Migratory locust (Locusta migratoria)

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Migratory locust (Locusta migratoria)
Migratory locust
Distribution
Afghanistan, Uzbekistan, Georgia (country), Armenia, Tajikistan, Russian Federation, Kyrgyzstan, Turkmenistan, Kazakhstan, Azerbaijan, Madagascar, India, Malawi, Mozambique, South Africa, Zambia, Namibia, Botswana, Angola, Zimbabwe, Australia, New Zealand, Pakistan, Philippines, Cambodia, Thailand, Indonesia, Malaysia, China, Romania, Viet Nam, Tanzania, Laos, Eswatini, Turkey
Taxonomy
Family:Acrididae
Subfamily:Oedipodinae
Genus:Locusta
Additional resources
Full taxonomy from OSF

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The Migratory locust (Locusta migratoria) is a notorious agricultural pest and the most widely distributed grasshopper species in the world[1]. It is present across the entire temperate and tropical Eastern Hemisphere. In China, records of Migratory Locust plagues go back to 200 BC. Today it is one of the most important agricultural pests in Russia, Kazakhstan, and Uzbekistan.

Nomenclature

Locusta migratoria (Linnaeus, 1758). For full nomenclature, see this taxon's page on Orthoptera Species File.


Recent finding shows only two true subspecies: Asian migratory locust, L. migratoria migratoria (Linnaeus, 1758) and African migratory locust, L. migratoria migratorioides (Reiche and Fairmaire, 1849) [2]

Identification

Distinctive for its large size, absence of prosternal tubercule, and smoky colored wings at the apex (radial sector) and black veins in the anal sector. In profile, the pronotum is high for solitary adults and saddle-shaped for gregarious. The transversal furrow is well marked for gregarious individuals. [3]
Unique features: Dark mouth, hairy underside of thorax. Its large size, clear rear wing, and raised thorax profile distinguish it from the Australian plague locust. Clear rear wing, lighter tibia, strongly marked forewings (compared to mottled), and flatter thorax profile, distinguish it from similar yellow-winged locust.[4] Compare photos

Phase Stage Color Wings Legs
Gregarious nymph Orange and black in later instars. Dark stripe from antenna through eye at a 45 degree angle, becomes horizontal before fading into the thorax, which has a small but prominent dark spot. Two oblique dark bands on rear femur.
Gregarious immature adult
Gregarious mature adult bluish body with a yellow head and legs Folded forewings are dark with thin light markings. Hind wings are clear, smoky at the apex (radial sector) and distinct black anal veins [1]
Solitarious nymph Green but grey in the 1st instar. Dark stripe from antenna through eye at a 45 degree angle, becomes horizontal before fading into the thorax, which has a small but prominent dark spot. Two oblique dark bands on rear femur.
Solitarious immature adult
Solitarious adult usually brown or green (sometimes yellowish-green or grey) Folded forewings are dark with thin light markings. Hind wings are clear, smoky at the apex (radial sector) and distinct black anal veins [1]

Additional information
Male body length:  varying from 35 to 50 mm
Female body length: 45 to 55 mm
Head (in profile): slightly low​​er than the thorax but both head and thorax are raised above the wing line.
Mandibles: blue, dark mouth
Elytra (the outer tough pair of wings) are reflective and long: 43.5-56.0 (males), 49.0-61.0 mm (females), clearly protruding past the abdominal area [3]
Hind femora color: brownish, bluish to black on bottom internal side [3]
Hind femur length: 22.0-26.0 mm (males) and 20.0-32.0 mm (females). Relatively long in solitarious individuals [3]
Hind tibia color: yellowish, beige or red [3]

Identification resources

Title Link Descriptive keyword Language Geographic purview Author Year
Agriculture Victoria Australian plague locust identification, biology and behaviour View URL Species identification, Biology, Behavior English Australia Agriculture Victoria 2022
APLC locust and grasshopper identification guide View URL Management, Species identification English Australia Australian Plague Locust Commission
Australian plague locust online learning module View URL Species identification, Biology, Behavior, Management English Australia Agriculture Victoria 2022
FAO locust handbook identification key View URL Species identification, Locusts English FAO Desert Locust Information Service, Food and Agriculture Organization of the United Nations
Locust identification in Niger View URL Management, Species identification, Agriculturist guide French, Zarma, Hausa Niger Global Locust Initiative, United States Agency for International Development, Directorate of Plant Protection, United Nations AGRHYMET Regional Center, Centre National de Lutte Anti-Acridienne, National Network of Agriculture Chambers of Niger 2022
Locusts of Australia View URL Management, Species identification English Australia Queensland Department of Agriculture and Fisheries 2020
Orthoptera Species File View URL Species identification, Biology English Africa, Americas, Asia, Europe, Oceania Orthoptera Species File Online
Pest locusts View URL History, Archive, Species identification French, English Africa, Americas, Asia, Europe, Oceania The French Agricultural Research Centre for International Development



Distribution

The migratory locust has the largest global distribution of all locusts and grasshoppers. It is present in all temperate and tropical regions of the Eastern hemisphere. The southern edge of the coniferous forest of Eurasia marks its northern limit and it extends as far as South as New Zealand. It can be found from the Azores in the West to Fiji in the East. [1]

Their presence in a variety of climates and environmental conditions resulted in the assumption of many subspecies. However, Ma et al. (2012) conducted a phylogeographic analysis and divided populations into just two lineages: the Northern lineage of the Eurasian continent Asian migratory locust, L. migratoria migratoria (Linnaeus, 1758) and the Southern lineage, African migratory locust, L. migratoria migratorioides (Reiche and Fairmaire, 1849), spanning Africa, southern Europe, the Arabian region, India, southern China, South-east Asia and Australia. L. m. migratorioides is distributed in Africa south of the Sahara and the offshore Atlantic islands. For more information and distribution records see [GBIF]

In China, records of the migratory locust outbreaks go back to 200 BC and are associated with droughts and flood events. [5] The Asian migratory locust is one of the most important agricultural pests in Russia, Kazhakhstan, and Uzbekistan. [6] Populations in Australia are more recent, starting in 1973, when an important outbreak occurred in the Central Highlands district of Queensland. Before then it was an uncommon species of the coastal and subcoastal region with a few scattered records from inland. [7]

For more information and distribution records see [GBIF]

Biology

Habitat and ecology

L. migratoria are common in areas of periodic flooding such as river deltas, coastal plains, and along lakes where they breed in wet grassland areas on light soils. [6] [8] Wet seasons (or years) following dry seasons can lead to population build up and the aggregation of locusts on drying soil or grasses. This mosaic of patchy vegetation has been attributed to the onset of gregarization. [6]

The Food and Agriculture Organization (FAO) lists permanent habitat areas of L. m. migratoria in the Russian Federation, Kazakhstan, and China with specific breeding hot spots in Balkhash-Alakol lakes, Amu Darya river and, more recently Northern Caspian and Dagestan regions.

The migratory locust L. m. migratoria is univoltine (having one brood or generation per year) but can have up to five generations per year in tropical zones under favorable conditions. Oviposition occurs in August-September and the eggs over winter. Each female can produce at least 1-3 egg pods (up to 5 in southern localities and with warm conditions) with an average of 60-80 eggs per pod.[3] Light sandy soils are preferred for egg laying. Egg pod is large, slightly bent, 50-85 mm in length, 7-10 mm in diameter; eggs are 7-8 mm long.[3] Hatching (completed in 4-5 days) takes place between early May and early June. There are 5 hopper instars, 7-8 days for each, and the total hopper development lasts 35-40 days. Adults emerge in June to early July, in warm areas they can remain as late as November. Mating starts 2-4 weeks after fledging, and females start laying eggs 2-3 weeks later (usually at the end of July). [3]

Almost all sub-Saharan countries are in the invasion zone of the African migratory locust (L. m. migratorioides) outbreaks. [3] The Niger River flood plains in Mali are the main African breeding area and to a lesser extent, Lake Chad basin, and the Blue Nile region in Sudan. [9] Four to five generations annually are possible in the Middle Niger and can form plagues in the area and surrounding Sahelian steppes. [1] The African migratory locust favors grasses Ctenium elegans, Cenchrus biflorus, Eragrostis tremula and 10–15% bare ground for breeding in the late rainy season. [8]

Gregarization is a slower process for the migratory locust relative to the desert locust. After 64 hours of crowding these locusts are only partially gregarized. [10] [11] [12]

  • Exhibits phase polymorphism and long-distance migration [13] [14] [15]

Land-use change

Historically, the migratory locust was not present in the drylands of the Sahara Desert however, agricultural expansion and human induced vegetation changes, particularly through the introduction of irrigation, has made for a more hospitable habitat and increased populations in this area. [16] [17]

Population upsurges have been seen in new areas due to rapid agricultural development of expansive cereal summer crops.[8] In the Sahel, shifting agricultural land has left plots abandoned with bare areas that still support millet, sorghum, and Cenchrus biflorus which are favored host plants. [8]

Outbreaks have been reported in Mali, Chad, Sudan and Madagascar.[18]

Pest status

Polyphagous but mainly graminivorous, preferring grasses such as Panicum sp. Phragmites communis, Artemisia sp. and Polygonum [8] Agricultural damages usually concern cereals (barley, millet, corn, maize, oats, rice, sugar cane, teff grass, wheat, also bamboo and sugar cane. Other plants when grasses are not available, like banana, pineapple leaves, and palms. [1]

Recent outbreaks

In January - June 2021, an outbreak of migratory locusts infested more than 48,000 hectares of land in the Grand Sud, Madagascar. [19]

In 2020, Southern African countries launched an emergency response to outbreaks of the African migratory locust which, according to the Food and Agriculture Organization of the United Nations (FAO), threatened the food security and livelihoods of millions of people in Botswana, Namibia, Zambia, and Zimbabwe.[20]

Southern African Development Community SADC Regional Appeal to contain the African Migratory Locust October 2020

ZAMBIA: African Migratory Locust FAO FLASH UPDATE – 26 October 2020

Madagascar 2014 - View photos of the outbreak here https://www.flickr.com/photos/faoemergencies/albums/72157644348724287/with/14228170412/

Media coverage

2012 Madagascar locust crisis [21]

Title Link Descriptive keyword Language Geographic purview Author Year
"Biblical plague" of locusts ruins crops in Tulcea View URL Media article, Locust outbreaks English Romania Romania Insider 2022
Besieged by locusts, South Africa's farmers fight back View URL Media article, Locust outbreaks English BBC News
Explainer: what's behind the locust swarms damaging crops in southern Africa View URL Media article, Locust outbreaks, Locusts, Outbreaks, Infestations, Rainfall, Weather, Climate Change, Agriculture, Ecology, Biology, Behavior, Education, General locust education, Monitoring, Chemical control English The Conversation 2020
Locust outbreaks threaten food security in southern Africa View URL Press release, Media article, Newsletter, Locust outbreaks, Locusts, Outbreaks, Infestations, Crop health, Food security, Livelihoods, Emergency response, Community development, Management English Food and Agriculture Organization of the United Nations
Locust Plague Devours 40,000 Tonnes of Vegetation View URL General locust education, Video English Madagascar BBC News 2022
Locusts now threatening parts of southern Africa, UN says View URL Media article, Locust outbreaks, Outbreaks, Locusts, Infestations, Crop health, Agriculture, Pesticides, Spraying, Chemical control, Environmental assessment, Management, Integrated pest management English phys.org 2020
On the frontlines: Battling Namibia’s worst locust crisis View URL Newsletter, Media article, Locust outbreaks, Locusts, Outbreaks, Infestations, Livelihoods, Food security, Crop health, Integrated pest management, Chemical control, Spraying, Governance, Community development, Collective movement, Grazing, Livestock English Namibia United Nations 2021
Organisation des Nations Unies pour l'alimentation et l'agriculture View URL Newsletter, Outbreaks, Locusts, Locust outbreaks, Emergency response, Management, Integrated pest management, Advocacy, Advisory note, Food security, Livelihoods, Agriculture, Rainfall, Crop health French Food and Agriculture Organization of the United Nations
Villagers In Sumba Chase Away Migratory Locusts With Loud Noises View URL Video, Locust outbreaks, Locusts, Outbreaks, Infestations, Management, Mechanical control, Spraying, Pesticides, Chemical control, Governance, Agriculture, Crop health English SEA Today News 2022



Organizations associated with the migratory locust

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Resources

Title Link Descriptive keyword Language Geographic purview Author Year
2014 evaluation of field trials data on the efficacy and selectivity of insecticides on locusts and grasshoppers View URL Pesticides, Management, Pesticide management, Chemical control English Commission for Controlling the Desert Locust in the Central Region, Food and Agriculture Organization of the United Nations 2014
2021 evaluation of field trials data on the efficacy and selectivity of insecticides on locusts and grasshoppers View URL Report, Pesticides, Pesticide management, Management, Monitoring, Public health English Africa, Europe, Asia, Americas, Oceania Locust Pesticide Referee Group, Food and Agriculture Organization of the United Nations 2021
Agriculture Victoria locust and grasshopper online reporting form View URL Report English Australia Agriculture Victoria
Australian Plague Locust Commission current locust situation View URL Locusts, Locust outbreaks, Outbreaks, Infestations, Distribution, Monitoring, Forecasting, Information hub, Weather, Rainfall English Australia Australian Plague Locust Commission
Australian Plague Locust Commission locust bulletin View URL Locusts, Monitoring, Migration, Forecasting, Advisory note, Weather, Rainfall, Behavior, Survey, Distribution, Map English Australia Australian Plague Locust Commission
CABI BioSpace project View URL Crop health, Invasive species, Forecasting, Meteorology, Remote sensing, Early warning English China, Laos Centre for Agriculture and Bioscience International 2020
CABI Green Muscle education videos View URL Video, Biological control, Biopesticide, Metarhizium acridum, Storage, Dosage, Application, PPE and cleaning Arabic, French, Russian, English Africa, Americas, Asia, Europe, Oceania 2021
Calendars 2023 on locust control safety measures Calendar, Management Azerbaijani, Armenian, Georgian Azerbaijan, Armenia, Georgia (country) FAO Locust Watch Locusts in Caucasus and Central Asia, Food and Agriculture Organization of the United Nations 2023
CCA locust bulletins Forecasting, Report, Locusts, Locust outbreaks English Kazakhstan, Kyrgyzstan, Tajikistan, Uzbekistan, Turkmenistan, Afghanistan, Georgia (country), Russian Federation, Armenia, Azerbaijan FAO Locust Watch Locusts in Caucasus and Central Asia, Food and Agriculture Organization of the United Nations
Control of the locust plague in Madagascar 2014 View URL Video, Management French Madagascar Food and Agriculture Organization of the United Nations 2014
FAO Locust Watch bulletin for Locusts in Caucasus and Central Asia View URL Forecasting, Information hub, Emergency alerts English Afghanistan, Georgia (country), Russian Federation, Uzbekistan, Armenia, Kazakhstan, Tajikistan, Azerbaijan, Kyrgyzstan, Turkmenistan FAO Locust Watch Locusts in Caucasus and Central Asia, Food and Agriculture Organization of the United Nations
India Locust Warning Bulletin View URL Early warning English India Indian Council of Agricultural Research
Locust Literature View URL Archive French, English Africa, Americas, Asia, Europe, Oceania The French Agricultural Research Centre for International Development
Locusts in Queensland View URL Taxonomy, History, Distribution, Biology, Ecology, Management English Australia Queensland Government 2003
Practical Guidelines on Pesticide Risk Reduction for Locust Control in CCA View URL Pesticides, Pesticide management, Management English, Georgian, Kyrgyz, Dari, Tajik, Uzbek, Azerbaijani Azerbaijan, Georgia (country), Kyrgyzstan, Tajikistan, Uzbekistan, Turkmenistan FAO Locust Watch Locusts in Caucasus and Central Asia, Food and Agriculture Organization of the United Nations 2019
Report Technical Workshop on Locusts In Caucasus and Central Asia View URL Report, Monitoring, Public health, Forecasting, Management English Afghanistan, Armenia, Azerbaijan, Georgia (country), Kazakhstan, Kyrgyzstan, Russian Federation, Tajikistan, Turkmenistan, Uzbekistan FAO Locust Watch Locusts in Caucasus and Central Asia, Food and Agriculture Organization of the United Nations 2022
Results of the three-year programme in response to the locust plague in Madagascar 1 View URL Video, Report English, French Madagascar Food and Agriculture Organization of the United Nations 2016
Southern Africa locust outbreak September 2020 View URL Summary, Locust outbreaks, Locusts, Outbreaks, Agriculture, Food security, Management, Integrated pest management, Pesticide management, Economics, Environmental assessment, Emergency response, Pesticides, Chemical control, Forecasting, Monitoring, Rainfall, Weather, Meteorology English 2020
Sustainable use and conservation of microbial and invertebrate biological control agents and microbial biostimulants View URL Management, Biological control, Biopesticide English Africa, Americas, Asia, Europe, Oceania Centre for Agriculture and Bioscience International 2023
USAID Transboundary Outbreak Pest ETOP bulletins View URL Information hub English Sub-Saharan Africa, Latin America and the Caribbean United States Agency for International Development


References

  1. 1.0 1.1 1.2 1.3 1.4 1.5 COPR (1982). The Locust and Grasshopper Agricultural Manual. London: Overseas Pest Research. 449.
  2. Ma C, Yang P, Jiang F, Chapuis M-P, Shali Y, Sword GA, Kang L (2012) Mitochondrial genomes reveal the global phylogeography and dispersal routes of the migratory locust: GLOBAL PHYLOGEOGRAPHY OF THE MIGRATORY LOCUST. Molecular Ecology 21: 4344–4358. https://doi.org/10.1111/j.1365-294X.2012.05684.x
  3. 3.0 3.1 3.2 3.3 3.4 3.5 3.6 3.7 3.8 FAO (2019) Asian Migratory Locust (LMI). Asian Migratory Locust (LMI) | Locust Watch in Caucasus and Central Asia. Available from: http://www.fao.org/locusts-cca/bioecology/asian-migratory-locust-lmi/en/ (November 18, 2019).
  4. APLC https://www.agriculture.gov.au/pests-diseases-weeds/locusts/about/id-guide/description_of_adults/3_migratory_locust_locusta_migratoria (accessed April 30, 2021)
  5. Stige LC, Chan K-S, Zhang Z, Frank D, Stenseth NC (2007) Thousand-year-long Chinese time series reveals climatic forcing of decadal locust dynamics. Proceedings of the National Academy of Sciences U.S.A. 104, 16188–16193. doi:10.1073/pnas.0706813104
  6. 6.0 6.1 6.2 Latchininsky AV (2013) Locusts and remote sensing: a review. Journal of Applied Remote Sensing 7: 075099. https://doi.org/10.1117/1.JRS.7.075099
  7. Farrow R A (1979) Causes of Recent Changes in the Distribution and Abundance of the Migratory Locust (Locusta migratoria L.) in Australia in Relation to Plagues.
  8. 8.0 8.1 8.2 8.3 8.4 Le Gall M, Overson R, Cease A (2019) A global review on locusts (Orthoptera: Acrididae) and their interactions with livestock grazing practices. Frontiers in Ecology and Evolution 7: 263. https://doi.org/10.3389/fevo.2019.00263
  9. Balança G, Gay P-E, Rachadi T, Lecoq M, Balanca G (1999a) Interpretation of Recent Outbreaks of the Migratory Locust Locusta migratoria migratorioides (Reiche and Fairmaire, 1850) [Orthoptera, Acrididae] in Lake Chad Basin According to Rainfall Data. Journal of Orthoptera Research: 83. https://doi.org/10.2307/3503430
  10. Guo W, Wang X, Ma Z, Xue L, Han J, Yu D (2011) CSP and takeout genes modulate the switch between attraction and repulsion during behavioral phase change in the migratory locust. PLoS Genet. 7:e1001291. doi:10.1371/journal.pgen.1001291
  11. Ma Z, Guo W, Guo X, Wang X, Kang L (2011) Modulation of behavioral phase changes of the migratory locust by the catecholamine metabolic pathway. Proceedings of the National Academy of Sciences U.S.A. 108: 3882–3887. doi:10.1073/pnas.1015098108
  12. Ma Z, Guo X, Lei H, Li T, Hao S, Kang L (2015) Octopamine and tyramine respectively regulate attractive and repulsive behavior in locust phase changes. Sci. Rep. 5:8036. doi:10.1038/srep08036
  13. Uvarov B (1966) Grasshoppers and Locusts: A Handbook of General Acridology. 1. Anatomy, Physiology, Development, Phase Polymorphism, Introduction to Taxonomy. Cambridge: University Press.
  14. Uvarov B (1977) Grasshoppers and Locusts. A Handbook of General Acridology Vol. 2. Behaviour, Ecology, Biogeography, Population Dynamics. Cambridge: Centre for Overseas Pest Research.
  15. Pener M P and Simpson SJ (2009) “Locust phase polyphenism: an update,” in Advances in Insect Physiology, eds S. J. Simpson and M. P. Pener (London:Academic Press), 1–272. doi:10.1016/S0065-2806(08)36001-9
  16. Benfekih L, Chara B, Doumandji-Mitiche B (2002) Influence of anthropogenic impact on the habitats and swarming risks of Dociostourus morocconus and Locusto migratoria (Orthoptera, Acrididae) in the Algerian Sahara and the semiarid zone. Journal of Orthoptera Research 11: 243–250. https://doi.org/10.1665/1082-6467(2002)011[0243:IOAIOT]2.0.CO;2
  17. Benfekih L, Petit D (2010) The annual cycle of Saharan populations of Locusta migratoria cinerascens (Orthoptera: Acrididae: Oedipodinae) in Algeria. Annales de la Société entomologique de France (N.S.) 46: 351–358. https://doi.org/10.1080/00379271.2010.10697674
  18. Farrow RA (1987) Effects of changing land use on outbreaks of tropical migratory locust, Locusta migratoria migratorioides (R. and F.). International Journal of Tropical Insect Science, 8 969–975. doi:10.1017/S1742758400023262
  19. OCHA Madagascar: Grand Sud Humanitarian Snapshot (August 2021) file:///Users/miraword/Downloads/Madagascar_20210827_GrandSud_HumanitarianSnapshot%20(1).pdf
  20. FAO Regional Office for Africa. (September 4, 2020) Locust outbreaks threaten food security in southern Africa. http://www.fao.org/africa/news/detail-news/en/c/1306167/
  21. Anon FAO and the locust crisis in Madagascar : FAO in Emergencies. Available from: http://www.fao.org/emergencies/crisis/madagascar-locust/intro/en/ (November 20, 2019).


Additional Papers Hunter DM, Strong K, Spurgin PA (1998) Management of Populations of the Spur-Throated Locust, Austracris guttulosa (Walker), and Migratory Locust, Locusta migratoria (L.) (Orthoptera: Acrididae), in Eastern Australia during 1996 and 1997. Journal of Orthoptera Research: 173–178. https://doi.org/10.2307/3503515

Lomer CJ, Bateman RP, Johnson DL, Langewald J, Thomas M (2001) BIOLOGICAL CONTROL OF LOCUSTS AND GRASSHOPPERS. Annual Review of Entomology 46: 667–702. https://doi.org/10.1146/annurev.ento.46.1.667

Steedman A (1990) Other African locusts. In: Locust Handbook. Natural Resources Institute, Chatham, 204.

Bei-Bienko G.Ya. 1932. A manual of locust survey. Leningrad: Upravlenie Sluzhby ucheta Gos. OBV Narkozema SSSR. 159 pp.

Bei-Bienko G.Ya., Mishchenko L.L. 1951. Locusts of the fauna of the USSR and adjacent countries. Keys to Fauna of USSR, N38, parts I & II. Moscow & Leningrad: AN SSSR. 668 pp.

Latchininsky A.V., Sergeev M.G., Childebaev M.K., Chernyakhovsky M.E., Lockwood J.A.,

Kambulin V.E., Gapparov F.A. 2002. Locusts of Kazakhstan, Central Asia and adjacent territories. Larami: Association for Applied Akridology International, University of Wyoming. 387 pp.

Mishchenko L.L. 1952. Locusts (Catantopinae). Fauna of the USSR. Orthopterous insects. Leningrad: AN SSSR. V. 4(2): 610.

Qing, Y. Q. L. Z. L., and Chunxian, Y. G. F. C. J. (2008). Feeding habits and economic threshold of Locusta migratoria tibetensis. Acta Phytophylac. Sin. 5.

Sergeev M.G. 1986. Regularities in distribution of orthopterous insects of Northern Asia. Novosibirsk: Nauka. 238 pp.

Tanaka, S., and Zhu, D.-H. (2005). Outbreaks of the migratory locust Locusta migratoria (Orthoptera: Acrididae) and control in China. Appl. Entomol. Zool. 40, 257–263. doi: 10.1303/aez.2005.257

Tsyplenkov E.P. 1970. Locust pests in the USSR. Leningrad: Kolos. 272 pp.

Wang, Xian-Hui, and Le Kang. "Differences in egg thermotolerance between tropical and temperate populations of the migratory locust Locusta migratoria (Orthoptera: Acridiidae)." Journal of Insect Physiology 51.11 (2005): 1277-1285.