Grasshopper

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Grasshoppers
Temporal range: Induan–Holocene, 250–0  Ma
American Bird Grasshopper.jpg
American grasshopper ( Schistocerca americana )
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Orthoptera
Suborder: Caelifera
Infraorder: Acrididea
Informal group: Acridomorpha
Dirsh, 1966
Superfamilies

Grasshoppers are a group of insects belonging to the suborder Caelifera. They are amongst what are possibly the most ancient living groups of chewing herbivorous insects, dating back to the early Triassic around 250 million years ago.

Contents

Grasshoppers are typically ground-dwelling insects with powerful hind legs which allow them to escape from threats by leaping vigorously. Their front legs are shorter and used for grasping food. As hemimetabolous insects, they do not undergo complete metamorphosis; they hatch from an egg into a nymph or "hopper" which undergoes five moults, becoming more similar to the adult insect at each developmental stage. [1] The grasshopper hears through the tympanal organ which can be found in the first segment of the abdomen attached to the thorax; while its sense of vision is in the compound eyes, a change in light intensity is perceived in the simple eyes (ocelli). At high population densities and under certain environmental conditions, some grasshopper species can change colour and behavior and form swarms. Under these circumstances, they are known as locusts.

Grasshoppers are plant-eaters, with a few species at times becoming serious pests of cereals, vegetables and pasture, especially when they swarm in the millions as locusts and destroy crops over wide areas. They protect themselves from predators by camouflage; when detected, many species attempt to startle the predator with a brilliantly coloured wing flash while jumping and (if adult) launching themselves into the air, usually flying for only a short distance. Other species such as the rainbow grasshopper have warning coloration which deters predators. Grasshoppers are affected by parasites and various diseases, and many predatory creatures feed on both nymphs and adults. The eggs are subject to attack by parasitoids and predators. Grasshoppers are diurnal insects, meaning they are most active during the day time.

A Grey Bird Grasshopper at Mungo National Park. Grasshopper at Mungo.jpg
A Grey Bird Grasshopper at Mungo National Park.

Grasshoppers have had a long relationship with humans. Swarms of locusts can have devastating effects and cause famine, having done so since Biblical times. [2] Even in smaller numbers, the insects can be serious pests. They are used as food in countries such as Mexico and Indonesia. They feature in art, symbolism and literature. The study of grasshopper species is called acridology.

Phylogeny

Grasshoppers belong to the suborder Caelifera. Although "grasshopper" has been used as a common name for the suborder in general, [3] [4] [5] modern sources restrict it to the more "evolved" families. [6] They may be placed in the infraorder Acrididea [7] and have been referred to as "short-horned grasshoppers" in older texts [8] to distinguish them from the also-obsolete term "long-horned grasshoppers" (now bush-crickets or katydids) with their much longer antennae. The phylogeny of the Caelifera, based on mitochondrial ribosomal RNA of thirty-two taxa in six out of seven superfamilies, is shown as a cladogram. The Ensifera (crickets, etc.), Caelifera and all the superfamilies of grasshoppers except "Pamphagoidea" appear to be monophyletic. [9] [10]

Orthoptera
Ensifera  (crickets, katydids, etc.)

[6 superfamilies] Gryllus campestris MHNT.jpg

Caelifera
Fossil grasshoppers at the Royal Ontario Museum Fossil grasshoppers - Royal Ontario Museum - DSC00013.JPG
Fossil grasshoppers at the Royal Ontario Museum

In evolutionary terms, the split between the Caelifera and the Ensifera is no more recent than the Permo-Triassic boundary; [11] the earliest insects that are certainly Caeliferans are in the extinct families Locustopseidae and Locustavidae from the early Triassic, roughly 250 million years ago. The group diversified during the Triassic and have remained important plant-eaters from that time to now. The first modern families such as the Eumastacidae, Tetrigidae and Tridactylidae appeared in the Cretaceous, though some insects that might belong to the last two of these groups are found in the early Jurassic. [12] [13] Morphological classification is difficult because many taxa have converged towards a common habitat type; recent taxonomists have concentrated on the internal genitalia, especially those of the male. This information is not available from fossil specimens, and the palaeontological taxonomy is founded principally on the venation of the hindwings. [14]

The Caelifera includes some 2,400 valid genera and about 11,000 known species. Many undescribed species probably exist, especially in tropical wet forests. The Caelifera have a predominantly tropical distribution with fewer species known from temperate zones, but most of the superfamilies have representatives worldwide. They are almost exclusively herbivorous and are probably the oldest living group of chewing herbivorous insects. [14]

The most diverse superfamily is the Acridoidea, with around 8,000 species. The two main families in this are the Acrididae (grasshoppers and locusts) with a worldwide distribution, and the Romaleidae (lubber grasshoppers), found chiefly in the New World. The Ommexechidae and Tristiridae are South American, and the Lentulidae, Lithidiidae and Pamphagidae are mainly African. The Pauliniids are nocturnal and can swim or skate on water, and the Lentulids are wingless. [12] Pneumoridae are native to Africa, particularly southern Africa, and are distinguished by the inflated abdomens of the males. [15]

Characteristics

Grasshoppers have the typical insect body plan of head, thorax, and abdomen. The head is held vertically at an angle to the body, with the mouth at the bottom. The head bears a large pair of compound eyes which give all-round vision, three simple eyes which can detect light and dark, and a pair of thread-like antennae that are sensitive to touch and smell. The downward-directed mouthparts are modified for chewing and there are two sensory palps in front of the jaws. [16]

The thorax and abdomen are segmented and have a rigid cuticle made up of overlapping plates composed of chitin. The three fused thoracic segments bear three pairs of legs and two pairs of wings. The forewings, known as tegmina, are narrow and leathery while the hindwings are large and membranous, the veins providing strength. The legs are terminated by claws for gripping. The hind leg is particularly powerful. The legs of these species are so powerful that they can jump quite a long distance. they also use this to flee from danger. [17] The femur is robust and has several ridges where different surfaces join and the inner ridges bear stridulatory pegs in some species. The posterior edge of the tibia bears a double row of spines and there are a pair of articulated spurs near its lower end. The interior of the thorax houses the muscles that control the wings and legs. [16]

Two differential grasshoppers, with visible spines along the tibia of the hind legs Two grasshoppers utah.jpg
Two differential grasshoppers, with visible spines along the tibia of the hind legs
Ensifera, like this great green bush-cricket Tettigonia viridissima, somewhat resemble grasshoppers but have over 20 segments in their antennae and different ovipositors. Heupferd fg01.jpg
Ensifera, like this great green bush-cricket Tettigonia viridissima , somewhat resemble grasshoppers but have over 20 segments in their antennae and different ovipositors.

The abdomen has eleven segments, the first of which is fused to the thorax and contains the tympanal organ and hearing system. Segments two to eight are ring-shaped and joined by flexible membranes. Segments nine to eleven are reduced in size; segment nine bears a pair of cerci and segments ten and eleven have the reproductive organs. Female grasshoppers are normally larger than males, with short ovipositors. [16] The name of the suborder "Caelifera" comes from the Latin and means chisel-bearing, referring to the shape of the ovipositor. [18]

The grasshopper's auditory organs are located on its abdomen, rather than on its head. These organs consist of a pair of membranes, each positioned on either side of the first abdominal segment and tucked under the wings. Known as tympanal organs, these simple eardrums vibrate in response to sound waves, enabling the grasshopper to hear the songs of other grasshoppers. [19]

Those species that make easily heard noises usually do so by rubbing a row of pegs on the hind legs against the edges of the forewings (stridulation). These sounds are produced mainly by males to attract females, though in some species the females also stridulate. [20]

Grasshoppers may be confused with crickets, but they differ in many aspects; these include the number of segments in their antennae and the structure of the ovipositor, as well as the location of the tympanal organ and the methods by which sound is produced. [21] Ensiferans have antennae that can be much longer than the body and have at least 20–24 segments, while caeliferans have fewer segments in their shorter, stouter antennae. [20]

Biology

Diet and digestion

Structure of mouthparts Grasshopper mouth anatomy.svg
Structure of mouthparts

Most grasshoppers are polyphagous, eating vegetation from multiple plant sources, [22] but some are omnivorous and also eat animal tissue and animal faeces. [23] In general their preference is for grasses, including many cereals grown as crops. [24] The digestive system is typical of insects, with Malpighian tubules discharging into the midgut. Carbohydrates are digested mainly in the crop, while proteins are digested in the ceca of the midgut. Saliva is abundant but largely free of enzymes, helping to move food and Malpighian secretions along the gut. Some grasshoppers possess cellulase, which by softening plant cell walls makes plant cell contents accessible to other digestive enzymes. [25] Grasshoppers can also be cannibalistic when swarming. [26] [27] [28]

A differential grasshopper eating the leaf of a climbing pea plant. Cellulase in its digestive tract allows it to digest cellulose in the cell walls of plants. Grasshopper eating leaf.jpg
A differential grasshopper eating the leaf of a climbing pea plant. Cellulase in its digestive tract allows it to digest cellulose in the cell walls of plants.

Sensory organs

Frontal view of Egyptian locust (Anacridium aegyptium) showing the compound eyes, tiny ocelli and numerous setae Anacridium aegyptium - 01.jpg
Frontal view of Egyptian locust ( Anacridium aegyptium ) showing the compound eyes, tiny ocelli and numerous setae

Grasshoppers have a typical insect nervous system, and have an extensive set of external sense organs. On the side of the head are a pair of large compound eyes which give a broad field of vision and can detect movement, shape, colour and distance. There are also three simple eyes (ocelli) on the forehead which can detect light intensity, a pair of antennae containing olfactory (smell) and touch receptors, and mouthparts containing gustatory (taste) receptors. [29] At the front end of the abdomen there is a pair of tympanal organs for sound reception. There are numerous fine hairs (setae) covering the whole body that act as mechanoreceptors (touch and wind sensors), and these are most dense on the antennae, the palps (part of the mouth), and on the cerci at the tip of the abdomen. [30] There are special receptors (campaniform sensillae) embedded in the cuticle of the legs that sense pressure and cuticle distortion. [31] There are internal "chordotonal" sense organs specialized to detect position and movement about the joints of the exoskeleton. The receptors convey information to the central nervous system through sensory neurons, and most of these have their cell bodies located in the periphery near the receptor site itself. [30]

Circulation and respiration

Like other insects, grasshoppers have an open circulatory system and their body cavities are filled with haemolymph. A heart-like structure in the upper part of the abdomen pumps the fluid to the head from where it percolates past the tissues and organs on its way back to the abdomen. This system circulates nutrients throughout the body and carries metabolic wastes to be excreted into the gut. Other functions of the haemolymph include wound healing, heat transfer and the provision of hydrostatic pressure, but the circulatory system is not involved in gaseous exchange. [32] Respiration is performed using tracheae, air-filled tubes, which open at the surfaces of the thorax and abdomen through pairs of valved spiracles. Larger insects may need to actively ventilate their bodies by opening some spiracles while others remain closed, using abdominal muscles to expand and contract the body and pump air through the system. [33]

Jumping

Grasshoppers jump by extending their large back legs and pushing against the substrate (the ground, a twig, a blade of grass or whatever else they are standing on); the reaction force propels them into the air. [34] A large grasshopper, such as a locust, can jump about a metre (20 body lengths) without using its wings; the acceleration peaks at about 20 g. [35]

They jump for several reasons; to escape from a predator, to launch themselves into flight, or simply to move from place to place. For the escape jump in particular there is strong selective pressure to maximize take-off velocity, since this determines the range. This means that the legs must thrust against the ground with both high force and a high velocity of movement. A fundamental property of muscle is that it cannot contract with high force and high velocity at the same time. Grasshoppers overcome this by using a catapult mechanism to amplify the mechanical power produced by their muscles. [36]

The jump is a three-stage process. [37] First, the grasshopper fully flexes the lower part of the leg (tibia) against the upper part (femur) by activating the flexor tibiae muscle (the back legs of the grasshopper in the top photograph are in this preparatory position). Second, there is a period of co-contraction in which force builds up in the large, pennate extensor tibiae muscle, but the tibia is kept flexed by the simultaneous contraction of the flexor tibiae muscle. The extensor muscle is much stronger than the flexor muscle, but the latter is aided by specialisations in the joint that give it a large effective mechanical advantage over the former when the tibia is fully flexed. [38] Co-contraction can last for up to half a second, and during this period the extensor muscle shortens and stores elastic strain energy by distorting stiff cuticular structures in the leg. [39] The extensor muscle contraction is quite slow (almost isometric), which allows it to develop high force (up to 14 N in the desert locust), but because it is slow only low power is needed. The third stage of the jump is the trigger relaxation of the flexor muscle, which releases the tibia from the flexed position. The subsequent rapid tibial extension is driven mainly by the relaxation of the elastic structures, rather than by further shortening of the extensor muscle. In this way the stiff cuticle acts like the elastic of a catapult, or the bow of a bow-and-arrow. Energy is put into the store at low power by slow but strong muscle contraction, and retrieved from the store at high power by rapid relaxation of the mechanical elastic structures. [40]

Stridulation

Gnome-mime-sound-openclipart.svg
Grasshopper stridulation
Several unidentified grasshoppers stridulating
Problems playing this file? See media help.

Male grasshoppers spend much of the day stridulating, singing more actively under optimal conditions and being more subdued when conditions are adverse; females also stridulate, but their efforts are insignificant when compared to the males. Late-stage male nymphs can sometimes be seen making stridulatory movements, although they lack the equipment to make sounds, demonstrating the importance of this behavioural trait. The songs are a means of communication; the male stridulation seems to express reproductive maturity, the desire for social cohesion and individual well-being. Social cohesion becomes necessary among grasshoppers because of their ability to jump or fly large distances, and the song can serve to limit dispersal and guide others to favourable habitat. The generalised song can vary in phraseology and intensity, and is modified in the presence of a rival male, and changes again to a courtship song when a female is nearby. [41] In male grasshoppers of the family Pneumoridae, the enlarged abdomen amplifies stridulation. [15]

Life cycle

Six stages (instars) of development, from newly hatched nymph to fully winged adult Grasshoppermetasnodgrass.svg
Six stages (instars) of development, from newly hatched nymph to fully winged adult
Romalea microptera grasshoppers: female (larger) is laying eggs, with male in attendance. Two eastern Lubber grasshopers (Romalea microptera), mating.jpg
Romalea microptera grasshoppers: female (larger) is laying eggs, with male in attendance.

In most grasshopper species, conflicts between males over females rarely escalate beyond ritualistic displays. Some exceptions include the chameleon grasshopper ( Kosciuscola tristis ), where males may fight on top of ovipositing females; engaging in leg grappling, biting, kicking and mounting. [42]

The newly emerged female grasshopper has a preoviposition period of a week or two while she increases in weight and her eggs mature. After mating, the female of most species digs a hole with her ovipositor and lays a batch of eggs in a pod in the ground near food plants, generally in the summer. After laying the eggs, she covers the hole with soil and litter. [16] Some, like the semi-aquatic Cornops aquaticum , deposit the pod directly into plant tissue. [43] The eggs in the pod are glued together with a froth in some species. After a few weeks of development, the eggs of most species in temperate climates go into diapause, and pass the winter in this state. Diapause is broken by a sufficiently low ground temperature, with development resuming as soon as the ground warms above a certain threshold temperature. The embryos in a pod generally all hatch out within a few minutes of each other. They soon shed their membranes and their exoskeletons harden. These first instar nymphs can then jump away from predators. [44]

Grasshoppers undergo incomplete metamorphosis: they repeatedly moult, each instar becoming larger and more like an adult, with the wing-buds increasing in size at each stage. The number of instars varies between species but is often six. After the final moult, the wings are inflated and become fully functional. The migratory grasshopper, Melanoplus sanguinipes, spends about 25 to 30 days as a nymph, depending on sex and temperature, and lives for about 51 days as an adult. [44]

Swarming

Millions of plague locusts on the move in Australia CSIRO ScienceImage 7007 Plague locusts on the move.jpg
Millions of plague locusts on the move in Australia

Locusts are the swarming phase of certain species of short-horned grasshoppers in the family Acrididae. Swarming behaviour is a response to overcrowding. Increased tactile stimulation of the hind legs causes an increase in levels of serotonin. [45] This causes the grasshopper to change colour, feed more and breed faster. The transformation of a solitary individual into a swarming one is induced by several contacts per minute over a short period. [46]

Following this transformation, under suitable conditions dense nomadic bands of flightless nymphs known as "hoppers" can occur, producing pheromones which attract the insects to each other. With several generations in a year, the locust population can build up from localised groups into vast accumulations of flying insects known as plagues, devouring all the vegetation they encounter. The largest recorded locust swarm was one formed by the now-extinct Rocky Mountain locust in 1875; the swarm was 1,800 miles (2,900 km) long and 110 miles (180 km) wide, [47] and one estimate puts the number of locusts involved at 3.5 trillion. [48] An adult desert locust can eat about 2 g (0.1 oz) of plant material each day, so the billions of insects in a large swarm can be very destructive, stripping all the foliage from plants in an affected area and consuming stems, flowers, fruits, seeds and bark. [49]

Predators, parasites, and pathogens

Cottontop tamarin monkey eating a grasshopper Cottontop tamarin.JPG
Cottontop tamarin monkey eating a grasshopper

Grasshoppers have a wide range of predators at different stages of their lives; eggs are eaten by bee-flies, ground beetles and blister beetles; hoppers and adults are taken by other insects such as ants and , robber flies and sphecid wasps, by spiders, and by many birds and small mammals including dogs and cats. [50]

The eggs and nymphs are under attack by parasitoids including blow flies, flesh flies, and tachinid flies. External parasites of adults and nymphs include mites. [50] Female grasshoppers parasitised by mites produce fewer eggs and thus have fewer offspring than unaffected individuals. [51]

Grasshopper with parasitic mites Unidentified Phlaeoba, near Godean Street, Yogyakarta, 2015-03-17.jpg
Grasshopper with parasitic mites

The grasshopper nematode (Mermis nigrescens) is a long slender worm that infects grasshoppers, living in the insects' hemocoel. Adult worms lay eggs on plants and the host becomes infected when the foliage is eaten. [52] Spinochordodes tellinii and Paragordius tricuspidatus are parasitic worms that infect grasshoppers and alter the behaviour of their hosts. When the worms are sufficiently developed, the grasshopper is persuaded to leap into a nearby body of water where it drowns, thus enabling the parasite to continue with the next stage of its life cycle, which takes place in water. [53] [54]

Locusts killed by the naturally occurring fungus, Metarhizium, an environmentally friendly means of biological control. CSIRO, 2005 CSIRO ScienceImage 1367 Locusts attacked by the fungus Metarhizium.jpg
Locusts killed by the naturally occurring fungus, Metarhizium , an environmentally friendly means of biological control. CSIRO, 2005

Grasshoppers are affected by diseases caused by bacteria, viruses, fungi and protozoa. The bacteria Serratia marcescens and Pseudomonas aeruginosa have both been implicated in causing disease in grasshoppers, as has the entomopathogenic fungus Beauveria bassiana . This widespread fungus has been used to control various pest insects around the world, but although it infects grasshoppers, the infection is not usually lethal because basking in the sun has the result of raising the insects' temperature above a threshold tolerated by the fungus. [56] The fungal pathogen Entomophaga grylli is able to influence the behaviour of its grasshopper host, causing it to climb to the top of a plant and cling to the stem as it dies. This ensures wide dispersal of the fungal spores liberated from the corpse. [57]

The fungal pathogen Metarhizium acridum is found in Africa, Australia and Brazil where it has caused epizootics in grasshoppers. It is being investigated for possible use as a microbial insecticide for locust control. [56] The microsporidian fungus Nosema locustae , once considered to be a protozoan, can be lethal to grasshoppers. It has to be consumed by mouth and is the basis for a bait-based commercial microbial pesticide. Various other microsporidians and protozoans are found in the gut. [56]

Anti-predator defences

Grasshoppers exemplify a range of anti-predator adaptations, enabling them to avoid detection, to escape if detected, and in some cases to avoid being eaten if captured. Grasshoppers are often camouflaged to avoid detection by predators that hunt by sight; some species can change their coloration to suit their surroundings. [58]

Several species such as the hooded leaf grasshopper Phyllochoreia ramakrishnai (Eumastacoidea) are detailed mimics of leaves. Stick grasshoppers (Proscopiidae) mimic wooden sticks in form and coloration. [59] Grasshoppers often have deimatic patterns on their wings, giving a sudden flash of bright colours that may startle predators long enough to give time to escape in a combination of jump and flight. [60]

Some species are genuinely aposematic, having both bright warning coloration and sufficient toxicity to dissuade predators. Dictyophorus productus (Pyrgomorphidae) is a "heavy, bloated, sluggish insect" that makes no attempt to hide; it has a bright red abdomen. A Cercopithecus monkey that ate other grasshoppers refused to eat the species. [61] Another species, the rainbow or painted grasshopper of Arizona, Dactylotum bicolor (Acridoidea), has been shown by experiment with a natural predator, the little striped whiptail lizard, to be aposematic. [62]

Relationship with humans

Detail of grasshopper on table in Rachel Ruysch's painting Flowers in a Vase, c. 1685. National Gallery, London Grasshopper detail in Rachel Ruysch Flowers in a Vase c 1685.jpg
Detail of grasshopper on table in Rachel Ruysch's painting Flowers in a Vase, c. 1685. National Gallery, London

In art and media

Grasshoppers are occasionally depicted in artworks, such as the Dutch Golden Age painter Balthasar van der Ast's still life oil painting, Flowers in a Vase with Shells and Insects, c. 1630, now in the National Gallery, London, though the insect may be a bush-cricket. [64]

Another orthopteran is found in Rachel Ruysch's still life Flowers in a Vase, c. 1685. The seemingly static scene is animated by a "grasshopper on the table that looks about ready to spring", according to the gallery curator Betsy Wieseman, with other invertebrates including a spider, an ant, and two caterpillars. [65] [66]

Grasshoppers are also featured in cinema. The 1957 film Beginning of the End portrayed giant grasshoppers attacking Chicago. [67] In the 1998 Disney/Pixar animated film A Bug's Life , the antagonists are a gang of grasshoppers, with their leader Hopper serving as the main villain. [68]

The protagonists of the 1971 tokusatsu series Kamen Rider primarily carry a grasshopper motif (for example Kamen Rider Black's Batta Man form), which continues to serve as the baseline visual template for most entries in the media franchise it has given birth to since.

Symbolism

Sir Thomas Gresham's gilded grasshopper symbol, Lombard Street, London, 1563 Londres - Lombard Street.JPG
Sir Thomas Gresham's gilded grasshopper symbol, Lombard Street, London, 1563

Grasshoppers are sometimes used as symbols. [69] During the Greek Archaic Era, the grasshopper was the symbol of the polis of Athens, [70] possibly because they were among the most common insects on the dry plains of Attica. [70] Native Athenians for a while wore golden grasshopper brooches to symbolise that they were of pure Athenian lineage with no foreign ancestors. [70] In addition, Peisistratus hung the figure of a kind of grasshopper before the Acropolis of Athens as apotropaic magic. [71]

Another symbolic use of the grasshopper is Sir Thomas Gresham's gilded grasshopper in Lombard Street, London, dating from 1563; [a] the building was for a while the headquarters of the Guardian Royal Exchange, but the company declined to use the symbol for fear of confusion with the locust. [72]

Grasshoppers appearing in dreams have been interpreted as symbols of "Freedom, independence, spiritual enlightenment, inability to settle down or commit to decision". Locusts are taken literally to mean devastation of crops in the case of farmers; figuratively as "wicked men and women" for non-farmers; and "Extravagance, misfortune, & ephemeral happiness" by "gypsies". [73]

As food

Fried grasshoppers from Gunung Kidul, Yogyakarta, Indonesia Fried grasshoper.jpg
Fried grasshoppers from Gunung Kidul, Yogyakarta, Indonesia
Sweet-and-salty grasshoppers dish in Japan (Inago no Tsukudani) Inago no tsukudani 02.jpg
Sweet-and-salty grasshoppers dish in Japan ( Inago no Tsukudani )

In some countries, grasshoppers are used as food. [74] In southern Mexico, grasshoppers, known as chapulines , are eaten in a variety of dishes, such as in tortillas with chilli sauce. [75] Grasshoppers are served on skewers in some Chinese food markets, like the Donghuamen Night Market. [76] Fried grasshoppers (walang goreng) are eaten in the Gunung Kidul Regency, Yogyakarta, Java in Indonesia. [77] Grasshoppers are a beloved delicacy in Uganda; they are usually eaten fried (most commonly in November and May after the rains). [78] In America, the Ohlone burned grassland to herd grasshoppers into pits where they could be collected as food. [79]

It is recorded in the Bible that John the Baptist ate locusts and wild honey (Greek: ἀκρίδες καὶ μέλι ἄγριον, akrídes kaì méli ágrion) while living in the wilderness. [80] However, because of a tradition of depicting him as an ascetic, attempts have been made to explain that the locusts were in fact a suitably ascetic vegetarian food such as carob beans, notwithstanding the fact that the word ἀκρίδες means plainly grasshoppers. [81] [82]

In recent years, with the search for alternative healthy and sustainable protein sources, grasshoppers are being cultivated by commercial companies operating grasshopper farms and are being used as food and protein supplements.

As pests

Crop pest: grasshopper eating a maize leaf Grasshopper eating the maize leaf.jpg
Crop pest: grasshopper eating a maize leaf

Grasshoppers eat large quantities of foliage both as adults and during their development, and can be serious pests of arid land and prairies. Pasture, grain, forage, vegetable and other crops can be affected. Grasshoppers often bask in the sun, and thrive in warm sunny conditions, so drought stimulates an increase in grasshopper populations. A single season of drought is not normally sufficient to stimulate a major population increase, but several successive dry seasons can do so, especially if the intervening winters are mild so that large numbers of nymphs survive. Although sunny weather stimulates growth, there needs to be an adequate food supply for the increasing grasshopper population. This means that although precipitation is needed to stimulate plant growth, prolonged periods of cloudy weather will slow nymphal development. [83]

Grasshoppers can best be prevented from becoming pests by manipulating their environment. Shade provided by trees will discourage them and they may be prevented from moving onto developing crops by removing coarse vegetation from fallow land and field margins and discouraging thick growth beside ditches and on roadside verges. With increasing numbers of grasshoppers, predator numbers may increase, but this seldom happens rapidly enough to have much effect on populations. Biological control is being investigated, and spores of the protozoan parasite Nosema locustae can be used mixed with bait to control grasshoppers, being more effective with immature insects. [84] On a small scale, neem products can be effective as a feeding deterrent and as a disruptor of nymphal development. Insecticides can be used, but adult grasshoppers are difficult to kill, and as they move into fields from surrounding rank growth, crops may soon become reinfested. [83]

Some grasshopper species, like the Chinese rice grasshopper, are a pest in rice paddies. Ploughing exposes the eggs on the surface of the field, to be destroyed by sunshine or eaten by natural enemies. Some eggs may be buried too deeply in the soil for hatching to take place. [85]

Locust plagues can have devastating effects on human populations, causing famines and population upheavals. They are mentioned in both the Qur’an and the Bible and have also been held responsible for cholera epidemics, resulting from the corpses of locusts drowned in the Mediterranean Sea and decomposing on beaches. [49] The FAO and other organisations monitor locust activity around the world. Timely application of pesticides can prevent nomadic bands of hoppers from forming before dense swarms of adults can build up. [86] Besides conventional control using contact insecticides, [86] biological pest control using the entomopathogenic fungus Metarhizium acridum , which specifically infects grasshoppers, has been used with some success. [87]

Detection of explosives

In February 2020, researchers from Washington University in St. Louis announced they had engineered "cyborg grasshoppers" capable of accurately detecting explosives. In the project, funded by the US Office of Naval Research, researchers fitted grasshoppers with lightweight sensor backpacks that recorded and transmitted the electrical activity of their antennal lobes to a computer. According to the researchers, the grasshoppers were able to detect the location of the highest concentration of explosives. The researchers also tested the effect of combining sensorial information from several grasshoppers on detection accuracy. The neural activity from seven grasshoppers yielded an average detection accuracy rate of 80%, whereas a single grasshopper yielded a 60% rate. [88] [89]

In literature

Egyptian hieroglyphs "snhm" Egyptian-snhm 2.PNG
Egyptian hieroglyphs "snḥm"

The Egyptian word for locust or grasshopper was written snḥm in the consonantal hieroglyphic writing system. The pharaoh Ramesses II compared the armies of the Hittites to locusts: "They covered the mountains and valleys and were like locusts in their multitude." [90]

One of Aesop's Fables, later retold by La Fontaine, is the tale of The Ant and the Grasshopper . The ant works hard all summer, while the grasshopper plays. In winter, the ant is ready but the grasshopper starves. Somerset Maugham's short story "The Ant and the Grasshopper" explores the fable's symbolism via complex framing. [91] Other human weaknesses besides improvidence have become identified with the grasshopper's behaviour. [73] So an unfaithful woman (hopping from man to man) is "a grasshopper" in "Poprygunya", an 1892 short story by Anton Chekhov, [92] and in Jerry Paris's 1969 film The Grasshopper . [93] [94]

In mechanical engineering

A grasshopper beam engine, 1847 DMM 12236 Halbbalancier-Dampfmaschine.jpg
A grasshopper beam engine, 1847

The name "Grasshopper" was given to the Aeronca L-3 and Piper L-4 light aircraft, both used for reconnaissance and other support duties in World War II. The name is said to have originated when Major General Innis P. Swift saw a Piper making a rough landing and remarked that it looked like a grasshopper for its bouncing progress. [94] [95] [96]

Grasshopper beam engines were beam engines pivoted at one end, the long horizontal arm resembling the hind leg of a grasshopper. The type was patented by William Freemantle in 1803. [97] [98] [99]

Notes

  1. ↑ The symbol is a wordplay on the name Gresham and "grass". [69]

Related Research Articles

<span class="mw-page-title-main">Tettigoniidae</span> Family of insects

Insects in the family Tettigoniidae are commonly called katydids or bush crickets. They have previously been known as "long-horned grasshoppers". More than 8,000 species are known. Part of the suborder Ensifera, the Tettigoniidae are the only extant (living) family in the superfamily Tettigonioidea.

<span class="mw-page-title-main">Orthoptera</span> Order of insects including grasshoppers, crickets, weta and locusts

Orthoptera is an order of insects that comprises the grasshoppers, locusts, and crickets, including closely related insects, such as the bush crickets or katydids and wētā. The order is subdivided into two suborders: Caelifera – grasshoppers, locusts, and close relatives; and Ensifera – crickets and close relatives.

<span class="mw-page-title-main">Locust</span> Grasshopper that has a swarming phase

Locusts are various 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; this has evolved independently in multiple lineages, comprising at least 18 genera in 5 different subfamilies.

<span class="mw-page-title-main">Mayfly</span> Aquatic insects of the order Ephemeroptera

Mayflies are aquatic insects belonging to the order Ephemeroptera. This order is part of an ancient group of insects termed the Palaeoptera, which also contains dragonflies and damselflies. Over 3,000 species of mayfly are known worldwide, grouped into over 400 genera in 42 families.

<span class="mw-page-title-main">Anostostomatidae</span> Family of cricket-like animals

Anostostomatidae is a family of insects in the order Orthoptera, widely distributed in the southern hemisphere. It is named Mimnermidae or Henicidae in some taxonomies, and common names include king crickets in Australia and South Africa, and wētā in New Zealand. Prominent members include the Parktown prawn of South Africa, and the giant wētā of New Zealand.

<span class="mw-page-title-main">Mole cricket</span> Members of the insect family Gryllotalpidae

Mole crickets are members of the insect family Gryllotalpidae, in the order Orthoptera. Mole crickets are cylindrical-bodied, fossorial insects about 3–5 cm (1.2–2.0 in) long as adults, with small eyes and shovel-like fore limbs highly developed for burrowing. They are present in many parts of the world and where they have arrived in new regions, may become agricultural pests.

<span class="mw-page-title-main">Desert locust</span> Species of grasshopper

The desert locust is a species of locust, a periodically swarming, short-horned grasshopper in the family Acrididae. They are found primarily in the deserts and dry areas of northern and eastern Africa, Arabia, and southwest Asia. During population surge years, they may extend north into parts of Southern Europe, south into Eastern Africa, and east in northern India. The desert locust shows periodic changes in its body form and can change in response to environmental conditions, over several generations, from a solitary, shorter-winged, highly fecund, non-migratory form to a gregarious, long-winged, and migratory phase in which they may travel long distances into new areas. In some years, they may thus form locust plagues, invading new areas, where they may consume all vegetation including crops, and at other times, they may live unnoticed in small numbers.

<span class="mw-page-title-main">Caelifera</span> Suborder of insects

The Caelifera are a suborder of orthopteran insects. They include the grasshoppers and grasshopper-like insects, as well as other superfamilies classified with them: the ground-hoppers (Tetrigoidea) and pygmy mole crickets (Tridactyloidea). The latter should not be confused with the mole crickets (Gryllotalpidae), which belong to the other Orthopteran sub-order Ensifera.

<span class="mw-page-title-main">Roesel's bush-cricket</span> Species of cricket-like animal

Roesel's bush-cricket, Roeseliana roeselii is a European bush-cricket, named after August Johann Rösel von Rosenhof, a German entomologist.

<i>Eurycnema goliath</i> Species of stick insect

Eurycnema goliath, commonly known as the goliath stick insect, or the regal stick insect, is a large species of stick insect in the family Phasmatidae, endemic to Australia and considered one of the largest species of stick insects in the country. The species has the Phasmid Study Group number PSG14.

<span class="mw-page-title-main">Cricket (insect)</span> Small insects of the family Gryllidae

Crickets are orthopteran insects which are related to bush crickets, and, more distantly, to grasshoppers. In older literature, such as Imms, "crickets" were placed at the family level, but contemporary authorities including Otte now place them in the superfamily Grylloidea. The word has been used in combination to describe more distantly related taxa in the suborder Ensifera, such as king crickets and mole crickets.

<span class="mw-page-title-main">Insect</span> Class of arthropods

Insects are hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body, three pairs of jointed legs, compound eyes, and a pair of antennae. Insects are the most diverse group of animals, with more than a million described species; they represent more than half of all animal species.

<span class="mw-page-title-main">Senegalese grasshopper</span> Species of grasshopper

The Senegalese grasshopper is a medium-sized grasshopper species found in the Sahel region of Africa, the Canary Islands, Cape Verde Islands, and West Asia. Although not called a locust in English, this species shows gregarious behaviour and some morphological change on crowding. In many parts of the Sahel, this species may cause greater year-on-year crop damage than better-known locusts, attacking crops such as the pearl millet.

<span class="mw-page-title-main">Insect morphology</span> Description of the physical form of insects

Insect morphology is the study and description of the physical form of insects. The terminology used to describe insects is similar to that used for other arthropods due to their shared evolutionary history. Three physical features separate insects from other arthropods: they have a body divided into three regions, three pairs of legs, and mouthparts located outside of the head capsule. This position of the mouthparts divides them from their closest relatives, the non-insect hexapods, which include Protura, Diplura, and Collembola.

<i>Taeniopoda eques</i> Species of grasshopper

Taeniopoda eques, the western horse lubber grasshopper, is a relatively large grasshopper species of the family Romaleidae found in arid and semi-arid parts of southwestern United States to central and southwestern Mexico. Most populations are identifiable by their shiny black bodies with contrasting yellow markings, but some adults are mostly yellowish, orangish or greenish. The species is unique in using its black coloration to thermoregulate and in being chemically defended. The aposematic coloration warns vertebrate predators of its unpalatability and allows the grasshopper to roost conspicuously upon shrubs.

<i>Tetrix ceperoi</i> Species of grasshopper

Tetrix ceperoi, Cepero's groundhopper, is a member of the family Tetrigidae and is very similar to common grasshoppers. Grasshopper is defined as a "plant eating insect with long hind legs that are used for jumping and for producing a chirping sound". However, unlike the common grasshopper, the wings of T. ceperoi extend beyond its pronotum. The front wings have evolved throughout history to be stumps, and the back wings are very well developed. Thanks to the front wings only, T. ceperoi is capable of flying. Furthermore, T. ceperoi sports wide shoulders while covering its narrow abdomen beneath the pronotum. T. ceperoi are classified as Orthoptera, which describes crickets, grasshoppers, and locusts. Furthermore, these Orthoptera have incomplete metamorphosis, which also affects sexual dimorphism later. T. ceperoi reach an average length of about 10 millimetres (0.39 in). T. ceperoi is a multi-coloured ground dweller with the ability to blend into its surroundings. The ability of T. ceperoi to be different colours makes it able to evolve colour schemes better adapted to specific habitats. This cryptic nature of their outer layer provides protection from predators as it is able to blend into its surroundings. T. ceperoi is diurnal, which means it is solely active in the daytime. However, although it is diurnal, it continues to hibernate during a late nymphal instar or later on in life as adults.

<i>Conocephalus fuscus</i> Species of cricket-like animal

Conocephalus fuscus, the long-winged conehead, is a member of the family Tettigoniidae, the bush-crickets and is distributed through much of Europe and temperate Asia. This bush-cricket is native to the British Isles where it may confused with the short-winged conehead. These two species are phenotypically similar; however, the distinguishing factor between the two is the fully developed set of wings the long-winged conehead possesses that allows for flight. In the short-winged coneheads the hind wings are shorter than the abdomen, causing the wings to be vestigial and the species is incapable of flight. For this reason it is hard to discriminate between the two species during the early stages of their life cycle before the wings have fully developed. The colouration of the conehead is typically a grass green with a distinctive brown stripe down its back, though there are some brown phenotypes.

<span class="mw-page-title-main">Tridactylidae</span> Family of Caelifera

The Tridactylidae are a family in the insect order Orthoptera. They are small, mole-cricket-like insects, almost always less than 20 mm (0.79 in) long when mature. Generally they are shiny, dark or black, sometimes variegated or sandy-coloured. They commonly live in short tunnels and are commonly known as pygmy mole crickets, though they are not closely related to the true "mole crickets" (Ensifera), as they are included in the Caelifera suborder.

<i>Aularches</i> Genus of grasshopper

Aularches miliaris is a grasshopper species of the monotypic genus Aularches, belonging to the family Pyrgomorphidae. A native of South and Southeast Asia, the bright warning colours of this fairly large grasshopper keep away predators and their defense when disturbed includes the ejection of a toxic foam.

<i>Schistocerca americana</i> Species of grasshopper

Schistocerca americana is a species of grasshopper in the family Acrididae known commonly as the American grasshopper and American bird grasshopper. It is native to North America, where it occurs in the eastern United States, Mexico, and the Bahamas. Occasional, localized outbreaks of this grasshopper occur, and it is often referred to as a locust, though it lacks the true swarming form of its congener, the desert locust.

References

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