Locust

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Today 27/5/2020 in India Rajasthan more than 15000 locus were killed
This article is about the insect. For other uses, see Locust (disambiguation).
File:Locusta-migratoria-wanderheuschrecke.pg
Locusts are grasshoppers, such as this migratory locust (Locusta migratoria), that have entered into a migratory phase
of their life.

Garden locust, Acanthacris ruficornis, in Ghana

Millions of swarming Australian plague locusts on the move

Locusts are a collection of certain species of short-horned grasshoppers in the

family Acrididae that have a swarming phase. These insects are usually solitary, but under
certain circumstances they become more abundant and change their behaviour and habits,
becoming gregarious. No taxonomic distinction is made between locust and grasshopper
species; the basis for the definition is whether a species forms swarms under intermittently
suitable conditions.
These grasshoppers are normally innocuous, their numbers are low, and they do not pose a
major economic threat to agriculture. However, under suitable conditions of drought followed
by rapid vegetation growth, serotonin in their brains triggers a dramatic set of changes: they
start to breed abundantly, becoming gregarious and nomadic (loosely described
as migratory) when their populations become dense enough. They form bands of
wingless nymphs which later become swarms of winged adults. Both the bands and the
swarms move around and rapidly strip fields and cause damage to crops. The adults are
powerful fliers; they can travel great distances, consuming most of the green vegetation
wherever the swarm settles.[1]
Locusts have formed plagues since prehistory. The ancient Egyptians carved them on their
tombs and the insects are mentioned in the Iliad, the Bible and the Quran. Swarms have
devastated crops and been a contributory cause of famines and human migrations. More
recently, changes in agricultural practices and better surveillance of locations where swarms
tend to originate, have meant that control measures can be used at an early stage. The
traditional means of control are based on the use of insecticides from the ground or the air,
but other methods using biological control are proving effective.
Swarming behaviour decreased in the 20th century, but despite modern surveillance and
control methods, the potential for swarms to form is still present, and when suitable climatic
conditions occur and vigilance lapses, plagues can still occur.[2][3] Locusts are large insects
and convenient for use in research and the study of zoology in the classroom. They are
also edible insects; they have been eaten throughout history and are considered
a delicacy in many countries.
The word "locust" is derived from the Vulgar Latin locusta, meaning grasshopper.[4]

Contents

 1Swarming grasshoppers
o 1.1Solitary and gregarious phases
 2Distribution and diversity
 3Interaction with humans
o 3.1Ancient times
o 3.2Recent times
o 3.3Monitoring
o 3.4Control
o 3.5As experimental models
o 3.6As food
 4See also
 5Notes
 6References
 7External links

Swarming grasshoppers[edit]
Main articles: Grasshopper, Swarm behaviour, and Predator satiation

A desert locust ovipositing in sand

Locusts are the swarming phase of certain species of short-horned grasshoppers in the
family Acrididae. These insects are usually solitary, but under certain circumstances become
more abundant and change their behaviour and habits, becoming gregarious.[5][6][7]
Desert locusts in copulation

No taxonomic distinction is made between locust and grasshopper species; the basis for the
definition is whether a species forms swarms under intermittently suitable conditions. In
English, the term "locust" is used for grasshopper species that change morphologically and
behaviourally on crowding, forming swarms that develop from bands of immature stages
called hoppers. The change is referred to in the technical literature as "density-
dependent phenotypic plasticity".
These changes are examples of phase polymorphism; they were first analysed and
described by Boris Uvarov, who was instrumental in setting up the Anti-Locust Research
Centre.[8] He made his discoveries during his studies of the migratory locust in Caucasus,
whose solitary and gregarious phases had previously been thought to be separate species
(Locusta migratoria and L. danica L.). He designated the two phases
as solitaria and gregaria.[9] These are also referred to as statary and migratory morphs,
though strictly speaking, their swarms are nomadic rather than migratory. Charles Valentine
Riley and Norman Criddle were also involved in achieving the understanding and control of
locusts.[10][11]
Swarming behaviour is a response to overcrowding. Increased tactile stimulation of the hind
legs causes an increase in levels of serotonin.[12] This causes the locust to change colour, eat
much more, and breed much more easily. The transformation of the locust to the swarming
form is induced by several contacts per minute over a four-hour period.[13] A large swarm can
consist of billions of locusts spread out over an area of thousands of square kilometres, with
a population of up to 80 million per square kilometre (200 million per square mile).[14] When
desert locusts meet, their nervous systems release serotonin, which causes them to become
mutually attracted, a prerequisite for swarming.[15][16]
The initial bands of gregarious hoppers are known as "outbreaks", and when these join
together into larger groups, the event is known as an "upsurge". Continuing agglomerations
of upsurges on a regional level originating from a number of entirely separate breeding
locations are known as "plagues".[17] During outbreaks and the early stages of upsurges, only
part of the locust population becomes gregarious, with scattered bands of hoppers spread
out over a large area. As time goes by, the insects become more cohesive and the bands
become concentrated in a smaller area. In the desert locust plague in Africa, the Middle
East, and Asia that lasted from 1966 to 1969, the number of locusts increased from two to
30 billion over two generations, but the area covered decreased from over 100,000 square
kilometres (39,000 sq mi) to 5,000 square kilometres (1,900 sq mi).[18]
Solitary and gregarious phases[edit]
Solitaria (grasshopper) and gregaria (swarming) phases of the desert locust

One of the greatest differences between the solitary and gregarious phases is behavioural.
The gregaria nymphs are attracted to each other, this being seen as early as the
second instar. They soon form bands of many thousands of individuals. These groups
behave like cohesive units and move across the landscape, mostly downhill, but making
their way around barriers and merging with other bands. The attraction between the insects
seems to be largely visual, but also involves olfactory cues, and the band seem to navigate
using the sun. They pause to feed at intervals before resuming their march, and may cover
tens of kilometres over a few weeks.[9]
Also, differences in morphology and development are seen. In the desert locust and the
migratory locust, for example, the gregaria nymphs become darker with strongly contrasting
yellow and black markings, they grow larger, and have longer developmental periods. The
adults are larger with different body proportions, less sexual dimorphism, and
higher metabolic rates. They mature more rapidly and start reproducing earlier, but have
lower levels of fecundity.[9]
The mutual attraction between individual insects continues into adulthood, and they continue
to act as a cohesive group. Individuals that get detached from a swarm fly back into the
mass. Others that get left behind after feeding, take off to rejoin the swarm when it passes
overhead. When individuals at the front of the swarm settle to feed, others fly past overhead
and settle in their turn, the whole swarm acting like a rolling unit with an ever-changing
leading edge. The locusts spend much time on the ground feeding and resting, moving on
when the vegetation is exhausted. They may then fly a considerable distance before settling
in a location where transitory rainfall has caused a green flush of new growth.[9]

Distribution and diversity[edit]

Main article: List of locust species
Several species of grasshoppers swarm as locusts in different parts of the world, on all
continents except Antarctica and North America:[19][20][21][a] For example, the Australian plague
locust (Chortoicetes terminifera) swarms across Australia.[19]
The desert locust (Schistocerca gregaria) is probably the best known species owing to its
wide distribution (North Africa, Middle East, and Indian subcontinent)[19] and its ability
to migrate over long distances. A major infestation covered much of western Africa in 2003-
4, after unusually heavy rain set up favourable ecological conditions for swarming. The first
outbreaks occurred in Mauritania, Mali, Niger, and Sudan in 2003. The rain allowed swarms
to develop and move north to Morocco and Algeria, threatening croplands.[23][24] Swarms
crossed Africa, appearing in Egypt, Jordan and Israel, the first time in those countries for 50
years.[25][26] The cost of handling the infestation was put at US$122 million, and the damage to
crops at up to $2.5 billion.[27]
The migratory locust (Locusta migratoria), sometimes classified into up to 10 subspecies,
swarms in Africa, Asia, Australia, and New Zealand, but has become rare in Europe.[28] In
2013, the Madagascan form of the migratory locust formed many swarms of over a billion
insects, reaching "plague" status and covering about half the country by March
2013.[29] Species such as the Senegalese grasshopper (Oedaleus senegalensis)[30] and
the African rice grasshopper (Hieroglyphus daganensis), both from the Sahel, often display
locust-like behaviour and change morphologically on crowding.[30]
North America is currently the only continent besides Antarctica without a native locust
species. The Rocky Mountain locust was formerly one of the most significant insect pests
there, but it became extinct in 1902. In the 1930s, during the Dust Bowl, a second species of
North American locust, the High Plains locust (Dissosteira longipennis) reached plague
proportions in the American Midwest. Today, the High Plains locust is a rare species, leaving
North America with no regularly swarming locusts.

Interaction with humans[edit]

Ancient times[edit]

Locust detail from a hunt mural in the grave-chamber of Horemhab, Ancient Egypt, circa 1422–1411 BC

Study of literature shows how pervasive plagues of locusts were over the course of history.
The insects arrived unexpectedly, often after a change of wind direction or weather, and the
consequences were devastating. The Ancient Egyptians carved locusts on tombs in the
period 2470 to 2220 BC, and a devastating plague is mentioned in the Book of Exodus in the
Bible, as taking place in Egypt around 1446 BC.[18][31] The Iliad mentions locusts taking to the
wing to escape fire.[32] Plagues of locusts are also mentioned in the Quran.[14] In the ninth
century BC, the Chinese authorities appointed anti-locust officers.[33]
Aristotle studied locusts and their breeding habits and Livy recorded a devastating plague
in Capua in 203 BC. He mentioned human epidemics following locust plagues which he
associated with the stench from the putrifying corpses; the linking of human disease
outbreaks to locust plagues was widespread. A pestilence in the northwestern provinces of
China in 311 AD that killed 98% of the population locally was blamed on locusts, and may
have been caused by an increase in numbers of rats (and their fleas) that devoured the
locust carcases.[33]
Recent times[edit]
Locusts which swarmed over England in 1748: Drawing by De la Cour; engraved by R. White, in Thomas Pennant's A
Tour in Wales, 1781

During the last two millennia, locust plagues continued to appear at irregular intervals with
the main recorded outbreaks of the desert and migratory locusts occurring in Africa, the
Middle East, and Europe. Other species of locusts caused havoc in North and South
America, Asia, and Australasia; 173 outbreaks over a period of 1924 years have been
recorded in China.[33] The Bombay locust (Nomadacris succincta) was a major pest in India
and southeastern Asia in the 18th and 19th centuries, but has seldom swarmed since the
last plague in 1908.[34]
In the spring of 1747 locusts arrived outside Damascus eating the majority of the crops and
vegetation of the surrounding countryside. One local barber, Ahmad al-Budayri, recalled the
locusts "came like a black cloud. They covered everything: the trees and the crops. May God
Almighty save us!"[35]
The extinction of the Rocky Mountain locust has been a source of puzzlement. It had
swarmed throughout the west of the United States and parts of Canada in the 19th
century. Albert's swarm of 1875 was estimated to cover 198,000 square miles (510,000 km2)
(greater than the area of California) and to weigh 27.5 million tons, with some 12.5 trillion
insects.[36] The last specimen was seen alive in Canada in 1902. Recent research suggests
the breeding grounds of this insect in the valleys of the Rocky Mountains came under
sustained agricultural development during the large influx of gold miners,[37] destroying the
underground eggs of the locust.[38][39]
Monitoring[edit]

Eugenio Morales Agacino on expedition monitoring locusts in the desert of Spanish Sahara, 1942

Early intervention is a more successful means of dealing with locusts than later action when
swarms have already built up. The technology to control locust populations is now available,
but the organisational, financial, and political problems may be difficult to overcome.
Monitoring is the key to reducing damage, with the early detection and eradication of bands
being the objective. Ideally, a sufficient proportion of nomadic bands can be treated with
insecticide before the swarming phase is reached. Reaching this objective may be possible
in richer countries like Morocco and Saudi Arabia, but neighbouring poorer countries
(Mauritania, Yemen, etc.) lack the resources and may act as a source of locust swarms that
threaten the whole region.[14]
Several organisations around the world monitor the threat from locusts. They provide
forecasts detailing regions likely to suffer from locust plagues in the near future. In Australia,
this service is provided by the Australian Plague Locust Commission.[40] It has been very
successful in dealing with developing outbreaks, but has the great advantage of having a
defined area to monitor and defend without locust invasions from elsewhere.[41] In Central
and Southern Africa, the service is provided by the International Locust Control Organization
for Central and Southern Africa.[42] In West and Northwest Africa, the service is co-ordinated
by the Food and Agriculture Organization's Commission for Controlling the Desert Locust in
the Western Region, and executed by locust control agencies belonging to each country
concerned.[43] The FAO also monitors the situation in the Caucasus and Central Asia, where
over 25 million hectares of cultivated land are under threat.[44] In February 2020, in an effort
to end massive locust outbreaks, India decided to use drones and special equipment to
monitor locusts and spray insecticides.[45]
Control[edit]

Preparing to flame locusts in Palestine, 1915

Historically, people could do little to protect their crops from being devastated by locusts,
although eating the insects may have been some consolation. By the early 20th century,
efforts were being made to disrupt the development of the insects by cultivating the soil
where eggs were laid, collecting hoppers with catching machines, killing them with
flamethrowers, trapping them in ditches, and crushing them with rollers and other
mechanical methods.[18] By the 1950s, the organochloride dieldrin was found to be an
extremely effective insecticide, but it was later banned from use in most countries because
of its persistence in the environment and its bioaccumulation in the food chain.[18]

Cessna of the International Red Locust Control Organization spraying red locusts in Iku Katavi National Park, Tanzania,
2009

In years when locust control is needed, the hoppers are targeted early by applying water-
based, contact pesticides using tractor-based sprayers. This is effective but slow and labour-
intensive, and where possible, spraying concentrated insecticide solutions from aircraft over
the insects or the vegetation on which they feed is preferable.[46] The use of ultralow-volume
spraying of contact pesticides from aircraft in overlapping swathes is effective against
nomadic bands and can be used to treat large areas of land swiftly.[41] Other modern
technologies used for planning locust control include GPS, GIS tools, and satellite imagery,
and computers provide rapid data management and analysis.[47][48]

Locusts killed by the naturally occurring fungus Metarhizium, an environmentally friendly means of biological control [49]

A biological pesticide to control locusts was tested across Africa by a multinational team in
1997.[50] Dried fungal spores of a Metarhizium acridum sprayed in breeding areas pierce the
locust exoskeleton on germination and invade the body cavity, causing death.[51] The fungus
is passed from insect to insect and persists in the area, making repeated treatments
unnecessary.[52] This approach to locust control was used in Tanzania in 2009 to treat around
10,000 hectares in the Iku-Katavi National Park infested with adult locusts. The outbreak
was contained and the elephants, hippopotamuses, and giraffes present in the area were
unharmed.[42]
The ultimate goal in locust control is the use of preventive and proactive methods that
disrupt the environment to the least possible extent. This would make agricultural production
easier and more secure in the many regions where growing crops is of vital importance to
the survival of the local people.[17]
As experimental models[edit]
The locust is large and easy to breed and rear, and is used as an experimental model in
research studies. It has been used in evolutionary biology research and to discover to what
degree conclusions reached about other organisms, such as the fruit fly (Drosophila) and
the housefly (Musca), are applicable to all insects.[53][54] It is a suitable school laboratory
animal because of its robustness and the ease with which it can be grown and handled.[55]
As food[edit]
See also: Kosher locust

Skewered locusts in Beijing, China

Locusts are edible insects. Several cultures throughout the world consume insects, and
locusts are considered a delicacy and eaten in many African, Middle Eastern, and Asian
countries. They have been used as food throughout history.[56]
They can be cooked in many ways, but are often fried, smoked, or dried.[57] The Bible records
that John the Baptist ate locusts and wild honey (Greek: ἀκρίδες καὶ μέλι
ἄγριον, romanized: akrides kai meli agrion) while living in the wilderness.[58] Attempts have been
made to explain the locusts as some ascetic vegetarian food such as carob beans, but the
plain meaning of akrides is the insects.[59][60]
The Torah, although disallowing the use of most insects as food, permits the consumption of
certain locusts; specifically, the red, the yellow, the spotted grey, and the white are
considered permissible.[61][62] In Islamic jurisprudence, eating locusts is
considered halal.[63][62] The Islamic prophet, Muhammad, was reported to have eaten locusts
during a military raid with his companions.[64]
Locusts are eaten in the Arabian Peninsula, including Saudi Arabia,[65] where consumption of
locusts spiked around Ramadan especially in the Al-Qassim Region in 2014, since many
Saudis believe they are healthy to eat. The Saudi Ministry of Health warned that pesticides
they used against the locusts made them unsafe.[66][67] Yemenis also consume locusts, and
expressed discontent over governmental plans to use pesticides to control them.[68] ʻAbd al-
Salâm Shabînî described a locust recipe from Morocco.[69] 19th century European travellers
observed Arabs in Arabia, Egypt, and Morocco selling, cooking, and eating locusts.[70] They
reported that in Egypt and Palestine locusts were consumed.[71] They reported that in
Palestine, around the River Jordan, in Egypt, in Arabia, and in Morocco that Arabs ate
locusts, while Syrian peasants did not eat locusts.
In the Haouran region, Fellahs who were in poverty and suffered from famine ate locusts
after removing the guts and head, while locusts were swallowed wholesale by
Bedouins.[72] Syrians, Copts, Greeks, Armenians, and other Christians and Arabs themselves
reported that in Arabia locusts were eaten frequently and one Arab described to a European
traveler the different types of locusts which were favored as food by Arabs.[73][74] Persians use
the Anti-Arab racial slur "Arabe malakh-khor" (Persian: ‫عرب ملخ خور‬, literally Arab locust
eater) against Arabs.[75][76][77][78]
Locusts yield about five times more edible protein per unit of fodder than cattle, and produce
lower levels of greenhouse gases in the process.[79] The feed conversion rate of orthopterans
is 1.7 kg/kg,[80] while for beef it is typically about 10 kg/kg.[81] The protein content in fresh
weight is between 13–28 g/100g for adult locust, 14–18 g/100g for larvae, as compared to
19–26 g/100g for beef.[82][83] The calculated protein efficiency ratio is low, with 1.69 for locust
protein compared to 2.5 for standard casein.[84] A serving of 100 g of desert locust provides
11.5 g of fat, 53.5% of which is unsaturated, and 286 mg of cholesterol.[84] Among the fatty
acids, palmitoleic, oleic and linolenic acids were found to be the most abundant. Varying
amounts of potassium, sodium, phosphorus, calcium, magnesium, iron, and zinc were
present.[84]

See also[edit]
 List of locust swarms
 Australian Plague Locust Commission
 LUBILOSA

Notes[edit]
1. ^ The American locust (Schistocerca americana) does not swarm.[22]
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External links[edit]
Look up locust in
Wiktionary, the free
dictionary.

Wikimedia Commons has

media related to Locusta.

 Visual neuron of the locust, Ri Channel video, October 2011

  FAO Locust Watch
 FAO EMPRES
 FAO eLocust3e suite
 eLocust3M android app
 Desert Locust Meteorological Monitoring at Sahel Resources
 Locust Video
 USAID Supplemental Environmental Assessment of the
Eritrean Locust Control Program
 USAID Supplemental Environmental Assessment: Pakistan
Locust Control Programs, August 1993
 footage on YouTube
 When The Skies Turned To Black, The Locust Plague of
1875
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