Desert locust

A gregarious phase adult

The genus Schistocerca consists of more than 30 species, distributed in Africa, Asia, and North and South America, and many species are difficult to identify due to the presence of variable morphs. It is the only genus within the Cyrtacanthacridinae that occurs in both the New and Old World. Most species have the fastigium deflexed and lack lateral carinae on the pronotum. The hind tibiae have smooth margins with numerous spines, but have no apical spine on the outer margin. The second tarsal segment is half as long as the first. Males in the genus have broad anal cerci and a split subgenital plate. The genus is thought to have originated in Africa and then speciated in the New World after a dispersal event that took place 6 to 7 million years ago.[8][9][10]

The solitary phase insects (or solitaria morph) are greenish in early instars, while older adults are greyish. The solitary phase grasshoppers lack black patterning as present in the gregarious (or gregaria morph) phase, where the body is pinkish in early instars and yellow in adults.[11]

The second and third pairs of spiracles are permanently open.[12]

The lifecycle of the desert locust consists of three stages, the egg, the nymph known as a hopper, and the winged adult. Copulation takes place when a mature male hops onto the back of a mature female and grips her body with his legs. Sperm is transferred from the tip of his abdomen to the tip of hers, where it is stored. The process takes several hours and one insemination is sufficient for a number of batches of eggs.[13]

The female locust then seeks suitable soft soil in which to lay her eggs. It needs to be the right temperature and degree of dampness and be in close proximity to other egg-laying females. She probes the soil with her abdomen and digs a hole into which an egg pod containing up to 100 eggs is deposited. The egg pod is 3 to 4 cm (1+1⁄8 to 1+5⁄8 in) long and the lower end is about 10 cm (4 in) below the surface of the ground. The eggs are surrounded by foam and this hardens into a membrane and plugs the hole above the egg pod. The eggs absorb moisture from the surrounding soil. The incubation period before the eggs hatch may be two weeks, or much longer, depending on the temperature.[13]

Dense hopper band in Sudan

The newly hatched nymph soon begins to feed, and if it is a gregarious individual, is attracted to other hoppers and they group together. As it grows, it needs to moult (shed its exoskeleton). Its hard cuticle splits and its body expands, while the new exoskeleton is still soft. The stages between moulting are called instars and the desert locust nymph undergoes five moults before becoming a winged adult. Immature and mature individuals in the gregarious phase form bands that feed, bask, and move as cohesive units, while solitary-phase individuals do not seek conspecifics.[13][14]

After the fifth moult, the insect is not yet mature. It is at first soft and pink with drooping wings, but over the course of a few days, the cuticle hardens and haemolymph is pumped into the wings, which stiffens them. Maturation can occur in 2–4 weeks when the food supply and weather conditions are suitable but may take as long as 6 months when they are less ideal. Males start maturing first and give off an odour that stimulates maturation in the females. On maturing, the insects turn yellow and the abdomens of the females start swelling with developing eggs.[13]

Solitary (top) and gregarious (bottom) desert locust nymphs

„Schwarm der Wanderheuschrecke“, Chromolithographie nach Emil Schmidt (Brehm, A.E., 1882–1884, Bd. 9: 551)

Desert locusts have a solitary phase and a gregarious phase, a type of polyphenism. Solitary locusts nymphs and adults can behave gregariously within a few hours of being placed in a crowded situation, while gregarious locusts need one or more generations to become solitary when reared in isolation.[14] Differences in morphology and behaviour are seen between the two phases. In the solitary phase, the hoppers do not group together into bands but move about independently. Their colouring in the later instars tends to be greenish or brownish to match the colour of their surrounding vegetation. The adults fly at night and are also coloured so as to blend into their surroundings, the immature adults being grey or beige and the mature adults being a pale yellowish colour. In the gregarious phase, the hoppers bunch together and in the later instars develop a bold colouring with black markings on a yellow background. The immatures are pink and the mature adults are bright yellow and fly during the day in dense swarms.[13]

Desert locusts preparatory to mating, London Zoo

The change from an innocuous solitary insect to a voracious gregarious one normally follows a period of drought, when rain falls and vegetation flushes occur in major desert locust breeding locations. The population builds up rapidly and the competition for food increases.[15] As hoppers get more crowded, the close physical contact causes the insects' hind legs to bump against one another. This stimulus triggers a cascade of metabolic and behavioral changes that causes the insects to transform from the solitary to the gregarious phase. When the hoppers become gregarious, their colouration changes from largely green to yellow and black, and the adults change from brown to pink (immature) or yellow (mature). Their bodies become shorter, and they give off a pheromone that causes them to be attracted to each other, enhancing hopper band and subsequently swarm formation. The nymphal pheromone is different from the adult one. When exposed to the adult pheromone, hoppers become confused and disoriented, because they can apparently no longer "smell" each other, though the visual and tactile stimuli remain. After a few days, the hopper bands disintegrate and those that escape predation become solitary again. This effect could aid locust control in the future.

During quiet periods, called recessions, desert locusts are confined to a 16-million-square-kilometre (6.2-million-square-mile) belt that extends from Mauritania through the Sahara Desert in northern Africa, across the Arabian Peninsula, and into northwest India. Under optimal ecological and climatic conditions, several successive generations can occur, causing swarms to form and invade countries on all sides of the recession area, as far north as Spain and Russia, as far south as Nigeria and Kenya, and as far east as India and southwest Asia. As many as 60 countries can be affected within an area of 32 million square kilometres (12 million square miles), or about 20% of the Earth's land surface.

Locust swarms fly with the wind at roughly the speed of the wind. They can cover from 100 to 200 km (62 to 124 mi) in a day, and fly up to about 2,000 metres (6,600 ft) above sea level (the temperature becomes too cold at higher altitudes). Therefore, swarms cannot cross tall mountain ranges such as the Atlas, the Hindu Kush, or the Himalayas. They do not venture into the rain forests of Africa nor into central Europe. However, locust adults and swarms regularly cross the Red Sea between Africa and the Arabian Peninsula, and are even reported to have crossed the Atlantic Ocean from Africa to the Caribbean in 10 days during the 1987-89 plague.[16] A single swarm can cover up to 1,200 square kilometres (460 sq mi) and can contain between 40 and 80 million locusts per square kilometre (100,000,000 and 210,000,000 per square mile) (a total of around 50 to 100 billion locusts per swarm, representing 100,000 to 200,000 metric tons (98,000 to 197,000 long tons; 110,000 to 220,000 short tons), considering an average mass of 2 g per locust). The locust can live between 3 and 6 months, and a 10- to 16-fold increase in locust numbers occurs from one generation to the next.

Desert locusts consume an estimated equivalent of their body weight (2 g (0.07 oz)) each day in green vegetation. They are polyphagous and feed on leaves, shoots, flowers, fruit, seeds, stems, and bark. Nearly all crops and noncrop plants are eaten, including pearl millet, maize, sorghum, barley, rice, pasture grasses, sugarcane, cotton, fruit trees, date palms, banana plants, vegetables, and weeds.[15]

Locusts feeding

Crop loss from locusts was noted in the Bible and Qur'an; these insects have been documented as contributing to the severity of a number of Ethiopian famines. Since the early 20th century, desert locust plagues occurred in 1926–1934, 1940–1948, 1949–1963, 1967–1969, 1987–1989, 2003–2005, and 2019–2020.[17] In March–October 1915, a plague of locusts stripped Ottoman Palestine of almost all vegetation.[18] The significant crop loss caused by swarming desert locusts exacerbates problems of food shortage, and is a threat to food security.

Early warning and preventive control is the strategy adopted by locust-affected countries in Africa and Asia to try to stop locust plagues from developing and spreading. In the 1920s-1930s, locust control became a major field for international cooperation. The International Agricultural Institute developed several programmes aimed at exchanging data about the desert locust and international conferences were held in the 1930s: Rome in 1931, Paris in 1932, London in 1934, Cairo in 1936, and Brussels in 1938. Colonial empires were heavily involved in these attempts to control locust pests, which affected heavily the Middle East and parts of Africa.[19] The USSR also used locust control as a way to expand its influence in the Middle East and Central Asia.[20]

FAO's Desert Locust Information Service (DLIS) in Rome monitors the weather, ecological conditions, and the locust situation on a daily basis. DLIS receives results of survey and control operations carried out by national teams in affected countries. The teams use a variety of innovative digital devices, eLocust3, to collect, record and transmit standardized data in real-time to their national locust centres for decision-making. This data is automatically integrated into SWARMS, the global monitoring and early warning system operated by DLIS. Within this system, the field data are combined with the latest satellite imagery to actively monitor rainfall, vegetation and soil moisture conditions in the locust breeding area from West Africa to India. This is supplemented by sub-seasonal and seasonal temperature and rainfall predictions up to six months in advance as well as other weather forecasts and data from NOAA and ECMWF. Models are used to estimate egg and hopper development rates and swarm trajectories (NOAA HYSPLIT) and dispersion (UK Met Office NAME). DLIS uses a custom GIS to analyze the field data, satellite imagery, weather predictions and model results to assess the current situation and forecast the timing, scale, and location of breeding and migration up to six weeks in advance. The situation assessments and forecasts are published in monthly locust bulletins that date back to the 1970s. These are supplemented by warnings and alerts to affected countries and the international community. This information is available on the FAO Locust Watch website. DLIS continuously adopts the latest technologies as innovative tools, including drones, to improve monitoring and early warning. FAO also provides information and training to affected countries and coordinates funding from donor agencies in case of major upsurges and plagues.

The desert locust is a difficult pest to control, and control measures are further compounded by the large and often remote areas (16–30 million square kilometres (6.2–11.6 million square miles)) where locusts can be found. Undeveloped basic infrastructure in some affected countries, limited resources for locust monitoring and control, and political turmoil within and between affected countries further reduce the capacity of a country to undertake the necessary monitoring and control activities.

At present, the primary method of controlling desert locust infestations is with insecticides applied in small, concentrated doses by vehicle-mounted and aerial sprayers at ultra-low volume rates of application. The insecticide is acquired by the insect directly, meaning that control must be precise. Control is undertaken by government agencies in locust-affected countries or by specialized regional aerial organizations such as the Desert Locust Control Organization for East Africa (DLCO-EA).

The desert locust has natural enemies such as predatory wasps and flies, parasitoid wasps, predatory beetle larvae, birds, and reptiles. These may be effective at keeping solitary populations in check but are of limited effects against gregarious desert locusts because of the enormous numbers of insects in the swarms and hopper bands.[15]

Farmers often try mechanical means of killing locusts, such as digging trenches and burying hopper bands, but this is very labour-intensive and is difficult to undertake when large infestations are scattered over a wide area. Farmers also try to scare locust swarms away from their fields by making noise, burning tires, or other methods. This tends to shift the problem to neighbouring farms, and locust swarms can easily return to reinfest previously visited fields.

Biopesticides include fungi, bacteria, neem extract, and pheromones. The effectiveness of many biopesticides equals that of conventional chemical pesticides, but two distinct differences exist. Biopesticides in general take longer to kill insects, plant diseases, or weeds, usually between 2 and 10 days.

The two types of biopesticides are biochemical and microbial. Biochemical pesticides are similar to naturally occurring chemicals and are nontoxic, such as insect pheromones used to locate mates, while microbial biopesticides, come from bacteria, fungi, algae, or viruses that either occur naturally or are genetically altered. Entomopathogenic fungi generally suppress pests by mycosis - causing a disease that is specific to the insect.

Biological control products have been under development since the late 1990s; Green Muscle and NOVACRID are based on a naturally occurring entomopathogenic fungus, Metarhizium acridum. Species of Metarhizium are widespread throughout the world, infecting many groups of insects, but show low risk to humans, other mammals, and birds. The species M. acridum has specialised in short-horned grasshoppers, to which these locusts belong, so has been chosen as the active ingredient of the product.

The product is available in Australia under the name Green Guard and in Africa, it used to be available as Green Muscle. However, since Green Muscle seems to have disappeared from the market, another product, NOVACRID, was developed for Africa, Central Asia, and the Middle East. These products are applied in the same way as chemical insecticides, but do not kill as quickly. At recommended doses, the fungus can take up to two weeks to kill up to 90% of the locusts. For that reason, it is recommended for use mainly against hoppers, the wingless early stages of locusts. These are mostly found in the desert, far from cropping areas, where the delay in death does not result in damage. The advantage of the product is that it affects only grasshoppers and locusts, which makes it much safer than chemical insecticides. Specifically, it allows the natural enemies of locusts and grasshoppers to continue their beneficial work. These include birds, parasitoid and predatory wasps, parasitoid flies, and certain species of beetles. Though natural enemies cannot prevent plagues, they can limit the frequency of outbreaks and contribute to their control. Biopesticides are also safer to use in environmentally sensitive areas such as national parks or near rivers and other water bodies.

Green Muscle was developed under the LUBILOSA programme, which was initiated in 1989 in response to environmental concerns over the heavy use of chemical insecticides to control locusts and grasshoppers during the 1987-89 plague. The project focused on the use of beneficial disease-causing microorganisms (pathogens) as biological control agents for grasshoppers and locusts. These insects were considered to be too mobile and to reproduce too fast to be readily controlled by classical biological control. Pathogens have the advantage that many can be produced in artificial culture in large quantities and be used with ordinary spraying equipment. Entomopathogenic fungi were traditionally seen as needing humid conditions to work well. However, the LUBILOSA programme found a way to avoid this by spraying fungal spores in an oil formulation. Even under desert conditions, Green Muscle can be used to kill locusts and other acridid pests, such as the Senegalese grasshopper. During trials in Algeria and Mauritania in 2005 and 2006, various natural enemies, but especially birds, were abundant enough to eliminate treated hopper bands in about a week, because the diseased hoppers became sluggish and easy to catch.

In the 1900s, there were six major desert locust plagues, one of which last almost 13 years.

The swarming pheromone guaiacol is produced in the gut of desert locusts by the breakdown of plant material. This process is undertaken by the gut bacterium Pantoea (Enterobacter) agglomerans. Guaiacol is one of the main components of the pheromones that cause locust swarming.[25] Pheromones also accelerate S. gregaria development.[26] Mahamat et al., 1993 find that an undifferentiated mix of several volatiles derived from the males of the species (including guaiacol) speed up the maturation process of both immature males and females.[26]

S. gregaria was one of the organisms examined by McNeill and Hoyle 1967 and found to have thinner muscle filaments than those before found. This contributed greatly to the development of the sliding filament theory.[27]

Westerman[28] showed that exposure of S. gregaria males to a dose of X-rays during the S-phase (DNA synthesis phase) of spematogonial mitoses and during the early stages of meiosis (leptotene-early zygotene stages) caused a significant increase in chiasmata frequency when scored at the later stages (diplotene-diakinesis stages) of meiosis. These results indicated that the formation of chiasmata is not an isolated event but the end product of an interrelated series of processes initiated at some earlier stage of meiosis.[28]

Given the long history of desert locust, it is to be expected that references of the world's most dangerous migratory pest have crept into popular film and literature as well as many of the world's religions.

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Wikimedia Commons has media related to Desert locust.

• Desert Locust crisis in the Horn of Africa - FAO Website Archived 2021-08-18 at the Wayback Machine
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• Columbia University IRI Climate and Desert Locust Archived 2012-12-21 at the Wayback Machine
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• Modelling insect wings using the finite element method