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cyst of Gregarina blattarum endoparasitises intestine of Schistocerca gregaria

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sporont of Gregarina garnhami endoparasitises gut lumen of Schistocerca gregaria

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Hammerschmidtiella diesingi endoparasitises intestine of Schistocerca gregaria

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Leidynema appendiculata endoparasitises intestine of Schistocerca gregaria

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cyst of Malamoeba locustae endoparasitises rectum of Schistocerca gregaria

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larva of Mermis nigrescens endoparasitises body cavity of Schistocerca gregaria


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Evolution and Systematics

Functional Adaptations

Functional adaptation

Energy stored to amplify power: locusts

The hindlegs of grasshoppers and locusts store energy in chitonous apodemes (muscle attachment sites) in order to amplify jumping power.

  "And that's where energy storage comes in. Down to the size of a trout or a squid tentacle, unaided muscle can do a decent job with nothing more than ordinary leverage. Below that, muscle needs help; in practice, energy is put in slowly and stored elastically. Some kind of trigger then releases it at a higher rate. Work and energy may be conserved, but power gets amplified…A locust or grasshopper jumping with its hindlegs stores up work in chitinous apodemes and gets a tenfold power amplification (Bennet-Clark 1975)…Each of these creatures has some kind of a mechanical catch to prevent premature extension while the work is being put in; the specific arrangements, though, are different for each case." (Vogel 2003:476)
  Learn more about this functional adaptation.
  • Bennet-Clark, HC. 1975. The energetics of the jump of the locust Schistocerca gregaria. Journal of Experimental Biology. 63: 53-83.
  • Steven Vogel. 2003. Comparative Biomechanics: Life's Physical World. Princeton: Princeton University Press. 580 p.
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Functional adaptation

Congregating and physical stimulation trigger swarming: desert locusts

Solitary forms of desert locusts transform into those that swarm when environmental conditions force the locusts to congregate in groups, increasing physical stimulation that sets off a metabolic and behavioral cascade.

  "The desert locust is especially destructive, but like other locusts it lives a sort of Jekyll and Hyde existence. For much of the time, it is a harmless grasshopper, living a solitary life, doing no harm to agriculture, and laying eggs that hatch into fresh generations of normal grasshoppers, wingless at first but gradually developing wings as they mature.

"If environmental conditions confine these grasshoppers into concentrated breeding sites, however, a disturbing transformation occurs. The juvenile grasshoppers begin forming groups. Then, groups close to one another combine to form ever-larger congregations, and eventually an immense 'army' is produced. These gregarious yellow grasshoppers develop longer-than-normal wings, enabling them to fly farther and faster. Once airborne, they move through the skies in vast clouds, guided by the Sun and wind, seeking food and able to fly across a continent to find it." (Shuker 2001:73)

"Desert locusts, Schistocerca gregaria, show extreme phenotypic plasticity, transforming between a little-seen solitarious phase and the notorious swarming gregarious phase depending on population density. An essential tipping point in the process of swarm formation is the initial switch from strong mutual aversion in solitarious locusts to coherent group formation and greater activity in gregarious locusts. We show here that serotonin, an evolutionarily conserved mediator of neuronal plasticity, is responsible for this behavioral transformation, being both necessary if behavioral gregarization is to occur and sufficient to induce it. Our data demonstrate a neurochemical mechanism linking interactions between individuals to large-scale changes in population structure and the onset of mass migration." (Anstey et al. 2009:627)

  Learn more about this functional adaptation.
  • Anstey ML; Rogers SM; Ott SR; Burrows M; Simpson SJ. 2009. Serotonin mediates behavioral gregarization underlying swarm formation in desert locusts. Science. 323(5914): 627-630.
  • Hassanali A; Bashir MO; Njagi PGN; Ely SO. 2005. Desert locust gregarization: a conceptual kinetic model. Journal of Orthoptera Research. 14(2): 223-226.
  • Shuker, KPN. 2001. The Hidden Powers of Animals: Uncovering the Secrets of Nature. London: Marshall Editions Ltd. 240 p.
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Functional adaptation

Ovipositor drills holes: locust

Ovipositors of locusts drill deep holes by use of two reciprocating rotating elements.

  "When females of locust species such as Locusta migratoria, Schistocerca gregaria (Forskal), S.peregrina (Olivier), Anacridium aegyptium and a number of other Acrididae dig oviposition holes, they stretch the intersegmental membranes between abdominal segments IV, V, VI and VII and thus make a hole considerably deeper than could otherwise be achieved. The ovipositor valves provide the force for this extension. The ovipositor is a boring machine which, once set in motion with its prongs against the soil, must automatically bury itself and in doing so it will stretch the easily extended abdomen to its full length, so long as the insect maintains it hold on the ground. The findings of the study provide an insight to the functioning of the locust apparatus, and suggested the possibility to develop an innovative digging system composed by two reciprocating rotating elements." (Courtesy of the Biomimicry Guild)
  Learn more about this functional adaptation.
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Molecular Biology and Genetics

Molecular Biology

Barcode data: Schistocerca gregaria

The following is a representative barcode sequence, the centroid of all available sequences for this species.

There are 4 barcode sequences available from BOLD and GenBank.

Below is a sequence of the barcode region Cytochrome oxidase subunit 1 (COI or COX1) from a member of the species.

See the BOLD taxonomy browser for more complete information about this specimen and other sequences.

-- end --

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Statistics of barcoding coverage: Schistocerca gregaria

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 10
Specimens with Barcodes: 16
Species With Barcodes: 1
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Desert locust

The desert locust (Schistocerca gregaria) is a species of locust. Plagues of desert locusts have threatened agricultural production in Africa, the Middle East, and Asia for centuries. The livelihood of at least one-tenth of the world’s human population can be affected by this voracious insect.

The desert locust is potentially the most dangerous of the locust pests because of the ability of swarms to fly rapidly across great distances. It has two to five generations per year. The last major desert locust upsurge in 2004–05 caused significant crop losses in West Africa and had a negative impact on food security in the region. While the desert locust alone is not responsible for famines, it can be an important contributing factor.

Ecology and swarming[edit]

The desert locust lives a solitary life until it rains. Rain causes vegetation growth and allows the female to lay eggs in the sandy soil. The new vegetation provides food for the newly hatched locusts and provides them with shelter as they develop into winged adults.

Solitary (top) and gregarious (bottom) desert locust nymphs
Desert locusts preparatory to mating, London Zoo

When vegetation is distributed in such a way that allow the nymphs, usually called hoppers, to congregate, and there has been sufficient rain for most eggs to hatch, 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 cause the insects to transform from the solitary to the gregarious phase. When the hoppers become gregarious, their coloration changes from largely green to yellow and black, and the adults change from brown to red (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. Interestingly, 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. It is possible that this effect could aid locust control in the future.

During quiet periods, called recessions, desert locusts are confined to a 16-million-square-kilometer 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 kilometers, or approximately 20 percent 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 kilometers in a day, and will fly up to about 2,000 meters above sea level (thereafter, it becomes too cold). Therefore, swarms cannot cross tall mountain ranges such as the Atlas, the Hindu Kush or the Himalayas. They will 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 ten days during the 1987-89 plague.[1] A single swarm can cover up to 1200 square kilometers and can contain between 40 and 80 million locusts per square kilometer. The locust can live between three to six months, and there is a ten to 16-fold increase in locust numbers from one generation to the next.

Crop loss[edit]

Desert locusts can consume the approximate equivalent of their body mass each day (2 g) in green vegetation: leaves, flowers, bark, stems, fruit, and seeds. Nearly all crops, and noncrop plants, are at risk, including pearl millet, rice, maize, sorghum, sugarcane, barley, cotton, fruit trees, date palm, vegetables, rangeland grasses, acacia, pines, and banana. What is more, locust droppings are toxic, and spoil any stored food that is left uneaten.[citation needed]

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. During the twentieth century, desert locust plagues occurred in 1926-1934, 1940–1948, 1949–1963, 1967–1969, 1987–1989 and 2003-2005. In March–October 1915, a plague of locusts stripped Ottoman Palestine of almost all vegetation.[2] The significant crop loss caused by swarming desert locusts exacerbates problems of food shortage, and is a threat to food security.


Early warning and preventative 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.[3] The USSR also used locust control as a way to expand its influence in the Middle East and Central Asia.[4]

FAO's Desert Locust Information Service (DLIS) in Rome, Italy 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, and combines this information with satellite data such as MODIS, rainfall estimates and seasonal temperature and rainfall predictions 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. Those since the 1990s are available on the FAO Locust Watch web site. 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 km²) 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 (ULV) rates of application. The insecticide is acquired by the insect directly or via secondary pickup (i.e. walking over or eating the residue on a plant). Control is undertaken by government agencies in locust-affected countries or by specialized organizations such as the Desert Locust Control Organization for East Africa (DLCO-EA).

Natural enemies such as predatory and parasitic wasps and flies, predatory beetle larvae, birds, and reptiles may have limited effects on desert locusts because they can be easily overwhelmed by the sheer magnitude of most swarms and hopper bands. On the other hand, they may be effective in keeping solitary populations in check.

Farmers often try mechanical means of killing locusts, such as digging trenches and burying hopper bands, but this is very labor-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 reinfest previous fields.


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

There are two types of biopesticides - 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 like Green Muscle 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.

A biological control product has been under development since the late nineties. It is based on a naturally occurring entomopathogenic fungus (i.e. insects-infecting fungus), Metarhizium acridum. Species of Metarhizium are widespread throughout the world, infecting many groups of insects, but show a low risk to humans, other mammals and birds. The species M. acridum has specialised on short-horned grasshoppers, to which group locusts belong, and has therefore been chosen as the active ingredient of the product.

The product is available in Africa under the name Green Muscle and in Australia as Green Guard. It is applied in the same way as chemical insecticides, but does not kill as quickly. At recommended doses, the fungus can take up 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 recent trials in Algeria and Mauritania (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.

2003–2005 desert locust upsurge[edit]

Main article: 2004 locust outbreak

From October 2003 to May 2005, West Africa faced the largest desert locust outbreak in 15 years. The upsurge started as small independent outbreaks that developed in Mauritania, Mali, Niger and Sudan in the autumn of 2003. Two days of unusually heavy rains that stretched from Dakar, Senegal to Morocco in October allowed breeding conditions to remain favourable for the next six months and the desert locusts rapidly increased. Lack of rain and cold temperatures in the winter breeding area of northwest Africa in early 2005 slowed the development of the locusts and allowed the locust control agencies to stop the cycle. During the upsurge, nearly 130,000 km² were treated by ground and aerial operations in more than 20 countries. The costs of fighting this upsurge have been estimated by the FAO to have exceeded US$400 million, and harvest losses were valued at up to US$2.5 billion, which had disastrous effects on the food security situation in West Africa.

The countries affected by the 2004 outbreak were Algeria, Burkina Faso, the Canary Islands, Cape Verde, Chad, Egypt, Ethiopia, the Gambia, Greece, Guinea, Guinea Bissau, Israel, Jordan, Lebanon, Libyan Arab Jamahiriya, Mali, Mauritania, Morocco, Niger, Saudi Arabia, Senegal, Sudan, Syria and Tunisia.

In culture[edit]

Owing to the destructive habits of locusts, they have been a representation of famine in many Middle Eastern cultures, and is seen in the movies The Mummy (1999) and The Bible (1966). In the pre-oil era of the Arab states of the Persian Gulf, locusts were considered as a food delicacy.[citation needed]

Locust pheromone[edit]

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.[5]


  1. ^ "Biological control of locusts". Food and Agriculture Organization. 31 July 2006. Retrieved 29 March 2013. 
  2. ^ The Locust Plague
  3. ^ Antonio Buj, « International Experimentations and Control of the Locust Plague – Africa in the First Half of the 20th Century », in Yvon Chatelin, Christophe Bonneuil (eds.), Les sciences hors d’Occident au XXe siècle, Vol. 3 : Nature et environnement, Paris, Orstom Editions, 1995, pp. 93-105.
  4. ^ Etienne Forestier-Peyrat, "Fighting Locusts Together: Pest Control and the Birth of Soviet Development Aid, 1920-1939", ″Global Environment″, Vol. 7, No. 2, 2014, pp. 536-571
  5. ^ Nature, Pheromones: Exploitation of gut bacteria in the locust


  • AFROL News, Stronger efforts to fight West Africa's locusts Oct. 1, 2004 [1]
  • FAO Locust Watch (Desert Locust Information Service) [2]
  • Lindsey, R. 2002. Locust![3]
  • Lomer C.J., Bateman R.P., Johnson D.L., Langwald, J. and Thomas, M. 2001 Biological Control of Locusts and Grasshoppers. Annual Review of Entomology 46: 667-702.
  • OECD, The Desert Locust Outbreak in West Africa - Sept. 23, 2004 [4]
  • Programme on biological control of locusts and grasshoppers (LUBILOSA) [5]
  • Nature Magazine Article on combating desert locust through natural enemies [6]
  • Jahn, G. C. 1993. Supplementary environmental assessment of the Eritrean Locust Control Program. USAID, Washington DC. [7]

Further reading[edit]

  • Abdin, A. Stein & A. van Huis, 2001. Spatial distribution of the desert locust, Schistocerca gregaria, in the plains of the Red Sea coast of Sudan during the winter of 1999.
  • Ceccato, P., K. Cressman, A. Giannini, S. Trzaska. 2007. The desert locust upsurge in West Africa (2003–2005): Information on the desert locust early warning system and the prospects for seasonal climate forecasting. Intl J Pest Management 53(1): 7-13.
  • Cressman, K. 1996. Current methods of desert locust forecasting at FAO. Bulletin OEPP/EPPO Bulletin 26: 577-585.
  • Cressman, K. 2008. The use of new technologies in Desert Locust early warning. Outlooks on Pest Management (April, 2008): 55-59.
  • Guershon, M. & A. Ayali, 2012. Innate phase behavior in the desert locust, Schistocerca gregaria. Insect Science.
  • Symmons, P. & A. van Huis, 1997. Desert Locust Control campaign studies: operations guidebook. Wageningen University. 167 pp. & CD-Rom, 19 floppy disks.
  • Van Huis, A. 1994. Desert locust control with existing techniques: an evaluation of strategies. Proceedings of the Seminar held in Wageningen, the Netherlands, 6–11 December 1993. 132 pp. ISBN 90-6754-364-0.
  • Van Huis, A. 1995. Desert locust plagues. Endeavour, 19(3): 118-124.
  • Van Huis, A. 1997. Can we prevent desert locust plagues? In: New strategies in locust control (Eds.: S. Krall, R. Preveling and D.B. Diallo), pp. 453–459. Birkhäuser Verlag, Basel. 522 pp.
  • Van Huis, A., K. Cressman, J. Magor. 2007. Preventing desert locust plagues: optimizing management interventions. Entomologia Experimentalis et Applicata 122: 191-214.
  • Werf, W. van der, G. Woldewahid, T. Abate, M. Butrous, O. Abdalla, A.M. Khidir, B. Mustafa, I. Magzoub, O.
  • Vallebona C, Genesio L, Crisci A, Pasqui M, Di Vecchia A, Maracchi G (2008). Large-scale climatic patterns forcing desert locust upsurges in West Africa. CLIMATE RESEARCH (2008) 37:35-41. ISSN: 1616-1572.
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