Available via license: CC BY 4.0
Content may be subject to copyright.
Journal of orthoptera research 2021, 30(2)
Journal of Orthoptera Research 2021, 30(2): 145–154
Abstract
The recent upsurge of the desert locust Schistocerca gregaria (Forskål,
1775) has had an impact on East Africa and the Middle East as far as India.
It has affected and slowed down many aspects of the Pakistani economy.
Swarms of locusts have infested many areas and caused immense dam-
age to all types of crops. Both farmers and economists are concerned and
are trying to get the most up-to-date information on the best strategy to
manage this pest. This paper is an attempt to (i) provide insight into the
dynamics of this upsurge internationally as well as in the various regions
of Pakistan, (ii) briefly assess its local impact and locust control measures,
and (iii) clarify the role of the various stakeholders in the management,
both nationally and internationally, suggesting various improvements for
the future.
Keywords
control strategies, crop damage, desert locust, outbreak, pest, Schistocerca
gregaria
Introduction
In 2019 and 2020, large swarms of desert locusts again threat-
ened parts of East Africa and large areas as far as India and Pakistan
via the Arabian Peninsula. The Food and Agriculture Organization
(FAO) of the United Nations has described this locust situation
as the most serious in decades (FAO 2019, 2020a). The swarms
reached Kenya, Uganda, and Tanzania, which had not faced a
threat of this magnitude for 70 years. Although we have already
seen classic images of these devastating swarms in the past, their
impact is still impressive. Trees can twist and branches can break
under the weight of locusts. Without adequate means of control,
farmers are made desperate by the loss of their crops. Equipped
with manual sprayers and often poorly protected against insec-
ticides, technicians try to fight against these insects where only
aerial means would be effective. Fortunately, even if detected too
late, this upsurge quickly became the subject of major control op-
erations, with the assistance of various donors and under the co-
ordination of the FAO. However, after two years of intensive fight-
ing, the situation is still not under control. In early 2021, calm
returned to Southwest Asia, and in particular Pakistan, but these
swarms have yet to be contained in the Horn of Africa (Dowlatch-
ahi et al. 2020b).
The desert locust Schistocerca gregaria (Forskål, 1775) (Insecta:
Orthoptera: Acrididae) is considered a serious agricultural pest
in West and North Africa, the Middle East, and Southwest Asia
(Steedman 1990, Cressman 2016, Lecoq 2019), and regular inva-
sions of this insect pose a real threat to agricultural production
and have devastating consequences for food security in more than
50 countries (Lecoq 2003, 2004, 2005, Brader et al. 2006). The
social impact of an invasion can be visible in the long term, even
after 20 years (De Vreyer et al. 2014). Like other locust species,
the desert locust exhibits phase polyphenism, a plastic response
to population density associated with several changes in behavio-
ral, morphological, anatomical, and physiological traits. Isolated,
harmless, and hidden solitary locusts transform into huge hopper
bands and devastating swarms of the gregarious form under condi-
tions of overpopulation (Uvarov 1921, 1966, Pener and Simpson
2009, Piou et al. 2017). Of the 31 million km2 that can be invaded
by the desert locust (the invasion area), the remission area (where
low-density solitary-phase populations exist during calm peri-
ods) covers only 15 million km2. In this zone, the outbreak areas
(where the first outbreaks that could lead to invasions occur due to
appropriate ecological characteristics) occupy an even smaller area
of about 1.7 million km2 (Sword et al. 2010, Gay P.E. p.c.).
The lifespan of a locust generation, under optimal conditions, is
40–50 days, and the annual number of generations varies between
two and three. Young adults may remain immature (quiescent) for
several months until they find moist conditions favorable for egg
laying, with 20–25 mm of rainfall being normally sufficient (Du-
ranton and Lecoq 1990, Symmons and Cressman 2001). As rainfall
is seasonally distributed throughout the habitat area, this results
in the existence of three main breeding seasons—spring, summer,
and winter—between which the imagos undertake seasonal migra-
tions to benefit from favorable breeding conditions (COPR 1982,
The 2019–2020 upsurge of the desert locust and its impact in Pakistan
riffat sultana1, santosh Kumar2, ahmed ali sameJo1, samiallah soomro1, michel lecoq3
1 Department of Zoology, University of Sindh, Jamshoro, Sindh, Pakistan.
2 Department of Zoology, Cholistan University of Veterinary and Animal Sciences, Bahawalpur, Punjab, Pakistan.
3 CIRAD, UMR CBGP, F-34398 Montpellier, France.
Corresponding author: Riffat Sultana (riffat.sultana@usindh.edu.pk)
Academic editor: Daniel Petit | Received 15 March 2021 | Accepted 10 May 2021 | Published 7 October 2021
http://zoobank.org/66ADFDBD-4372-4F8C-8374-60DD4C76CFA2
Citation: Sultana R, Kumar S, Samejo AA, Soomro S, Lecoq M (2021) The 2019–2020 upsurge of the desert locust and its impact in Pakistan. Journal of
Orthoptera Research 30(2): 145–154. https://doi.org/10.3897/jor.30.65971
Research Article
Journal of orthoptera research 2021, 30(2)
R. SULTANA, S. KUMAR, A.A. SAMEJO, S. SOOMRO AND M. LECOQ
146
Duranton and Lecoq 1990, Steedman 1990, Symmons and Cress-
man 2001). During periods of remission, solitary populations are
dispersed in desertic areas. There, gregarization (transformation of
locusts from solitary to gregarious) begins in the outbreak areas,
thanks to rainy sequences favorable to reproduction in grouped
vegetation and on mainly sandy or sandy-clay soils (Collett et al.
1998, Despland et al. 2000, Cissé et al. 2013). Outbreaks develop
and then, if good conditions persist, follow increasingly severe
phases of upsurge and plague (classic terminology defined by FAO
2009a) as populations and the number of occupied sites increase.
The invasions develop intermittently and, in the past, have fre-
quently persisted for 5, 10, or more years (Sword et al. 2010).
Since the 1960s, a preventive control strategy has been recom-
mended by the FAO based on the monitoring of outbreak areas
and ecological conditions (Showler et al. 2021, Lecoq 2003, 2004,
Sword et al. 2010), followed, if necessary, by early intervention and
thus limited use of pesticides. The implementation of this strat-
egy helps to maintain low densities and to stop any outbreak as
soon as possible (Duranton and Lecoq 1990, Martini et al. 1998,
Magor et al. 2008, Sword et al. 2010). Consequently, and with 60
years of hindsight, it is clear that invasions are now less frequent,
smaller in scale and, if they cannot be stopped at an early stage,
shorter and better managed (Magor et al. 2008, Sword et al. 2010).
However, financial and political uncertainties, as well as recurrent
insecurity in many areas of desert locust distribution, continue to
maintain the threat, and some outbreaks cannot be stopped at an
early stage, as was observed again recently (Meynard et al. 2020,
Showler and Lecoq 2021, Showler et al. 2021).
Pakistan has historically been subject to periodic swarm inva-
sions. The country also contains outbreak areas, where particu-
larly ecological conditions can favor, when suitable rains occur,
the concentration, reproduction, and intensive multiplication of
locusts and give rise to outbreaks and plagues. In recent years, the
greatest outbreaks were noted in 1993 and 1997. These invasions
have caused incalculable damage to crops, sometimes leading to
severe famines. The recent upsurge in 2019–2020 seriously affected
the country (Dowlatchahi 2020a). Here, we present a summary of
these two years of upsurge by focusing on its impact in Pakistan, the
damage caused in this country, and the surveillance and control op-
erations undertaken. Furthermore, we try to clarify, both nationally
and internationally, the role of the various stakeholders in the man-
agement of this pest, suggesting some improvements for the future.
Materials and methods
The general pattern of the current global upsurge was taken
from the Desert Locust Bulletin produced monthly by FAO-DLIS
(Desert Locust Information Service) in Rome based on informa-
tion from all the countries within the desert locust habitat area
(FAO 2019, 2020a). These bulletins also provide information
about the likely migration of swarms based on the study of the
meteorological situation and the use of migration trajectory mod-
els. Meteorological data and remote sensing imagery are used to
help estimate rainfall, detect green vegetation, and identify areas
where ecological conditions may be favorable for locust breeding
(Cressman 2008). Regarding Pakistan, close contacts were main-
tained with the Department of Plant Protection (DPP)—the lead
institution tasked with monitoring and managing the desert lo-
cust threat in Pakistan—to obtain information on the ongoing
invasion and damage in various regions of the country. Desert lo-
cust field information was collected by DPP survey teams accord-
ing to a standardized procedure recommended by FAO-DLIS and
was used to produce monthly locust situation maps (Cressman
2001). Contacts were also maintained with inhabitants, farmers,
and local entomologists to obtain more daily information on
the presence and migration of locusts. In addition, field surveys
were carried out by the authors in different locust-affected locali-
ties, mainly in Sindh province; locust samples were collected, and
photos were taken in different affected areas of Sindh to document
the situation on the ground.
Results
General course of the desert locust upsurge in 2019–2020
The last major desert locust plague ended in 1962 (Magor et al.
2008, Sword et al. 2010). In recent years, for Pakistan, the greatest
outbreaks were in 1993 and 1997 (FAO 1993, 1997). Globally, the
last major upsurge was in 2004–2005 (FAO 2004, 2005). At the
end of 2018, the situation was calm throughout the desert locust
habitat area (FAO 2018). The FAO situation bulletins contained
almost no observations; in particular, no gregarious formations
were reported, and there was a single report of a group of hoppers
in northern Somalia during September. Then suddenly, in Decem-
ber 2018, laying swarms were reported on the coast of the Red Sea,
in Sudan, and in Eritrea. In January 2019, such laying swarms were
also seen on both sides of the Red Sea, and immature swarms were
seen in Saudi Arabia (FAO 2019). Iran and Pakistan were warned
of the possible migration of swarms. The situation quickly escalat-
ed in the following months (Figs 1, 2). Southern Iran was affected
in February, Pakistan in March, and India in May. The desert locust
situation continued to worsen across the Arabian Peninsula. In
June, the Horn of Africa (Sudan, Eritrea, Djibouti, and Somalia)
was affected by swarms originating from Yemen. In this region, the
cyclonic rains and floods of October and November 2019 created
good conditions in which the desert locust could continue to mul-
tiply. Kenya was infested as of December 2019. From June to De-
cember, the locust situation remained serious in southwest Asia,
particularly in Pakistan and western India. In 2020, the desert lo-
cust situation continued to worsen (FAO 2020a). Strong spring
breeding occurred in April, May, and June. The locust situation
remained critical in the eastern region in Iran, Pakistan, and India
until August and was only brought under control beginning in
September. This was largely a result of the capacity and experience
of these countries to monitor and control desert locusts in their
outbreak areas. On the other hand, the desert locust situation did
not improve on both sides of the Red Sea and the Horn of Africa,
where, in December 2020, it was deemed very critical. In the end,
and for the moment, only West Africa has been spared.
Although not seen until January 2019, the development of this
upsurge was the result of favorable conditions for the desert lo-
cust, which were occurring as early as 2018. Two cyclones brought
heavy rains in the Rub al Khali, or Empty Quarter of the Arabic
peninsula, in May and October 2018 (FAO 2020b). Rains were
very heavy in Yemen, Oman, Djibouti, northern Somalia, east-
ern Ethiopia, and southern Saudi Arabia (Meynard et al. 2020).
Favorable conditions for desert locust breeding were maintained
for many consecutive months, allowing at least three successive
generations to develop. However, locusts went undetected and
unchecked for a significant amount of time, mainly due to the
insecure conditions in the areas of origin, particularly in Yemen
(Showler and Lecoq 2021). Had the initial outbreak in Saudi Ara-
bia’s Rub al Khali been detected in the early stage and controlled
in the summer of 2018, swarms may not have reached Yemen, the
R. SULTANA, S. KUMAR, A.A. SAMEJO, S. SOOMRO AND M. LECOQ 147
Journal of orthoptera research 2021, 30(2)
African Red Sea coast, and parts of Iran. In addition, an initially
weak response in Iran (where very heavy flooding in the south-
west of the country allowed two generations of breeding) allowed
swarms to move to Pakistan and India (FAO 2020c, d), where an
unusually long summer monsoon resulted in three generations of
breeding along both sides of the Indo-Pakistan border.
The 2019–2020 upsurge dynamic in Pakistan
In early 2019, no locusts were reported in Pakistan. It was
not until March that isolated solitarious adults first appeared on
the Baluchistan coast in the Uthal region west of Karachi, pre-
sumably coming from Iran, which had been invaded in previ-
ous months. On the 16th March, a mature swarm and groups of
mating and laying adults were seen on the coast at Pasni, on the
Iranian border, and in the Kulanch valley region, west of Pas-
ni (Fig. 3). These arrivals continued in April, and the migrant
populations continued to lay eggs. In April, the first larvae of
the spring reproduction appeared in Balochistan in the coastal
areas of Pasni. Hoppers continued to emerge and develop in May
between Turbat and Gwadar, near Uthal, and in the interior near
Kharan. Groups of gregarious hoppers of all stages were then
found, mixed with scattered adults. The spring breeding ended
in early June in Balochistan, with a last report of a laying swarm
Fig. 1. General situation of the desert locust from January to December 2019 (modified from FAO 2019). Swarms or hopper bands:
immature adults, red square; mature adults, blue triangle; maturity unknown, black triangle up; egg laying or eggs, black triangle down;
hoppers, black circle; hoppers and adults, combined symbols. Groups of adults or hoppers: same symbols but hollowed out. Density
unknown: same symbols, but partial.
Journal of orthoptera research 2021, 30(2)
R. SULTANA, S. KUMAR, A.A. SAMEJO, S. SOOMRO AND M. LECOQ
148
on 1st June near Lasbela, while hoppers and hopper groups per-
sisted near Lasbela, Turbat, Gwadar, and in the northern interior
near Dalbandin.
The adults from the spring breeding gradually migrated to the
summer breeding area (June–November) on the Indo-Pakistan
border. Some swarms may have originated from the Horn of Af-
rica after migrating over the Indian Ocean. This summer breeding
started at the end of May with scattered gregarious adults that
appeared during the last week of May near the Indian border
southeast of Chaman starting to lay. It developed mainly in June,
July, and August in the Nara, Cholistan, and Thar deserts east of
the Indus Valley. In Cholistan, egg laying continues until August.
Thus, from mid-August, outbreaks of a second generation caused
locust numbers to further increase. This second generation de-
veloped mainly in September, October, and November. Wide-
spread breeding was then observed in the deserts of Cholistan,
Nara, and Thar, where numerous hopper bands were forming,
giving rise to numerous swarms. During November and Decem-
ber, a third generation of breeding occurred in the Thar, Nara,
and Cholistan deserts, where numerous hopper groups formed,
resulting in numerous adult groups and immature swarms (FAO
2019) (Fig. 4).
The swarms then began to move westward to the winter-spring
breeding areas (February–June). Cross-border movements of
swarms from the summer breeding areas of Rajasthan in India oc-
curred. On 11 November, an immature westward swarm was seen
flying over Karachi. In southern Balochistan, immature swarms
from the summer breeding areas started to arrive in December.
Fig. 2. General situation of the desert locust from January to December 2020 (modified from FAO 2020a). Swarms or hopper bands:
immature adults, red square; mature adults, blue triangle; maturity unknown, black triangle up; egg laying or eggs, black triangle down;
hoppers, black circle; hoppers and adults, combined symbols. Groups of adults or hoppers: same symbols but hollowed out. Density
unknown: same symbols, but partial.
R. SULTANA, S. KUMAR, A.A. SAMEJO, S. SOOMRO AND M. LECOQ 149
Journal of orthoptera research 2021, 30(2)
In January 2020, a few nymphs of the 3rd generation continued
to molt. Groups of immature adults persisted in the Thar, Nara,
and Cholistan deserts. Cross-border movements of immature
swarms continued westward. On 21 February, three swarms report-
edly arrived in the Afghan province of Khost from adjacent areas
in northwest Pakistan.
A new spring breeding started in March 2020 and went until
May. During March, breeding took place mainly in Balochistan
(Khuzdar, Nushki, Washuk, Kharan and Dalbandin, Chagai, Pan-
jgur, Turbat, and Pasni) and in the Indus valley (Rajanpur, Kash-
more, Sukkur, Dera Ismail Khan, and Rohri), as well as in the
plains of Punjab. Breeding continued into April and May, and a
second generation of laying began in mid-April in the north near
Dalbandin in Balochistan. As a result of this breeding, an increas-
ing number of adult groups and immature swarms formed and
began to mature during May.
During June, as conditions dried out, these swarms moved
from the spring breeding areas eastward to the summer breeding
areas of the Cholistan, Nara, and Thar deserts in Punjab and Sindh
provinces. Some continued to India due to the too-dry conditions.
Summer breeding started in late June and continued into July
and August. Numerous first-generation hopper groups and bands
formed, especially in the Thar desert up to the Indian border in the
extreme southeast of Sindh. The imaginal molts began during the
first week of August, causing groups of immature adults to form
on the Indian border.
Then, in September, the situation improved dramatically. In
Sindh, a very limited second-generation breeding occurred in
September west of Hyderabad and in Tharparkar. Improvement
continued in October, and no locusts were seen in November and
December (Dowlatchahi et al. 2020b, FAO 2020a, d).
Damage and control measures in Pakistan
Damage.—The desert locust can consume most plant species and
crops (COPR 1982). Only a few plants are not eaten, such as the
neem Azadirachta indica A. Juss., Genista sp. (broom bush), and
Euphorbia hirta L. (asthma plant). Indeed, during this upsurge, a
great deal of damage was caused to all types of crops, including
wheat, cotton, rice, sugarcane, tobacco, corn, chickpea (gram)
sunflower, sorghum, pearl millet, mung bean (Vigna radiata (L.)
R. Wilczek), muth bean (Vigna aconitifolia Jacq.), sesame, cluster
bean (guar), potato, tomato, cabbage, cauliflower, carrot, peas,
onion, melon, cucumber, water-melon, chilies, eggplant (brinjal),
okra (lady finger), mango, citrus, apple, grapes, strawberry, peach-
es, banana, and guava.
This upsurge has been devastating for a country where agricul-
ture represents around 20% of the GDP and where 61% of the pop-
ulation lives and works in agricultural areas (FAO 2016). About 52
districts were reported to have suffered locust damage. According to
FAO estimates in May 2020 and assuming that the damage accounts
for about 25% of growing crops, losses could reach 353 billion Paki-
stani rupees (2.19 billion US$) for “rabi crops” (sown in winter and
harvested in spring) and about 464 billion Pakistani rupees (2.88
billion US$) for “kharif crops” (summer sown crops) (FAO 2020e).
The final balance has yet to be established and, in the end, the dam-
age from this upsurge will undoubtedly be much higher. In 2020,
the Government of Pakistan’s preliminary estimate of monetary
losses due to desert locusts over the two coming agricultural seasons
in 2020 and 2021 may range from 3.4 billion US$ to 10.21 billion
US$. More than 3 million people in Pakistan are facing severe acute
food insecurity, with the situation particularly precarious in Balo-
chistan. It is estimated that approximately 34,000 households will
need emergency livelihood and food security assistance due to crop
losses. Many more people may be indirectly affected by crop losses,
leading to price rises in key commodities (FAO 2020).
Control measures.—To better coordinate control operations, the
Government of Pakistan declared the locust invasions to be an
emergency. Many anticipatory measures have been taken in col-
laboration with the FAO, in coordination with neighboring coun-
tries, and with the support of international partners to face the
threat and be ready to respond quickly and effectively (FAO 2019,
2020a). In addition, the Space and Upper Atmosphere Research
Fig. 3. Desert locust breeding areas in Southern Pakistan during invasions (adapted from Symmons and Cressman 2001). Arrows: main
orientation of migrating swarms in March–July (blue) and August–October (red).
Journal of orthoptera research 2021, 30(2)
R. SULTANA, S. KUMAR, A.A. SAMEJO, S. SOOMRO AND M. LECOQ
150
Fig. 4. Photos of the desert locust in Pakistan. A–D. Outbreaks in various localities during the 2019–2020 upsurge; E, F. Individual and
group mating in the Thar desert (photos from the authors).
R. SULTANA, S. KUMAR, A.A. SAMEJO, S. SOOMRO AND M. LECOQ 151
Journal of orthoptera research 2021, 30(2)
Commission (SUPARCO) has helped through the use of remote
sensing used to delimit the areas more vulnerable to locust attacks
in the various affected districts of Pakistan based on vegetation,
soil type, and other factors.
Large areas, about 65 M ha, were surveyed by the DPP, and
in 2 years, based on data collected by the FAO (2019, 2020a), a
total of 566,390 ha was treated (FAO 2019, 2020b). Most of the
treatment was done using specialized ultra-low volume (ULV)
vehicle-mounted sprayers, mainly with malathion (ULV formula-
tion) in desert areas and lambda-cyhalothrin (EC formulation) for
protection against desert locust attacks in crop production areas;
543,370 ha were treated by land (Fig. 5). Only a small part was
treated by air (23,020 hectares). According to DPP, aerial spraying
was carried out on locust hoppers when large areas were involved;
otherwise, ground spraying was considered more effective. As with
other neighboring South Asian countries, Pakistan was unprepared
for the scale of the upsurge (Balakrishnan 2020). One challenge
DPP had to deal with was obsolete or non-functional equipment
for control operations, as the last serious desert locust outbreak was
over 25 years ago (Dowlatchahi et al. 2020b). Small aircrafts for
pesticide spraying were unavailable or not operational. Some also
expressed regret that control operations started too late and that
the federal government did not take the threat seriously enough
from the start in 2019, when locusts were confined to Balochistan
(Ellis-Petersen and Baloch 2020, Nawaz 2020). When operations
finally started, despite the efforts of government authorities, DPP,
local authorities and local inhabitants/farmers, it took 16 months,
from March 2019 to September 2020, for the upsurge to be brought
under control in Pakistan and in the whole eastern region, includ-
ing India and Iran. Ultimately, the control of this upsurge was a
success, which was the result of extensive, strategically planned, and
technically well-executed control operations in the country. Strong
coordination at federal and provincial levels and with all relevant
actors carried out under the National Locust Control Centre set
up in Islamabad increased the effectiveness of the response (Dow-
latchahi et al. 2020b). For all the countries affected by this upsurge,
4,891,150 ha were treated for the years 2019 and 2020 (Table 1).
However, at the start of 2021, the situation remained very worrying
in the Horn of Africa and in the Arabian Peninsula. It is not yet time
to put down our guard, and vigorous monitoring to detect any signs
of breeding desert locust is necessary (Dowlatchahi et al. 2020b).
Table 1. Areas sprayed with pesticides to control the desert lo-
cust upsurge in 2019–2020 over all affected regions (source FAO
2019, 2020a).
Countries ha sprayed Countries ha sprayed Countries ha sprayed
Afghanistan 2969 Jordania 2900 Saudi Arabia 505829
Algeria 1138 Kenya 168484 Somalia 170495
Bahrain 3 Kuwait 15841 Sudan 331368
Egypt 24206 Libya 70 South Sudan 250
Eritrea 113794 Mali 40 UAE 6102
Ethiopia 1177607 Mauritania 10 5 6 Uganda 7154
India 682790 Niger 3897 Yemen 58709
Iran 103 6 510 Oman 13907
Iraq 2610 Pakistan 566390 TOTAL 4891150
Fig. 5. Areas sprayed with pesticides in Pakistan to control the
desert locust upsurge in 2019–2020 (source FAO 2019, 2020a).
In some areas, local governments have announced compensa-
tion measures for farmers who have suffered from locust attacks.
Amid the current COVID-19 pandemic, farmers have found it diffi-
cult to control locusts on their own due to restrictions on transport
and communication. The supply of reliable, affordable pesticides
and spraying equipment has been insufficient. There are no crop
insurance programs in the country, and in some areas, farmers have
had to plant crops twice, as the first crops were completely eaten
up by locusts. Locusts were not only attacking crops, but also dam-
aging rangelands and other vegetation. Thus, livestock keepers and
nomadic communities were also suffering. Such damage was most
visible in arid regions like Balochistan, where rangelands were al-
ready in poor condition. Many affected areas were not treated due
to a lack of small airplanes that can be used for spraying pesticides.
Farmers have been known to adopt different ways to protect their
crops besides insecticide treatments. For instance, one measure taken
by many was beating drums at high volume to scare the locusts. In
some areas, farmers also used smoke from burning bushes and veg-
etation to repel them. According to some local people, since the last
major attack was 58 years ago, the current generation has no direct
experience of handling locusts using local knowledge (Nawaz 2020).
They may not be fully aware of methods to catch locusts or about
how to use them as a food source, a compensatory measure that can
reduce the number of locusts locally and provide a food supplement
to poor and undernourished rural populations (Samejo et al. 2021).
Discussion
The way forward—Institutional aspects and preventative
actions for the future
The situation Pakistan faced in 2019–2020 was the most se-
rious in many years. Nevertheless, desert locust invasions are
now better controlled, being less frequent, less important, and of
shorter duration than in the past (Sword et al. 2010, Zhang et al.
2019). The large invasions that followed one another with a high
frequency ended at the beginning of the 1960s with the establish-
ment of a proactive/preventive strategy and thanks to increasingly
effective surveillance and continuously improving control meth-
ods (Magor et al. 2008, Sword et al. 2010, Lecoq 2019).
Clearly, the problem remains. These invasions are, as always,
the result of exceptionally big rains that occurred in the past and
that are certain to continue to occur. Presently, climate change
cannot be blamed for the ongoing upsurge, even though it will
undoubtedly have consequences for outbreaks of this insect in the
future (Meynard et al. 2020). Thus, these desert locust invasions
will continue to occur. Upsurges over the past 50 years that were
Journal of orthoptera research 2021, 30(2)
R. SULTANA, S. KUMAR, A.A. SAMEJO, S. SOOMRO AND M. LECOQ
152
not stopped at an early stage were the result of gaps or a lack of vig-
ilance in the prevention system implemented at the international
level (Lecoq 2001, 2005). Most often, insecurity in key areas or the
too-late provision of emergency funds are the cause of these up-
surges. Then, control operations start too late, often in countries
that are still poorly prepared, and the swarms disseminate rapidly
(Showler and Lecoq 2021). Therefore, the countries concerned,
and Pakistan in particular, must remain mobilised to improve the
prevention system. Addressing these challenges requires invest-
ment into making the country capable of handling the menace
(Dowlatchahi et al. 2020a, b). For more information on preven-
tion system failures, see Showler (2019) and Showler et al. (2021).
Like other front-line countries, Pakistan contains certain desert
locust outbreak areas, located in desertic areas on the Indo-Pakistan
border and in the Makran region on the border with Iran (Symmons
and Cressman 2001, Cressman 2016). Pakistan, therefore, plays a
key role in the prevention strategy by conducting regular surveil-
lance of these areas. However, the country can also be invaded by
swarms that originate outside its borders, as was the case in 2019.
International cooperation is essential to better control these migra-
tory insects. As mandated by its Member States, the FAO ensures the
coordination of monitoring and control activities of the desert lo-
cust on an international scale (Lecoq 2003). Via its Desert Locust
Information Service (DLIS), it issues a monthly locust situation and
forecast bulletin (FAO 2009b). This bulletin is based on reports from
the affected countries, as well as on the analysis of the ecological
conditions in the habitats of the locust (using satellite remote sens-
ing data, weather reports related to rains and direction of the winds,
etc.) (Cressman 2008, 2013). In addition, the FAO provides a forum
for the meetings of the Desert Locust Control Committee (DLCC),
formed by representatives from all the countries affected by desert
locust as well as those that take part in locust control campaigns.
Since 1955, Pakistan has been a member of both the DLCC
and the South West Asia Commission (SWAC), established in
1964 under Article XIV of the FAO Constitution. SWAC has four
member states: Afghanistan, India, Iran, and Pakistan. All activi-
ties of SWAC contribute to the strengthening of the national ca-
pacities of its member countries in desert locust survey, control
operations, reporting, training, preparedness, contingency plan-
ning, emergency response, biopesticides, and health and safety
(FAO 2021a). SWAC also promotes cooperation among its mem-
ber countries and, in particular, the conduct of regular joint sur-
veys for desert locust surveillance and early warning, as well as
the exchange of information on the locust situation. It supports
training and capacity building activities and the promotion of
new technologies (FAO 2021a). SWAC is also collaborating with
the other two FAO commissions—CRC for the central region and
CLCPRO for the western region (FAO 2021b, c)—with regard to
the use of biopesticides, the development of risk management
plans, setting up inventory systems, and environmental moni-
toring. Obviously, all of these measures should be developed at
the level of each member state, and it is in Pakistan’s interest to
strengthen its cooperation with SWAC.
As a result, monitoring and preventative organization against
desert locust invasions is a leading example in the field of crop
protection (Hamouny 2021). However, while this prevention sys-
tem is genuinely effective, it also has its flaws, as the current situ-
ation unfortunately reminds us. Various reasons for failure have
been given: inexperience of field survey teams and campaign or-
ganizers, insufficient or inappropriate resources, inaccessibility of
some important breeding areas for security reasons, deterioration
of survey, and control capacities during recession periods (WMO
& FAO 2016). Some issues, such as security concerns, are beyond
Pakistan’s control. Others may find solutions locally. We give de-
tails on three main points below.
1. Desert locust require concerted monitoring and on-the-
ground control effort across borders, along with the resources, ex-
pertise, and infrastructure to support those actions. Moreover, these
efforts must be kept in place over the long term to build resilience,
despite the apparent lack of imminent threats (Lecoq 1991, Gay et
al. 2018). The lack of such coordinated and sustained efforts is like-
ly to put human populations at higher risk. Pakistan had to fight
the recent upsurge with insufficient funding, operational resources,
obsolete equipment, and an eroded expertise that left the DPP with
only a few high-level experts (Dowlatchahi et al. 2020a, b). The
highest priorities should therefore be (a) to ensure that the politi-
cal and socio-economic conditions are in place so that vulnerable
human populations can adapt to new large-scale threats and (b)
to maintain a long-term risk assessment culture with ongoing fi-
nancial, material, and expertise support (Meynard et al. 2020). Per-
haps maintaining funding mechanisms that provide sustainable
support during periods of recession, when priorities are elsewhere,
is one of the most difficult but key points to be solved. Yet, it has
been shown that funding institutions (governments, donors) could
considerably improve the effectiveness of the prevention system by
increasing their support by only a few percent (Gay et al. 2018).
2. Pakistan, as with all countries concerned with the desert
locust, must remain ready and develop compensatory measures
for the local populations in the event of an invasion that is not
controlled early on. Farmers are most often helpless in the face
of the threat from locusts. Prevention remains the best rampart,
but if this fails, local populations must have access to informa-
tion, advice, and support, both technical and financial. It is advis-
able, for instance, to develop desert locust control material for the
education of farmers and agriculture extension staff and organize
farmers’ schools for desert locust control.
3. Finally, current treatments are based almost exclusively
on traditional chemical insecticides that pose various risks to both
human health and the environment (Everts and Ba 1997, Sam-
ways and Lockwood 1998, van der Valk 1998, Peveling 2001, FAO
2014). The end result of these quick control measures is still mas-
sive damage, both to crops and, perhaps worse, to the ecosystem,
from the enormous amounts of pesticides sprayed (Balakrishnan
2020). Alternative products such as mycopesticides, which have
been used for some time by countries such as Australia and China,
should be able to find a larger audience globally (Lomer et al.
2001, Hunter 2004, 2010, Zhang and Hunter 2005, Zhang 2011).
These products are currently commercially available, and the
ongoing upsurge has given rise to their use in various countries
(Zhang et al. 2019). In 2020, mycopesticides were successfully ap-
plied to at least 10,845 ha in Somalia against desert locusts (FAO
2020g, h). In collaboration with the FAO, trials focused on the
introduction of mycopesticides in Pakistan have been done, which
should obviously be encouraged.
Acknowledgments
We greatly appreciate the constructive and valuable support of
Dr. M. Tariq Khan, Director Technical, Department of Plant Pro-
tection Karachi, for providing updated information and data. This
study was funded by the Higher Education Commission Islama-
bad, Pakistan (Project No. 6737 SINDH /NRPU /R & D/ HEC).
The authors also thank the Orthopterists’ Society for supporting
the publication of this article.
R. SULTANA, S. KUMAR, A.A. SAMEJO, S. SOOMRO AND M. LECOQ 153
Journal of orthoptera research 2021, 30(2)
References
Balakrishnan R (2020) India. Regional Reports. Metaleptea 40: 7–8.
https://orthsoc.org/2020/09/17/new-metaleptea-issue-403 [Upload-
ed 4 February 2021]
Brader L, Djibo H, Faye FG, Ghaout S, Lazar M, Luzietoso PN, Ould Ba-
bah MA (2006) Towards a more effective response to desert locusts
and their impacts on food security, livelihoods and poverty: multi-
lateral evaluation of the 2003–05 Desert Locust campaign. Food and
Agriculture Organization of the United Nations, Rome, 92 pp. http://
www.fao.org/ag/locusts/common/ecg/1913/en/DesertLocustEval-
ReportE.pdf [Accessed on 23 February 2021]
Cisse S, Ghaout S, Mazih A, Ould Babah Ebbe MA, Benahi AS, Piou C
(2013) Effect of vegetation on density thresholds of adult desert locust
gregarization from survey data in Mauritania. Entomologia Experi-
mentalis et Applicata 149: 159–165. https://doi.org/10.1111/eea.12121
Collett M, Despland E, Simpson SJ, Krakauer DC (1998) Spatial scales of de-
sert locust gregarization. Proceedings of the National Academy of Scienc-
es USA 95: 13052–13055. https://doi.org/10.1073/pnas.95.22.13052
COPR (1982) The Locust and Grasshopper Agricultural Manual. Centre
Overseas Pest Research, London, 690 pp.
Cressman K (2001) Desert Locust Guidelines 2. Survey. Food and Agricul-
ture Organization of the United Nations, Rome, 56 pp. http://www.
fao.org/ag/locusts/common/ecg/347_en_DLG2e.pdf [Uploaded 21
February 2021]
Cressman K (2008) The use of new technologies in desert locust early
warning. Outlooks on Pest Management 19: 55–59. https://doi.
org/10.1564/19apr03
Cressman K (2013) Role of remote sensing in desert locust early warn-
ing. Journal of Applied Remote Sensing 7: 075098. https://doi.
org/10.1117/1.JRS.7.075098
Cressman K (2016) Desert Locust. In: Shroder JF, Sivanpillai R (Eds) Biolog-
ical and Environmental Hazards, Risks, and Disasters. Elsevier, Amster-
dam, 87–105. https://doi.org/10.1016/B978-0-12-394847-2.00006-1
Despland E, Collett M, Simpson SJ (2000) Small-scale processes in de-
sert locust swarm formation: how vegetation patterns influence
gregarization. Oikos 88: 652–662. https://doi.org/10.1034/j.1600-
0706.2000.880322.x
De Vreyer P, Guilbert N, Mesple-Sompsa S (2014) Impact of natural dis-
asters on education outcomes: evidence from the 1987–89 locust
plague in Mali. Journal of African Economies 24: 57–100. https://doi.
org/10.1093/jae/eju018
Dowlatchahi M, Mubarik A, Cressman K (2020a) Desert locust situation
in Pakistan. Food and Agriculture Organization of the United Na-
tions, Rome. http://www.fao.org/pakistan/resources/in-depth/desert-
locust-situation-in-pakistan/en/ [Uploaded 4 February 2021]
Dowlatchahi M, Cressman K, Aamer I, Mubarik A (2020b) The Desert Lo-
cust upsurge in Southwest Asia has been stopped! Food and Agricul-
ture Organization of the United Nations, Rome. http://www.fao.org/
pakistan/resources/in-depth/the-desert-locust-upsurge-in-southwest-
asia-has-been-stopped/en/ [Uploaded 4 February 2021]
Duranton JF, Lecoq M (1990) Le Criquet pèlerin au Sahel. Comité Inter-
Etats de Lutte contre la Sécheresse dans le Sahel, Département de For-
mation en Protection des Végétaux, Niamey, 183 pp.
Ellis-Petersen H, Baloch SM (2020) ‘Many will starve’: locusts devour crops
and livelihoods in Pakistan. The Guardian. https://www.theguard-
ian.com/world/2020/may/25/many-will-starve-locusts-devour-crops-
and-livelihoods-in-pakistan [Uploaded 4 February 2021]
Everts JW, Ba L (1997) Environmental effects of locust control: state of
the art and perspectives. In: Krall S, Peveling R, Diallo DB (Eds) New
Strategies in Locust Control. Birkhäuser Verlag, Basel, 331–336. htt-
ps://doi.org/10.1007/978-3-0348-9202-5_49
FAO (1993) Desert locust bulletins, Nos. 179–183. Food and Agriculture Or-
ganization of the United Nations, Rome. http://www.fao.org/ag/locusts/
en/archives/archive/1367/1368/index.html [Uploaded 7 May 2021]
FAO (1997) Desert locust bulletins, Nos. 229–230. Food and Agriculture Or-
ganization of the United Nations, Rome. http://www.fao.org/ag/locusts/
en/archives/archive/1367/1997/index.html [Uploaded 7 May 2021]
FAO (2004) Desert locust bulletins, Nos. 303–314. Food and Agriculture Or-
ganization of the United Nations, Rome. http://www.fao.org/ag/locusts/
en/archives/archive/1366/2004/index.html [Uploaded 7 May 2021]
FAO (2005) Desert locust bulletins, Nos. 315–318. Food and Agriculture Or-
ganization of the United Nations, Rome. http://www.fao.org/ag/locusts/
en/archives/archive/1366/2005/index.html [Uploaded 7 May 2021]
FAO (2009a) Glossary on desert locust. Food and Agriculture Organiza-
tion of the United Nations, Rome, 334 pp. http://www.fao.org/3/
as983b/as983b.pdf [Uploaded 15 March 2021]
FAO (2009b) Locust watch. Desert locust. Food and Agriculture Organiza-
tion of the United Nations, Rome. http://www.fao.org/ag/locusts/en/
info/info/index.html [Uploaded 4 February 2021]
FAO (2014) Evaluation of field trials data on the efficacy and selectivity of
insecticides on locusts and grasshoppers: report to FAO by the Pesti-
cide Referee Group. Food and Agriculture Organization of the United
Nations, Rome, 66 pp. http://www.fao.org/ag/locusts/common/
ecg/2241/en/PRG10e.pdf [Uploaded 4 February 2021]
FAO (2016) Pakistan. Country fact sheet on food and agriculture policy
trends. Food and Agriculture Organization of the United Nations, Rome.
http://www.fao.org/3/i6054e/i6054e.pdf [Uploaded 7 May 2021]
FAO (2018) Desert locust bulletin, Nos. 472–483. Food and Agriculture
Organization of the United Nations, Rome. http://www.fao.org/ag/lo-
custs/en/archives/archive/1823/2415/index.html [Uploaded 29 Janu-
ary 2021]
FAO (2019) Desert Locust Bulletin, Nos. 484–495. Food and Agriculture Or-
ganization of the United Nations, Rome. http://www.fao.org/ag/locusts/
en/archives/archive/1823/index.html [Uploaded 29 January 2021]
FAO (2020a) Desert Locust Bulletin, Nos. 496–507. Food and Agriculture
Organization of the United Nations, Rome. http://www.fao.org/ag/lo-
custs/en/archives/archive/2521/index.html [Uploaded 2 February 2021]
FAO (2020b) Desert Locust upsurge in 2019–2020. Food and Agriculture
Organization of the United Nations, Rome. http://www.fao.org/ag/
locusts/en/info/2094/web18/index.html [Uploaded 29 January 2021]
FAO (2020c) Desert locust bulletin, 4 April 2020, No. 498. Food and Ag-
riculture Organization of the United Nations, Rome. http://www.fao.
org/ag/locusts/common/ecg/562/en/DL498e.pdf [Uploaded 24 Feb-
ruary 2021]
FAO (2020d) The Desert Locust Upsurge 2019–2020 in Pakistan. Food and
Agriculture Organization of the United Nations, Rome. http://www.
fao.org/ag/locusts/en/info/2094/2537/PAK/index.html [Uploaded 2
February 2021]
FAO (2020e) Pakistan readies for second battle against crop-devouring lo-
custs. https://reliefweb.int/report/pakistan/pakistan-readies-second-
battle-against-crop-devouring-locusts [Uploaded 4 February 2021]
FAO (2020f) Pakistan: Further desert locust damage forecast in coming
agricultural seasons. https://www.unocha.org/story/pakistan-further-
desert-locust-damage-forecast-coming-agricultural-seasons [Upload-
ed 4 February 2021]
FAO (2020g) Desert locust bulletin, 4 May 2020, No. 499. Food and Ag-
riculture Organization of the United Nations, Rome. http://www.fao.
org/ag/locusts/common/ecg/2551/en/DL499e.pdf [Uploaded 3 No-
vember 2020]
FAO (2020h) Desert locust bulletin, 4 Jun 2020, No. 500. Food and Agri-
culture Organization of the United Nations, Rome. http://www.fao.
org/ag/locusts/common/ecg/2555/en/DL500e.pdf [Uploaded 3 No-
vember 2020]
FAO (2021a) FAO Commission for Controlling the Desert Locust in South-
West Asia (SWAC). http://www.fao.org/ag/locusts/swac/swac-home/
en/ [Uploaded 6 February 2021]
FAO (2021b) The Commission for Controlling the Desert Locust in the Cen-
tral Region. http://desertlocust-crc.org/ [Uploaded 6 February 2021]
FAO (2021c) Commission de lutte contre le Criquet pèlerin dans la région
occidentale. http://www.fao.org/clcpro/apercu/historique/fr/ [Up-
loaded 6 February 2021]
Forskål (1775) Descriptiones Animalium Avium, Amphibiorum, Piscium,
Insectorum, Vermium; quae in Itinere Orientall observati Petrus For-
skal. Prof. Haun. Post morten Acutoris editt Carsten Nieburhr., Hau-
niae, 164 pp. https://doi.org/10.5962/bhl.title.2154
Journal of orthoptera research 2021, 30(2)
R. SULTANA, S. KUMAR, A.A. SAMEJO, S. SOOMRO AND M. LECOQ
154
Gay PE, Lecoq M, Piou C (2018) Improving preventive locust manage-
ment: insights from a multi-agent model. Pest Management Science
74: 46–58. https://doi.org/10.1002/ps.4648
Hamouny ML (2021) Commission for Controlling the Desert Locust in the
Western Region (CLCPRO) – A Success Story. Metaleptea 41: 20–21.
http://orthsoc.org/wp-content/uploads/2021/01/Metaleptea_41_1.
pdf? [Uploaded 6 February 2021]
Hunter DM (2004) Advances in the control of locusts (Orthoptera: Acrid-
idae) in eastern Australia: from crop protection to preventive con-
trol. Australian Journal of Entomology 43: 293–303. https://doi.
org/10.1111/j.1326-6756.2004.00433.x
Hunter DM (2010) Credibility of an IPM approach for locust and grass-
hopper control: the Australian example. Journal of Orthoptera Re-
search 19: 133–137. https://doi.org/10.1665/034.019.0108
Lecoq M (1991) Le Criquet pèlerin. Enseignements de la dernière inva-
sion et perspectives offertes par la biomodélisation. In: Essaid A
(Ed.) La lutte anti-acridienne. AUPELF-UREF, John Libbey Eurotext,
Paris, 71–98.
Lecoq M (2001) Recent progress in desert and migratory locust man-
agement in Africa: are preventative actions possible? Journal of
Orthoptera Research 10: 277–291. https://doi.org/10.1665/1082-
6467(2001)010[0277:RPIDAM]2.0.CO;2 [Uploaded 20 October 2020]
Lecoq M (2003) Desert locust threat to agricultural development and
food security and FAO’s international role in its control. Arab Jour-
nal of Plant Protection 21: 188–193. https://asplantprotection.org/
wp-content/uploads/2018/07/V21-2_188-193.pdf [Uploaded 3
November 2020]
Lecoq M (2004) Vers une solution durable au problème du criquet pèler-
in? Science et changements planétaires / Sécheresse 15: 217–224.
Lecoq M (2005) Desert locust management: from ecology to anthro-
pology. Journal of Orthoptera Research 14: 179–186. https://doi.
org/10.1665/1082-6467(2005)14[179:DLMFET]2.0.CO;2
Lecoq M (2019) Desert Locust Schistocerca gregaria (Forskål, 1775)
(Acrididae). In: Lecoq M, Zhang L (Eds) Encyclopedia of Pest Or-
thoptera of the World. China Agricultural University Press, Beijing,
204–212.
Lomer CJ, Bateman RP, Johnson DL, Langewald J, Thomas M (2001)
Biological control of locusts and grasshoppers. Annual Review
of Entomology 46: 667–701. https://doi.org/10.1146/annurev.
ento.46.1.667
Magor JI, Lecoq M, Hunter DM (2008) Preventive control and De-
sert Locust plagues. Crop Protection 27: 1527–1533. https://doi.
org/10.1016/j.cropro.2008.08.006
Martini P, Lecoq M, Soumaré L, Chara B (1998) Proposition de programme
de lutte contre le criquet pèlerin dans la partie occidentale de son aire
d’habitat. Food and Agriculture Organization of the United Nations,
Rome, 92 pp. http://www.fao.org/ag/locusts/oldsite/PDF/meetings/
EMPRESwest98f.pdf [Uploaded 20 October 2020]
Meynard CN, Lecoq M, Chapuis MP, Piou C (2020) On the relative role
of climate change and management in the current Desert Locust out-
break in East Africa. Global Change Biology 26: 3753–3755. https://
doi.org/10.1111/gcb.15137
Nawaz K (2020) Locust attacks in Pakistan. The Water Channel. https://
thewaterchannel.tv/thewaterblog/locust-attacks-in-pakistan/ [Up-
loaded 4 February 2021]
Pener MP, Simpson SJ (2009) Locust phase polyphenism: An update.
Advances in Insect Physiology 36: 1–272. https://doi.org/10.1016/
S0065-2806(08)36001-9
Piou C, Jaavar Bacar M, Babah Ebbe MAO, Chihrane J, Ghaout S, Cisse S,
Lecoq M, Ben Halima T (2017) Mapping the spatiotemporal distribu-
tions of the Desert Locust in Mauritania and Morocco to improve pre-
ventive management. Basic and Applied Ecology 25: 37–47. https://
doi.org/10.1016/j.baae.2017.10.002
Peveling R (2001) Environmental conservation and locust control—possible
conflicts and solutions. Journal of Orthoptera Research 10: 171–187.
https://doi.org/10.1665/1082-6467(2001)010[0171:ECALCP]2.0.CO;2
Samejo AA, Sultana R, Kumar S, Soomro S (2021) Could entomophagy be
an effective mitigation measure in desert locust management? Agron-
omy 11: 455. https://doi.org/10.3390/agronomy11030455
Samways MJ, Lockwood JA (1998) Orthoptera conservation: pests and
paradoxes. Journal of Insect Conservation 2: 143–149. https://doi.
org/10.1023/A:1009652016332
Showler AT (2019) Desert locust control. The effectiveness of proactive
interventions and the goal of outbreak prevention. American Ento-
mologist 65: 180–191. https://doi.org/10.1093/ae/tmz020
Showler AT, Lecoq M (2021) Incidence and ramifications of armed conflict
in countries with major desert locust breeding areas. Agronomy 11:
114. https://doi.org/10.3390/agronomy11010114
Showler AT, Ould Babah Ebbe MA, Lecoq M, Maeno KO (2021) Early
intervention against desert locusts: Current proactive approach and
the prospect of sustainable outbreak prevention. Agronomy 11: 312.
https://doi.org/10.3390/agronomy11020312
Steedman A (1990) Locust Handbook. 2nd edn. Overseas Development
Natural Resources Institute, London, 204 pp.
Sword GA, Lecoq M, Simpson SJ (2010) Phase polymorphism and preven-
tative locust management. Journal of Insect Physiology 56: 949–957.
https://doi.org/10.1016/j.jinsphys.2010.05.005
Symmons PM, Cressman K (2001) Desert Locust Guidelines 1. Biology
and behaviour. Food and Agriculture Organization of the United
Nations, Rome, 43 pp. http://www.fao.org/ag/locusts/common/
ecg/347_en_DLG1e.pdf [Uploaded 4 February 2021]
Uvarov BP (1921) A revision of the genus Locusta, L. (=Pachytylus, Fieb.),
with a new theory as to the periodicity and migrations of locusts.
Bulletin of Entomological Research 12: 135–163. https://doi.
org/10.1017/S0007485300044989
Uvarov BP (1966) Grasshoppers and Locusts, vol. 1. Cambridge University
Press, Cambridge, 481 pp.
van der Valk H (1998) The impact of locust and grasshopper control on
beneficial arthropods in West Africa. In: Haskell PT, McEwen P (Eds)
Ecotoxocology: Pesticides and Beneficial Organisms. Chapman & Hall,
London, 372–380. https://doi.org/10.1007/978-1-4615-5791-3_40
WMO & FAO (2016) Weather and Desert Locusts. World Meteorological
Organization and Food and Agriculture Organization of the United
Nations, Geneva, 38 pp.
Zhang L (2011) Advances and prospects of strategies and tactics of locust
and grasshopper management. Chinese Journal of Applied Entomol-
ogy 48: 804–810. [In Chinese]
Zhang L, Hunter DM (2005) Laboratory and field trials of Green
Guard Metarhizium anisopliae var. acridum (Deuteromycotina: Hy-
phomycetes) against the oriental migratory locust (Locusta mi-
gratoria manilensis) (Orthoptera: Acrididae) in China. Journal of
Orthoptera Research 14: 27–30. https://doi.org/10.1665/1082-
6467(2005)14[27:LAFTOG]2.0.CO;2 [Uploaded 4 February 2021]
Zhang L, Lecoq M, Latchininsky A, Hunter D (2019) Locust and grasshop-
per management. Annual Review of Entomology 64: 15–34. https://
doi.org/10.1146/annurev-ento-011118-112500
