesis.[37] Genital nerves in males extend from abdominal ganglia to accessory reproductive structures, facilitating copulatory behaviors, whereas female ganglia lack such exiting neuronal processes.[38] Body coloration shows no consistent sex-specific patterns beyond phase polyphenism influences, with both sexes displaying green or brown in the solitary phase and yellow-black in gregarious adults.[34]
Polyphenism and physiological adaptations
Solitary versus gregarious phases
The migratory locust, Locusta migratoria, displays density-dependent phase polyphenism, manifesting as solitary and gregarious phases that differ markedly in morphology, behavior, physiology, and ecology. These phases represent extremes of a continuous spectrum induced by population density, with the solitary phase predominant at low densities and the gregarious phase emerging under crowded conditions.[39]In the solitary phase, individuals exhibit cryptic coloration, typically green or brown, facilitating camouflage against predators, along with a longer hind femur and a straight or concave pronotal keel. Behaviorally, they are shy and sedentary, actively avoiding conspecifics, walking slowly with a low body posture, and preferring nocturnal flight. Physiologically, solitary locusts show elevated levels of dopamine, tyramine, and octopamine, particularly under isolation, and express higher levels of antioxidant-related genes. Ecologically, this phase supports dispersed, low-density populations with higher individual fecundity in some contexts, though overall reproductive output is lower than in outbreaks.[39][40]Conversely, the gregarious phase features conspicuous yellow and black patterning in nymphs, shorter hind femurs, and distinct pronotal shapes, serving as warning coloration during swarming. These locusts are highly active, forming cohesive hopper bands and adult swarms that aggregate over vast areas exceeding 1000 km², with rapid walking, elevated body posture, and diurnal flight. Crowding triggers increased serotonin levels, promoting attraction to conspecifics via pheromones like 4-vinylanisole, and upregulates stress-response genes alongside enhanced dopamine signaling through specific receptors that reinforce gregarization. Gregarious forms exhibit superior migration capabilities and dietary breadth, including tolerance for toxic plants containing hyoscyamine, but trade-offs include reduced individual size and potentially fewer generations per year (3 versus 4-5 in solitary).[39]
Aspect
Solitary Phase
Gregarious Phase
Morphology
Cryptic green/brown; longer hind femur; straight/concave pronotal keel
Yellow/black patterns; shorter hind femur; distinct pronotum
Swarming over large areas; pheromone-driven aggregation[39][41]
Triggers for phase change
The transition from the solitary to the gregarious phase in the migratory locust (Locusta migratoria) is primarily triggered by an increase in local population density, which induces behavioral and physiological changes through heightened social interactions.[42] This density-dependent polyphenism, known as gregarization, occurs when locusts aggregate, leading to sensory stimulation that overrides solitary tendencies.[43]
Tactile cues are the most potent initiators, with physical contact—particularly repeated touching of the hind femora between nymphs—mimicking crowding and rapidly promoting gregarious traits such as increased activity and attraction to conspecifics.[43] Laboratory experiments demonstrate that such stimulation can induce phase shifts within hours, especially during sensitive nymphal instars (typically third to fifth). Visual stimuli from seeing groups of locusts and olfactory signals, including aggregation pheromones released by crowded individuals, amplify these effects, facilitating cluster formation and band development.[43]
A critical density threshold of approximately 0.05 locusts per square centimeter destabilizes uniform distributions, promoting instability and the coalescence of small groups into larger hopper bands that sweep up additional solitarious individuals, accelerating widespread gregarization.[43] These triggers are reversible; reducing density allows solitarization, though the process is slower and primarily behavioral in adults. Environmental factors like resource availability can modulate density buildup, but sensory integration from crowding remains the core causal mechanism.[42]
Neurochemical mechanisms
The neurochemical mechanisms driving phase polyphenism in the migratory locust (Locusta migratoria) primarily involve biogenic amines such as serotonin, dopamine, octopamine, and tyramine, which exhibit dynamic changes in central nervous system (CNS) levels and receptor signaling in response to crowding or isolation, thereby modulating behavioral transitions between solitary and gregarious phases.[44][3] These neurotransmitters integrate sensory cues, like mechanostimulation from conspecifics, to alter locomotion, attraction/repulsion responses, and aggregation tendencies, with the catecholamine biosynthetic pathway upregulated in the gregarious phase to support swarm formation.[3]Serotonin levels in the thoracic ganglia surge ninefold within 4 hours of crowding solitary-phase locusts, facilitating the sensory processing that initiates gregarization, though levels decline toward baseline by 24 hours before stabilizing at higher gregarious values after a full instar.[44] However, serotonin signaling in the brain promotes solitariness: isolation of gregarious nymphs elevates brain serotonin, and injection accelerates the transition to solitary behavior (reaching 73.3% solitarious probability after 30 minutes), while 5-HT2 receptor activation during crowding inhibits gregarious traits in solitary individuals.[45] In contrast, serotonin precursors like 5-HTP, combined with brief crowding, partially induce gregarious behaviors but fail to sustain full phase change without additional factors.[3]Dopamine exhibits bidirectional regulation, with CNS levels rising within 4 hours of crowding (solitary to gregarious) and falling during isolation (gregarious to solitary), remaining elevated in long-term gregarious locusts.[44] A single brain injection of dopamine (80 μg/μL) or agonistapomorphine (10 μg/μL) shifts solitary nymphs to gregarious behaviors within 1 hour, while repeated dosing with crowding completes the transition over one instar; conversely, silencing dopaminebiosynthesis genes like pale reverts ~50% of gregarious nymphs to solitary.[3] This duality arises from two receptors: DA-Dop1 (D1-like) in the brain drives gregarization via increased attraction and locomotion (agonist SKF38393 promotes these in solitary locusts; antagonist or RNAi induces repulsion in gregarious), whereas DA-Dop2 signaling maintains solitariness (antagonist sulpiride blocks isolation-induced repulsion in gregarious locusts).[46]Octopamine and tyramine further fine-tune phase-specific responses, with octopamine levels higher in gregarious brains and injections (100 μg) enhancing attraction to conspecific volatiles in solitary locusts, mediated by OARα receptor signaling that sustains aggregative behavior during crowding.[47]Tyramine, elevated in solitary phases, induces repulsion via TAR receptors, with injections promoting solitary traits in gregarious locusts and knockdown preventing isolation-driven avoidance.[47] These amines interact antagonistically, where their ratio influences behavioral preference, underscoring a networked neurochemical control that integrates environmental triggers for adaptive phase shifts.[47]
Life cycle and reproduction
Egg development and oviposition
Adult females of Locusta migratoria select oviposition sites in moist, sandy loam soils, using their ovipositor to excavate a vertical chamber 5–10 cm deep.[48] They deposit eggs in a single vertical string within a foam-lined pod, with each pod containing 30–100 eggs measuring 6–8 mm in length.[49][50] The pod, approximately 100 mm long and 8.5 mm in diameter, is capped by a hardened frothy secretion that facilitates gas exchange, retains moisture, and deters predators.[49] A female typically produces 2–6 pods over her adult lifespan, with oviposition occurring at intervals of about 20 days and requiring soil temperatures above 21°C.[51][52]Embryonic development within the pod is strongly temperature-dependent, with optimal rates at 30°C yielding hatching in 10–12 days, whereas 20°C extends the period to 39 days and reduces viability.[53][54] Warmer incubation produces larger hatchlings with reduced melanism, resembling gregarious phase traits, while cooler conditions result in smaller, darker nymphs akin to solitary phase characteristics.[53] Soil moisture also modulates outcomes; drier conditions during development yield smaller, paler hatchlings from larger eggs, though nymphal instar number remains consistent at five.[53]Hatching from pods is synchronized, typically spanning a mean of 2.4 hours, with embryos responsive to cues from neighboring eggs to coordinate emergence.[55] Phase state influences oviposition; gregarious females lay approximately 17% fewer eggs per pod than solitarious females due to elevated oocyte oosorption rates of 21–31% in later developmental stages.[56] This reduction correlates with higher expression of regulatory genes like activinβ, targeted by microRNA-34, underscoring physiological trade-offs in reproductive allocation between phases.[56]
Nymphal growth and molting
The nymphal stage of the migratory locust (Locusta migratoria) consists of five instars, with rare occurrences of six or seven, during which hoppers grow through continuous feeding and periodic molting to replace the restrictive exoskeleton.[1] First-instar nymphs hatch measuring roughly 3-5 mm in length, wingless and initially pale, progressing to larger sizes with developing wing buds by the later instars. Each instar ends with ecdysis, a hormonally regulated process involving apolysis, new cuticle secretion, and shedding of the old integument, primarily controlled by ecdysteroids like ecdysone and facilitated by enzymes such as chitinases.[57][58]Nymphal growth relies on voracious herbivory, with body mass increasing exponentially across instars; for instance, fifth-instar nymphs can reach lengths of 40-50 mm before the final molt to adulthood.[26] The total duration of nymphal development typically spans 24-35 days under laboratory conditions at around 30°C, though it extends in cooler temperatures or solitary phase due to slower metabolic rates and reduced feeding efficiency.[27][1] In the gregarious phase, induced by high population density, development accelerates with shorter instar periods and heightened locomotion, enabling synchronized band formation as nymphs march collectively while feeding.[59]Molting vulnerability peaks during ecdysis, when nymphs are soft-bodied and immobile for several hours, increasing predation risk, though gregarious bands provide some protective aggregation. Successive molts not only enlarge the body but also refine morphological traits, such as darkening coloration and longer legs in gregarious forms, adapting them for swarming behavior post-fledging.[60] Environmental factors like temperature profoundly influence instar durations—higher optima (28-35°C) shorten them, while extremes cause developmental delays or mortality.[61] Phase polyphenism further modulates growth, with gregarious nymphs exhibiting larger head capsules and faster biomass gain per instar compared to solitary counterparts.[62]
Adult longevity and fecundity
Adult migratory locusts exhibit varying longevity depending on phase and environmental conditions. In the gregarious phase, male adults have a median lifespan of 22 days and a maximum of 33 days under laboratory conditions.[63] Gregarious individuals generally experience shorter lifespans compared to solitary ones, with the former being approximately 44% shorter due to density-dependent physiological stresses.[64] In field populations, adult longevity ranges from about 90 days in northern regions to 120-150 days in southern areas, influenced by climate and resource availability.[1]Fecundity in female migratory locusts is characterized by the production of multiple egg pods over their adult lifespan. A single female typically lays 3 to 5 egg pods, each containing 50 to 60 eggs, resulting in a total of 150 to 300 eggs per female.[65] Egg pod numbers average around 2.2 per female, with 42 to 68 eggs per pod in some populations. Phase differences affect output, with solitarious females producing more eggs than gregarious ones; the latter lay 17% fewer eggs due to higher oosorption rates in oocytes.[56] Total eggs per female can vary from 66 to 116 depending on phase polymorphism.[1] Factors such as male presence enhance reproductive parameters, with paired females producing up to 6 pods of 50 eggs each.[66]
Behavior and ecology
Solitary phase behaviors
In the solitary phase, Locusta migratoria individuals maintain behaviors that foster isolation and mimic those of non-plague grasshoppers, actively avoiding aggregation even at low densities. This phase is characterized by a pronounced aversion to conspecifics, with locusts responding to proximity by fleeing or orienting away, thereby minimizing physical contact and potential tactile stimulation that could trigger phase shifts.[67][68] Behavioral assays demonstrate that solitary nymphs and adults spend over 90% of their time isolated, showing rapid escape responses when approached within 5-10 cm by others, in contrast to the attraction observed in gregarious forms.[68][69]Locomotor activity is subdued, featuring slow walking speeds averaging 0.1-0.2 m/s, shorter stride lengths, and extended resting periods that enhance crypsis against predators through motionless postures aligned with vegetation.[70] Flight is rare and limited to short bursts for evasion, lacking the sustained, directed flights of the gregarious phase; adults cover distances under 1 km per day under natural conditions.[4]Foraging involves selective, individual consumption of foliage from diverse plant species, with daily intake rates 20-50% lower than in gregarious individuals, reflecting reduced metabolic demands and a strategy to avoid drawing attention through minimal disturbance.[71]Reproductive behaviors in solitary adults emphasize discretion, with males producing softer, less frequent calling songs and females ovipositing in scattered, low-density sites to prevent clustering of offspring; copulation rates are lower due to mutual avoidance, extending generation times to 4-5 per year in temperate regions.[8] These traits collectively sustain sparse populations, with densities typically below 1 adult per square meter, preventing the tactile and visual cues that precipitate gregarization.[72][73]
Gregarious phase and swarming dynamics
In the gregarious phase, Locusta migratoria individuals display heightened attraction to conspecifics through visual, olfactory, and tactile cues, resulting in rapid aggregation within 4–8 hours of crowding.[74][43] This behavioral shift promotes increased activity levels compared to the solitary phase, facilitating the formation of dense groups.[74]Nymphs in the gregarious phase form hopper bands starting from the first instar, a few days post-hatching, where they exhibit coordinated marching behavior.[74] These bands emerge when local densities exceed a critical threshold of approximately 0.05 locusts per cm², beyond which uniform distributions become unstable and transient clumps merge into propagating bands.[43] Marching occurs intermittently, typically twice daily during forenoon and afternoon periods, with only about 10% of hoppers actively moving at any time due to frequent pauses; individual speeds are 3–4 times greater than the overall band progression rate.[74] Quiescent bands achieve high densities, occupying 2–4 times less area than during active marching, which can span from hundreds of square meters to several kilometers.[74]Upon reaching adulthood, gregarious locusts transition to forming flying swarms characterized by collective downwind flight, with individuals maintaining separations of roughly 10 cm and synchronizing wing beats.[74] Swarm cohesion is sustained through edge-turning behaviors, where peripheral locusts redirect inward, supported by local interactions modeled as self-propelled particle systems exhibiting phase transitions from disorder to order.[74] Tactile stimuli, such as hind-leg stroking, reinforce gregarization via serotonin-mediated pathways, underpinning the sensory-driven dynamics of both hopper bands and adult swarms.[43]
Migration patterns and dispersal
Gregarious phase adults of the Locusta migratoria form swarms that enable long-distance migration, primarily during daylight hours, with flights oriented predominantly downwind to maximize dispersal efficiency.[1] These swarms can cover 5 to 130 km or more per day, influenced by wind assistance, airspeed of approximately 10-16 km/h, and endurance allowing sustained flight for several hours.[75][43] Ground speeds increase with tailwinds, enabling swarms to traverse regions spanning hundreds of kilometers in favorable conditions, as observed in outbreak dynamics where populations expand from core breeding areas.[43]Prior to adult flight, gregarious nymphs contribute to dispersal through coordinated marching bands, advancing up to 3 km per day in low-vegetation habitats, which facilitates local spread before fledging into migratory adults around 10 days post-molting.[5] This phased dispersal strategy, triggered by density-dependent phase polyphenism, allows outbreak populations to invade new territories, with swarms potentially spanning areas from less than 1 km² to several hundred km² containing 40-80 million individuals per km².[75][43]In contrast, solitarious phase individuals exhibit more limited dispersal, conducting nocturnal flights that support gene flow and recession-phase persistence without forming destructive swarms.[1] Overall, migration patterns reflect adaptations for exploiting transient resources, with wind patterns and phase shifts driving irregular, large-scale invasions rather than fixed routes, as evidenced by phylogeographic studies tracing historical dispersal via mitochondrial DNA.[76]
Habitat requirements and distribution
The migratory locust (Locusta migratoria) requires habitats with light sandy or loamy soils suitable for egg-laying, typically near standing water such as rivers, lakes, marshes, or seasonal floodplains, where females burrow pods 5-10 cm deep into moist but aerated ground.[1][6] Breeding is confined to areas experiencing periodic rainfall or flooding that maintains soil moisture for 2-6 weeks post-oviposition, enabling egg incubation at optimal temperatures of 25-35°C, with hatching times ranging from 10 to 40 days depending on conditions.[1] Vegetation preferences include dense stands of short, succulent grasses and reeds such as Phragmites communis and sedges along water margins, providing nutrition for nymphs and supporting hopper band formation during outbreaks; persistent solitarious populations occur in drier perennial grasslands and steppes adjacent to these sites.[6][77] Outbreak dynamics are driven by precipitation exceeding 25-50 mm triggering vegetation green-up in semi-arid zones, with population recessions tied to prolonged dry periods exceeding habitat water balance capacity.[6][78]Geographic distribution encompasses much of the Eastern Hemisphere, from the Atlantic coasts of North Africa eastward across southern Europe, the Middle East, and Asia (including India, China, and Japan) to Australia, Papua New Guinea, and New Zealand, with sporadic records in Madagascar.[6][79] Subspecies reflect regional variations: L. m. migratorioides in sub-Saharan Africa, L. m. migratoria in temperate Eurasia and Central Asia, L. m. rossica in Russian forest-steppes, and L. m. manilensis (Oriental migratory locust) in Southeast Asia.[6][80] Permanent breeding foci include river deltas (e.g., Danube, Amu Darya) emptying into the Black, Caspian, and Aral Seas, Lake Balkhash basins, and northern Caspian lowlands, with altitudinal occupancy from sea level to over 4,000 m in Central Asian highlands.[6] Northern limits align with the southern fringes of coniferous forests in Europe and Asia, while southern extents reach New Zealand's temperate zones.[6]Climate change projections indicate potential range expansions northward into higher latitudes, contingent on increased precipitation variability.[81]
Foraging and diet
The migratory locust (Locusta migratoria) is a polyphagous herbivore that consumes a diverse array of vegetation, including grasses from the Poaceae family such as wheat and barley, as well as broadleaf plants, crops like maize and sorghum, and occasionally trees or shrubs when other food is scarce.[82] Its diet supports rapid growth and reproduction, with nymphs and adults capable of ingesting up to their body weight in fresh plant matter daily under optimal conditions, though exact rates vary by instar, phase, and plant quality.[83] In laboratory settings, locusts thrive on diets supplemented with wheat bran or germ, which enhance fat content and survival compared to grass alone.[84]Locusts actively regulate macronutrient intake, preferentially balancing protein and digestible carbohydrates at a ratio of approximately 1:1 in nymphs to optimize growth and energy for locomotion.[85] They employ compensatory feeding mechanisms, increasing consumption of deficient nutrients while adjusting post-ingestive processes like excretion or metabolism to maintain homeostasis; for instance, high-carbohydrate diets lead to elevated lipid storage, aiding flight endurance in migratory phases.[86] Nutritional imbalances, such as excess protein relative to carbohydrates, suppress phase polyphenism and reduce flight performance.[87]Feeding preferences differ between solitary and gregarious phases: solitarious individuals select foods more selectively, avoiding alkaloid-laden plants to minimize toxicity risks aligned with cryptic antipredator strategies, whereas gregarious locusts exhibit greater tolerance for deterrents like hyoscyamine, enabling consumption of a broader, potentially riskier range that supports swarmdynamics.[88] Gregarious nymphs on carbohydrate-biased diets accumulate more lipids but show reduced survival compared to solitarious counterparts, reflecting phase-specific physiological priorities for collective movement over individual longevity.[89]Foraging relies on multimodal sensory cues, including visual detection of green foliage and olfactory attraction to plant volatiles like those from wheat grass, which elicit approach and biting responses even in hatchlings.[90] Associative learning allows locusts to link visual or odor cues with nutrient rewards, such as protein content, enhancing efficiency in heterogeneous environments.[91] In gregarious bands, coordinated marching amplifies foraging coverage, with hoppers advancing in a front that strips vegetation systematically, while adults descend on fields during flights, prioritizing accessible, high-biomass targets.[92]
Natural enemies and population regulation
The migratory locust (Locusta migratoria) is subject to predation, parasitism, and infection by diverse natural enemies across its life stages, with over 70 species documented to attack locusts and related grasshoppers, including predatory arthropods, vertebrates, fungi, bacteria, viruses, nematodes, and protozoans.[93] These factors exert density-dependent mortality, particularly during gregarious phases when high population densities facilitate pathogen transmission and predator aggregation, though their overall role in preventing plagues is debated due to the locust's mobility and rapid reproductive capacity outpacing enemy responses in outbreak conditions.[1][31]Predators include insectivorous birds (e.g., black kites, Milvus migrans), mammals, ants, and carabid beetles, which target eggs, nymphs, and adults; for instance, birds exploit hopper bands in open habitats, but efficacy diminishes as swarms migrate beyond localized predator ranges.[94][95] Parasitoids such as dipteran flies and hymenopteran wasps attack eggs and nymphs, while mites and nematodes infest all stages, with egg pod parasitism rates reaching up to 20-30% in some field studies.[31]Pathogens predominate in regulation during outbreaks: entomopathogenic fungi like Beauveria bassiana and Metarhizium anisopliae cause epizootics in dense nymphal bands under humid conditions, achieving 80-100% mortality in lab and field trials against L. migratoria; bacteria such as Photorhabdus luminescens and protozoans like Paranosema locustae disrupt gut microbiomes and swarming behavior, suppressing phase transitions and reducing fecundity via oosorption in infected females.[96][97][98] Viruses and microsporidians further amplify crashes in high-density aggregates, where transmission rates correlate inversely with rainfall-driven dispersal.[99]Population regulation integrates these biotic pressures with abiotic cues and endogenous phase polyphenism; solitary low-density phases evade enemies through cryptic behavior and dispersal, while gregarious outbreaks trigger self-limiting factors like slower development and increased pathogen susceptibility at densities exceeding 100 individuals per square meter, often culminating in 90%+ mortality from combined enemies before recession phases.[100] Empirical models indicate natural enemies stabilize populations below outbreak thresholds in recession areas but fail to halt plagues without intervention, as evidenced by historical data showing enemy impacts confined to 10-20% of total mortality during surges.[1]
Interactions with humans
Historical plagues and documented outbreaks
The migratory locust (Locusta migratoria) has been responsible for recurrent plagues across Eurasia for millennia, with the most extensive historical documentation originating from China, where over 800 outbreaks of its oriental subspecies (L. m. manilensis) have been recorded since 707 BC.[1] These events, often linked to favorable climatic conditions such as elevated temperatures and adequate rainfall promoting breeding, devastated agricultural regions and contributed to famines, as evidenced by a 1,910-year reconstructed time series derived from dynastic chronicles showing cyclical peaks in abundance every few centuries.[101] Chinese imperial records, spanning more than 3,000 years, highlight the insect's role as a perennial threat, with officials systematically noting swarm incursions that stripped crops and prompted early control measures like communal egg-pod destruction.[102]A particularly severe episode occurred from 1855 to 1859 across northern and central China, where prolonged droughts concentrated locust populations in remnant wetlands, leading to massive gregarious formations that consumed vast expanses of wheat, millet, and rice fields, exacerbating the Taiping Rebellion-era instability and causing widespread starvation.[103] In the Russian Empire and later Soviet territories, 19th- and early 20th-century plagues ravaged steppe grasslands and Volga basin farmlands, with swarms numbering billions documented in the 1860s and 1890s, destroying up to 80% of forage in affected areas and prompting the first large-scale pesticide applications by the 1920s.[104] European incursions, though less frequent, included notable invasions in the 17th and 18th centuries, such as the 1693 outbreak in the Czech lands where hopper bands denuded fields from the Danube to the Elbe, as chronicled in regional annals attributing the event to migratory influxes from eastern breeding grounds.[105] These historical patterns underscore the species' capacity for long-distance dispersal, with swarms traveling hundreds of kilometers to exploit transient vegetation booms following environmental perturbations like floods or mild winters.[106]
Economic and agricultural damages
The gregarious phase of Locusta migratoria involves hopper bands and adult swarms that rapidly defoliate vegetation, consuming up to their body weight in plant matter daily per individual, resulting in near-total crop destruction across swaths of farmland and pasture.[1] These outbreaks strip fields bare within hours to days, leaving behind barren landscapes incapable of supporting regrowth in the short term and exacerbating soil erosion in vulnerable areas.[1] While solitary phase individuals cause negligible harm by feeding selectively on grasses, phase transitions to gregariousness amplify feeding intensity, with bands of nymphs marching in unison to devour seedlings and mature plants alike.[1]Agricultural damage targets primarily graminaceous crops such as cereals, rice, maize, and wheat, though swarms opportunistically consume vegetables, legumes, and forage, leading to widespread yield failures.[1] In regions like southern Africa, the African migratory locust subspecies (L. m. migratorioides) has impacted over 2,000 hectares of cropland and 700,000 hectares of grazing land in documented outbreaks, severely reducing productivity of green leafy and staple food crops.[107] In Asia, particularly China and Indonesia, invasions have prompted farmers to abandon rice and maize cultivation after repeated total losses from 2–3 consecutive swarm passages.[108] Less susceptible crops like sunflower, chickpea, safflower, and cotton suffer minimal defoliation due to non-preferred chemistry or structure.[109]Economically, these damages impose direct costs through lost harvests and indirect burdens via heightened food prices, livestock starvation from depleted pastures, and disrupted rural livelihoods, particularly for subsistence farmers in outbreak-prone zones of Africa, Asia, and Australia.[107] Outbreaks can result in 100% yield reduction in affected fields, forcing reliance on imports or aid and amplifying vulnerability to famine in low-resilience areas.[1] While global-scale valuations are elusive due to sporadic plagues, regional control efforts underscore the scale, with L. migratoria recognized as a transboundary pest threatening nutritional security and agribusiness viability.[110] Recovery demands replanting and soil rehabilitation, compounding financial strain on under-resourced communities.[109]
Pest control methods: chemical, biological, and integrated
Chemical control of the migratory locust primarily involves the application of broad-spectrum insecticides such as organophosphates (e.g., malathion, fenitrothion), carbamates (e.g., carbaryl), and pyrethroids, delivered via ultra-low volume (ULV) aerial or vehicle-mounted spraying to target hopper bands and swarms efficiently.[1][111][112]Insect growth regulators (IGRs) like diflubenzuron are also used, particularly in barrier treatments where treated zones intercept migrating bands, reducing environmental impact compared to blanket applications.[113] These methods achieve rapid knockdown, with efficacy rates exceeding 90% against nymphs when timed to early instars, but they pose risks to non-target organisms, including beneficial insects and aquatic life, necessitating precise application to minimize ecological disruption.[114] Emerging alternatives include enantiomerically pure formulations of chiral insecticides like isocarbophos, which enhance potency against locusts while reducing toxicity to vertebrates.[115]Biological control leverages natural enemies and microbial agents, including entomopathogenic fungi such as Beauveria bassiana, which infect locusts via cuticle penetration and cause mortality within 7-14 days under humid conditions, with field trials showing up to 80% reduction in hopper densities.[116] Bacteria like Serratia and other indigenous entomopathogenic strains have demonstrated high virulence against Locusta migratoria, achieving 90-100% mortality in lab assays when applied to early-stage nymphs.[99] Protozoan pathogens such as Nosema locustae provide slower-acting, density-dependent suppression by reducing fecundity and longevity, suitable for preventive augmentation in outbreak-prone areas.[117] Predatory birds, reptiles, and insects (e.g., carabid beetles) contribute to regulation, though their impact is limited during plagues due to locust mobility and gregarious defenses; parasitic wasps and flies offer minor augmentation potential but are challenging to scale.[118] Biopesticides are most effective against younger instars in low-light conditions, promoting their integration for sustainable suppression without broad ecological harm.[119][120]Integrated pest management (IPM) for the migratory locust combines surveillance, early intervention, and selective tactics to minimize chemical reliance, emphasizing earth observation and ground teams for detecting hopper bands before fledging, as practiced in FAO-coordinated programs since the 1990s.[121] This approach prioritizes biological agents for prophylaxis and targeted ULV insecticides for outbreaks, achieving cost savings of 30-50% over reactive chemical campaigns by focusing on source areas.[122] Area-wide strategies incorporate habitat modification, such as controlled grazing to disrupt breeding sites, alongside biopesticide releases, though challenges persist in transboundary coordination and variable efficacy under arid conditions.[123][124] Recent reviews highlight IPM's shift toward predictive modeling and reduced-risk pesticides, yet chemical tools remain dominant due to biological methods' slower action against explosive plagues.[125][121]
Challenges in management and controversies
Managing migratory locust outbreaks presents significant challenges due to the insect's phase polyphenism, which allows solitary populations to rapidly transition into gregarious swarms capable of covering vast distances, complicating early detection in remote recession areas.[126]Surveillance is hindered by inaccessible terrains, limited field personnel, and fluctuating environmental conditions that delay hopper band formation until outbreaks escalate.[127] Predictive models struggle with time-lagged effects of rainfall and vegetation on breeding sites, often failing to forecast swarm trajectories accurately across international borders.[128]Chemical control, the dominant method since the mid-20th century, faces escalating pesticide resistance in Locusta migratoria populations, particularly to organophosphates like malathion (up to 57.5-fold resistance) and chlorpyrifos, driven by prolonged exposure and enhanced detoxification enzymes such as cytochrome P450s.[129][130] Application over large areas risks non-target impacts on beneficial insects, birds, and aquatic life, with residues persisting in soil and water, exacerbating environmental degradation in already fragile ecosystems.[131][125] Biological agents like mycopesticides offer promise but suffer from slower efficacy, weather dependency, and variable field performance, limiting their scalability during rapid swarm proliferation.[132]Funding inconsistencies and political barriers further impede proactive management, as post-outbreak urgency wanes, leading to under-resourced preventive programs despite recurrent plagues.[133]International coordination falters amid differing national priorities and border disputes over swarm incursions, while climate variability—intensified by erratic rainfall—amplifies breeding opportunities, rendering historical control thresholds obsolete.[134][135]Controversies surround the balance between aggressive chemical interventions and ecological preservation, with critics arguing that broad-spectrum pesticides cause disproportionate harm to biodiversity, as seen in post-spray declines in non-target arthropods, while proponents cite their necessity to avert famine-scale crop losses.[136] Debates persist over intervention timing and extent, with some African campaigns post-2000s questioned for overuse of unapproved formulations, raising human health risks from aerial spraying near communities.[137]Resistance evolution challenges the sustainability of reliance on a narrow insecticide repertoire, prompting calls for diversified integrated pest management, though implementation lags due to technological and logistical hurdles.[138][139]
Recent outbreaks and responses (2020-2025)
In 2020, outbreaks of the African migratory locust (Locusta migratoria subsp. migratorioides) emerged across southern Africa, affecting Angola, Botswana, Namibia, Zambia, and Zimbabwe, with initial reports from breeding areas in the Okavango Delta, Chobe wetlands, and Zambezi plains starting in February and intensifying by May.[140] The infestation spread to additional Southern African Development Community (SADC) member states including Eswatini, Malawi, Mozambique, South Africa, and Tanzania, impacting irrigated crops and the main planting season amid existing food insecurity affecting 44.8 million people, 75% of whom reside in rural areas.[141] In Botswana alone, the outbreak damaged 97,598 hectares of land, exacerbating threats to livelihoods and agriculture independent of concurrent desert locust incursions elsewhere on the continent.[142][143]Responses involved coordinated regional action, including a SADC-FAO-International Locust Control Organization for Central and Southern Africa (IRLCOCSA) technical cooperation project launched on September 4, 2020, to build emergency response capacity, enhance community-based monitoring, and deploy early warning systems.[141] Control measures emphasized spot spraying with environmentally friendly pesticides, procurement of equipment, and appeals for international support, with SADC issuing a regional funding request in November 2020 to mitigate resurgence risks.[144] These efforts focused on containing hopper bands and adult groups before widespread swarming, drawing on IRLCOCSA protocols for surveillance in traditional breeding sites.[145]By 2025, renewed swarms of African migratory locust appeared in Zambia starting in March, alongside red locust (Nomadacris septemfasciata), prompting implementation of a national response plan supported by the Centre for Agriculture and Biosciences International (CABI).[146] Interventions included targeted pesticide applications, aerial and ground surveys, and farmer training to limit crop damage in affected provinces.[146] In southern Ukraine, local outbreaks of Locusta migratoria were documented in 2025, primarily involving gregarious phases in steppe regions, managed through national monitoring amid broader agricultural pressures.[147] No large-scale plagues were reported in Asia for this species during the period, though ongoing surveillance continues in historical hotspots like northern China.[148]
Utilization as human food and nutritional benefits
The migratory locust (Locusta migratoria) has been consumed by humans in various cultures for millennia, with records of traditional use as food in 65 countries across Africa, Asia, and the Middle East.[149] In 2021, the European Commission authorized dried forms of the species for humanconsumption, recognizing its potential as a novel food source following safety assessments by the European Food Safety Authority.[150][151] Consumption typically involves harvesting during outbreaks or farming under controlled conditions to avoid contaminants like pesticides from agricultural spraying.[149]Nutritionally, L. migratoria offers high protein content, ranging from 50% to 65% on a dry weight basis, comparable to conventional meats and exceeding many plant-based sources.[152][153] Specific analyses report 50.79% protein and 34.93% fat in New Zealand-sourced specimens, with fats rich in polyunsaturated fatty acids beneficial for cardiovascular health.[152][154] The species also provides essential minerals such as iron, zinc, and magnesium, alongside vitamins influenced by diet—for instance, feeding on carrot-enriched substrates elevates vitamin A levels while slightly reducing protein.[149][155] Crude fat levels average 13–29%, supporting energy needs, and the overall composition positions it as a viable source for addressing protein deficiencies in regions prone to locust plagues.[153][155]Beyond basic nutrition, studies indicate bioactive potential, including antioxidant compounds from protein fractions that may offer anti-inflammatory effects, though human clinical trials remain limited.[152][155] As a farmed insect, it presents sustainability advantages over livestock, requiring less water and land while converting plant biomass efficiently into edible protein, potentially mitigating food insecurity during swarm events.[85][156] Risks include allergenicity for shellfish-sensitive individuals due to chitin and tropomyosin, necessitating processing like drying or powdering to enhance digestibility and palatability.[151]
Related species and terminology
Distinction from other locust species
The migratory locust (Locusta migratoria) is taxonomically unique as the sole species within the monotypic genusLocusta and subfamily Oedipodinae of the Acrididae family, differing from other prominent locust species such as the desert locust (Schistocerca gregaria), which resides in the distinct genusSchistocerca and subfamily Cyrtacanthacridinae.[157][158] This separation reflects evolutionary divergences, including variations in genetic pathways related to migration and phase polyphenism, with L. migratoria showing heightened selective pressures on metabolism and mitochondrial functions compared to non-Oedipodinae locusts.[158]Morphologically, adult L. migratoria display pronounced sexual dimorphism, with females reaching 50-60 mm in length and males 40-45 mm, characterized by a robust body, elongated hind legs suited for jumping, and phase-specific coloration—greenish or brownish in solitarious individuals versus pale yellow with black dorsal markings and red-tinged hind femora in gregarious forms.[1] In comparison, S. gregaria adults are similarly sized but feature a straighter, blunter prosternal tubercle and less domed solitary-phase thorax, while the Australian plague locust (Chortoicetes terminifera) is smaller (up to 45 mm), possesses dark spots on hind wing apices, and has red shanks on hind legs absent in L. migratoria.[159][160] Nymphal instars of L. migratoria (five across both phases) exhibit a hairy prothoracic "chest" and upright head posture in lateral view, contrasting with smoother thoracic regions and downward-projecting heads in species like Aiolopus thalassinus or the less robust build of Oedaleus australis, which also lacks the migratory locust's overall larger, more sturdy frame.[161][162][39]Ecologically, L. migratoria occupies diverse habitats from temperate grasslands to tropical wetlands across Eurasia, Africa, and Australia, with regional subspecies like L. m. migratorioides enabling adaptation to varied climates, unlike the arid-desert specialization of S. gregaria or the semi-arid Australian focus of C. terminifera.[157][31] While all locusts undergo density-dependent phase shifts from solitarious to gregarious forms involving behavioral, physiological, and morphological changes, L. migratoria's polyphenism includes consistent instar counts without the occasional variations seen in some Schistocerca populations, and its swarms emphasize sustained long-distance migration over the rapid, wind-aided flights more typical of desert locust upsurges.[39][70]
Species commonly misidentified as migratory locusts
Nymphs of Aiolopus thalassinus, commonly known as the spotted grasshopper or oblique-banded grasshopper, are frequently misidentified as those of the migratory locust (Locusta migratoria), particularly when A. thalassinus populations reach high densities in overlapping regions such as northern Africa, the Middle East, and southern Europe. This confusion arises from similarities in body size, coloration (often black and tan patterns), and gregarious behavior mimicking early swarming. However, L. migratoria nymphs possess a distinctive hairy underside (thorax), which is absent in A. thalassinus; additionally, A. thalassinus exhibits more pronounced oblique bands on the forewings and a less robust build in later instars.[161]Similarly, nymphs of Oedaleus species, such as O. nigeriensis or O. senegalensis (banded grasshoppers), are sometimes mistaken for migratory locust nymphs in high-density outbreaks across Africa and parts of Asia, due to comparable dark patterning and hopping band formation. Distinguishing traits include the prominent black facial stripe and banded hind femur in Oedaleus nymphs, contrasting with the smoother face and uniform hairy thorax of L. migratoria. Adult Oedaleus also feature more elongated pronota and less migratory flight patterns compared to the strong, sustained flights of gregarious L. migratoria.[163]In regions like Australia, where L. migratoria is exotic and subject to biosecurity surveillance, local acridids such as the Australian plague locust (Chortoicetes terminifera) or spur-throated locusts (Austracris spp.) may be erroneously reported as migratory locusts during perceived incursions, owing to superficial resemblances in swarm-like aggregations and yellowish hind wings. Accurate identification relies on L. migratoria's finer thoracic pubescence, shorter antennae relative to body length, and specific genital plate morphology, which differ from the coarser features and longer antennae in Australian endemics. Misidentifications can lead to unnecessary control efforts, as documented in agricultural monitoring protocols.[164][1]Due to L. migratoria's extensive distribution across Africa, Asia, and Europe, regional confusions extend to other grasshoppers like Calliptamus italicus (Italian locust) in temperate zones, where both exhibit yellowish swarming adults; differentiation involves C. italicus's shorter wings and more jagged pronotal edges versus the fully winged, smoother profile of L. migratoria. Entomological guides emphasize genitalia examination and phase-specific traits for resolution, as superficial morphology alone often fails in field settings.[1]