These ecosystems are subject to expanding agriculture, urbanization, energy development, and desertification, making them among the most threatened biomes on Earth Hoekstra et al. For some mix and grass-feeder species like the Moroccan locust or the red locust, this increase in anthropogenic change, alongside modern control practices, has led to a decrease in outbreak frequency Thindwa, ; Benfekih et al.
This also appears to have been the case for the Rocky Mountain grasshopper, Melanoplus spretus , which destroyed vast quantity of crops throughout the western and central portion of the United States and Canada during the mids.
The species declined rapidly in the late s and agricultural practices likes tillage, irrigation, introduction of cattle, plants, and birds into grasslands are thought to be responsible for its extinction Lockwood and Debrey, However, in many other cases because cropland expansion includes cereals that are readily consumed by mix- and grass-feeders, it yielded an increase in outbreak frequencies, as seen with the Bombay locust Bullen, , the spur-throated locust Casimir and Edge, , and the Central American locust Poot-Pech, Table 1.
Worldwide Locust Species, adapted from Cullen et al. One mechanistic link between livestock grazing and locust populations is the conversion of wooded areas to open fields, including patchy grasslands, more desirable to locusts Clark, The Mongolian locust, Oedaleus asiaticus , was rarely reported to have economically important outbreaks prior to the 's; increased outbreaks may coincide with increased livestock populations in Inner Mongolia, China Robinson et al.
Thus, land use and cover change has potentially contributed to the emergence of new locust species that previously did not exhibit strong locust phase change characteristics, likely in conjunction with shifting climate. For instance, the Senegalese grasshopper was not reported as a pest until it first started outbreaking in the 's and is now considered the main pest of millet in the Sahel, exhibiting locust characteristics like density-dependent color change and migrations Popov, ; Maiga et al.
Similarly, the Peruvian locust was only known as a non-swarming grasshopper until an upsurge in Peru in —84, subsequently gregarized individuals invaded several Peruvian states from to Duranton et al. Increasing grazing pressure and agricultural intensity may inadvertently be creating a nutritionally optimal environment for locusts through soil degradation. In addition to presenting bare areas favorable to egg laying and thermoregulation, degraded pastures may harbor plants with low protein:carbohydrate ratios that are favorable to locusts.
Soil erosion typically decreases soil nitrogen resulting in low nitrogen plants for a mechanistic description between grazing and plant nitrogen, see O. Since most nitrogen in plants, particularly grasses, is found in the form of protein, and protein and carbohydrate tend to be negatively correlated, this translates into low protein, high carbohydrate plants Loaiza et al.
In contrast to the N-limitation hypothesis White, , emerging evidence suggests that locusts are more likely to be carbohydrate-limited in field populations. For example, marching bands of S. Over recent decades, many lab studies have shown multiple locust species select a carbohydrate-biased diet when given the choice and show higher performance around their self-selected low protein, high carbohydrate diet Raubenheimer and Simpson, ; Chambers et al.
There are multiple factors that might explain why carbohydrate-biased diets are beneficial for locusts. First, carbohydrate-biased diets may be a requirement for long-distance flight Hunter et al. A defining characteristic of locusts is their capacity to undergo spectacular migrations: swarms can fly hundreds of kilometers in a single night and plagues can span continents. These long-distance flights are fueled primarily from lipid reserves Weis-Fogh, ; Jutsum and Goldsworthy, , which are built with dietary carbohydrates Walker et al.
Insect flying metabolic rates can be times that of resting animals and are among the highest known Rankin and Burchsted, This high energy demand means their food source pre-and during migration is critical to fueling flight. Second, carbohydrate-biased diets may be important for resistance against parasites and pathogens. For instance, Australian plague locusts are more likely to die of fungal infection when they are restricted to diets high in protein than when they are restricted to diets high in carbohydrates Graham et al.
Potentially because fungus are better at exploiting protein in the insect's hemolymph, or perhaps because immune function is correlated with lipid levels of the host Graham et al. Finally, these diets may help locusts persist in an arid environment because consuming more carbohydrate enhances lipid stores, which can in turn be used as a water reserve, as shown in the migratory locust Loveridge and Bursell, Although the link between land use, plant nutrient content, and locust performance has only experimentally been demonstrated for O.
The connection between plant nutritional landscape and locust outbreaks is thus a promising mechanism to explain the association between agricultural practices and locusts, and is an area for suggested research, particularly for non-model locusts see Table 1. Furthermore, the presence of locusts in grazed areas leads to strong competition between locusts and livestock, particularly for mix- and grass-feeders.
Even tree-feeders, like the Sahelian tree locust, were reported to compete with livestock by attacking shrubs that are a food source for domestic cattle and camels COPR, This competition for resources between livestock and locusts has the potential to further degrade soils in over-grazed areas as shown for the brown locust in South Africa Lea, a , b ; De Villiers, ; Price and Brown, The interactions between decisions regarding land use and locust biology has rippling effects: increased grazing pressure creates optimal environments for locusts, leading locusts to reach higher densities, which promotes gregarization.
Gregarization in turn increases locust diet breadth, leading to expansive and diverse crop damage COPR, ; Despland, One potential explanation for the diet expansion is that gregarious locusts may be more likely to encounter a variety of host plants with varying nutrient contents and, by eating from among these diverse food sources, they can balance nutrient requirements Simpson et al.
Another explanation is eating toxic plants can serve as an anti-predatory strategy, which is linked to the development of density-dependent aposematic coloration seen in some locust species e. Exploring how gregarious locusts expand their diet breadth is another understudied area. In many cases livestock grazing has positive effects on locust populations, which can lead to food insecurity.
However, grasshoppers, including locusts, are dominant herbivores and vital players in grassland ecosystems Branson et al. Furthermore, chemical control of grasshoppers and locusts has many undesirable side-effects on livestock health Botha, ; Tanaka and Zhu, , human health Houndekon and De Groote, ; Tingle et al.
Therefore, management programs that rely on population monitoring and early intervention, such as chemical treatment as soon as populations start to increase, are more likely to be sustainable Van Huis et al. For instance, a combination of satellite data to estimate rainfall and vegetation cover, and field survey data to indicate locust outbreaks, can be used to develop predictive models Cressman, ; Piou et al. These models assist managers in determining the probability of finding locusts in specific areas to more efficiently focus survey efforts.
With a greater understanding of the connections between agricultural practices and locust outbreaks, cultural control options may be viable components of future management programs. For example, habitat manipulation can slow nymphal development, reduce survival and reproduction, or decrease yearly variability in those life history traits Onsager and Olfert, ; Branson et al. Manipulations to limit pest damage can include direct competition from mammalian herbivores such as livestock for grasshoppers Onsager, , altering food quality for terrestrial herbivores Le Gall and Tooker, , manipulating plant composition, and increasing the abundance of natural enemies of grasshoppers Belovsky, ; Joern, These types of management practices are less developed for locusts but harbor great potential.
For example, in West Africa, fallow fields often have more Senegalese grasshoppers than other field types Toure et al. Due to limited land availability, fields may be left fallow only when the soil has been depleted and yield is limited, resulting in a low-N environment and carbohydrate-biased plants. Because locusts require a particularly carbohydrate-biased diet Table 1 , bottom up control using knowledge of locust nutritional requirements may be a viable option Le Gall and Tooker, ; Word et al.
Practices that improve soil fertility, such as reducing grazing pressure in Inner Mongolia, China Cease et al. However, livestock and farm managers may not have the resources, knowledge, or incentives to implement practices to decrease locust populations.
Therefore, cultural control options should be developed iteratively, in ways that integrate stakeholder perspectives, as well as the natural and social science Cullen et al. Such research linking land use to locust populations has revealed that this ecological feedback connects people across time and space producing both local and far-reaching environmental and social impacts Cease et al.
At a regional scale, locusts can spill from rangelands into croplands. For instance, the Australian plague locust is typically associated with grazing areas but during outbreaks will attack agricultural crops Watts et al. Decisions about stocking rates in one region, for example, have the potential to initiate outbreaks that could lead to swarms invading distant regions.
In addition to ecological links, distant regions are coupled through the economy e. Indeed, because locusts connect distant regions and span continents, a major challenge is implementing consistent management efforts across broad regions Cressman, ; Zhang et al.
This requires many countries to work together from the level of the producer to national and international government organizations Lockwood et al. This review highlights that humans are not passive players in their relationship with locusts. To effectively integrate land-use into management programs, important data on locust biology, such as nutritional preference and thermobiology, remain to be collected, particularly for non-model locust species, for which the expressions of density-dependent phase polyphenism in not well-understood Song, In addition, this biology research needs to be linked to broader landscape patterns and their management, including producer decisions as well as market and government constraints.
ML wrote the discussion and sections for nine species and oversaw paper cohesion. RO performed the initial literature review and wrote the method section and sections for seven species.
AC conceptualized the framework for the paper, wrote the introduction and sections for three species and edited all sections for consistency. All authors participated in revisions of the whole manuscript. The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
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They develop strong wings and an insatiable hunger, and become "gregarious", hoarding together in giant swarms as big as 10km x 10km 6 miles , and as dense as 50 locusts per cubic metre. If they end up out at sea, with no source of food, they eat each other. None of this gives locusts a good reputation, but "they taste very nice, I can tell you! Van Huis says it is completely normal for people in places that suffer from locust plagues to eat them. Insects of all sorts are eaten in most African countries, he says - and for good reason.
Locusts are a valuable source of income for women in Niger, who get up early to collect them from the millet fields, and then sell them at the market.
They make more money from the locusts on the millet than the millet itself, says van Huis. He argues that we are heading towards a meat crisis, and insects are an eco-friendly, nutritious, alternative. Locusts are high in protein, and also zinc and iron - minerals which many people around the world are lacking - and they emit very little in the way of greenhouse gases. But van Huis says it's impossible to eat your way out of a locust invasion. For one thing - there are simply too many.
And in a country like Israel, where they are being sprayed very heavily with insecticide, they could be contaminated, and he would advise against eating them. Although locusts are clearly described in the Torah as being kosher, there is much discussion in Israel, says Rabbi Ari Zivotofsky, as to whether all Jews can eat them - or only those Jews who have a tradition of eating them principally Yemenite Jews and those from North Africa.
Zivotofsky believes the only reason that Ashkenazi Jews don't have the tradition of eating locusts is because it's extremely rare to get locusts in Europe, so he has no problem - in principle, at least - with eating them. He organises special dinners encouraging Israelis to widen their horizons and try traditional foods that, although kosher, are rarely eaten. He's had locust on the menu before and it proved a hit. His children certainly enjoyed their own culinary creations, particularly the breaded specimens.
You can follow the Magazine on Twitter and on Facebook. Eucalyptus restaurant Jerusalem. Food Insects Newsletter. Locust swarms devastate crops and cause major agricultural damage, which can lead to famine and starvation. Locusts occur in many parts of the world, but today locusts are most destructive in subsistence farming regions of Africa. The desert locust Schistocerca gregaria is a notorious species.
Found in Africa, the Middle East, and Asia, this species inhabits an area of about six million square miles, or 30 countries, during a quiet period. During a plague, when large swarms descend upon a region, however, these locusts can spread out across some 60 countries and cover a fifth of Earth's land surface. Desert locust plagues threaten the economic livelihood of a tenth of humans. A desert locust swarm can be square miles in size and pack between 40 and 80 million locusts into less than half a square mile.
Each locust can eat its weight in plants each day, so a swarm of such size would eat million pounds of plants every day. To put it into context, a swarm the size of Paris can eat the same amount of food in one day as half the population of France. But experts can look at past weather patterns and historical records to identify the areas where swarms might occur and spray those areas with chemicals.
Some experts worry that locust plagues will worsen in a warming world. Rising sea temperatures are causing prolonged bouts of wet weather, including a surge of rare cyclones in eastern Africa and the Arabian Peninsula where desert locusts thrive.
All rights reserved. Common Name: Locusts. Scientific Name: Acrididae. Type: Invertebrates. Diet: Herbivore. Group Name: Swarm. Average Life Span: Several months. Size: 0.
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