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Grasshoppers are a group of
insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pa ...
s belonging to the
suborder Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...
Caelifera. They are among what is possibly the most ancient living group of chewing herbivorous insects, dating back to the
early Triassic The Early Triassic is the first of three epochs of the Triassic Period of the geologic timescale. It spans the time between Ma and Ma (million years ago). Rocks from this epoch are collectively known as the Lower Triassic Series, which is ...
around 250 million years ago. Grasshoppers are typically ground-dwelling insects with powerful hind legs which allow them to escape from threats by leaping vigorously. As hemimetabolous insects, they do not undergo
complete metamorphosis Holometabolism, also called complete metamorphosis, is a form of insect development which includes four life stages: egg, larva, pupa, and imago (or adult). Holometabolism is a synapomorphic trait of all insects in the superorder Endopterygota. ...
; they hatch from an egg into a
nymph A nymph ( grc, νύμφη, nýmphē, el, script=Latn, nímfi, label=Modern Greek; , ) in ancient Greek folklore is a minor female nature deity. Different from Greek goddesses, nymphs are generally regarded as personifications of nature, are ...
or "hopper" which undergoes five moults, becoming more similar to the adult insect at each developmental stage. 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, the 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 color 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
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill t ...
s by
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
; 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 Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defences which make the prey difficult to kill and eat, such as toxicity, venom, foul taste ...
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
parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...
s and predators. Grasshoppers have had a long relationship with humans. Swarms of locusts can have devastating effects and cause famine, having done so since
Biblical times The history of ancient Israel and Judah begins in the Southern Levant during the Late Bronze Age and Early Iron Age. "Israel" as a people or tribal confederation (see Israelites) appears for the first time in the Merneptah Stele, an inscripti ...
. 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" is sometimes used as a common name for the suborder in general, some sources restrict it to the more "advanced" groups. They may be placed in the infraorder Acrididea and have been referred-to as "short-horned grasshoppers" in older textsImms A.D., rev. Richards O.W. & Davies R.G. (1970) ''A General Textbook of Entomology'' 9th Ed. Methuen 886 pp. to distinguish them from the also-obsolete term "long-horned grasshoppers" (now
bush-cricket Insects in the family Tettigoniidae are commonly called katydids (especially in North America), or bush crickets. They have previously been known as "long-horned grasshoppers". More than 8,000 species are known. Part of the suborder Ensifera, ...
s or katydids) with their much longer antennae. The
phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological s ...
of the Caelifera, based on mitochondrial
ribosomal RNA Ribosomal ribonucleic acid (rRNA) is a type of non-coding RNA which is the primary component of ribosomes, essential to all cells. rRNA is a ribozyme which carries out protein synthesis in ribosomes. Ribosomal RNA is transcribed from riboso ...
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. In evolutionary terms, the split between the Caelifera and the Ensifera is no more recent than the Permo-Triassic boundary; 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 The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
, though some insects that might belong to the last two of these groups are found in the early Jurassic. 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 palaentological taxonomy is founded principally on the venation of the hindwings. 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. 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 The Romaleidae or lubber grasshoppers are a family of grasshoppers, based on the type genus ''Romalea''. The species in this family can be found in the Americas. Tribes and selected genera The Orthoptera Species File Online database lists two ...
(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. Pneumoridae are native to Africa, particularly southern Africa, and are distinguished by the inflated abdomens of the males.


Characteristics

Grasshoppers have the typical insect body plan of head,
thorax The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the c ...
and
abdomen The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the to ...
. 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 Jaws or Jaw may refer to: Anatomy * Jaw, an opposable articulated structure at the entrance of the mouth ** Mandible, the lower jaw Arts, entertainment, and media * Jaws (James Bond), a character in ''The Spy Who Loved Me'' and ''Moonraker'' * ...
.Pfadt, 1994. pp. 1–8 The thorax and abdomen are segmented and have a rigid cuticle made up of overlapping plates composed of
chitin Chitin ( C8 H13 O5 N)n ( ) is a long-chain polymer of ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is probably the second most abundant polysaccharide in nature (behind only cellulose); an estimated 1 billion tons of chit ...
. 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 femur is robust and has several ridges where different surfaces join and the inner ridges bear
stridulatory Stridulation is the act of producing sound by rubbing together certain body parts. This behavior is mostly associated with insects, but other animals are known to do this as well, such as a number of species of fish, snakes and spiders. The mech ...
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. The abdomen has eleven segments, the first of which is fused to the thorax and contains the
tympanal organ A tympanal organ (or tympanic organ) is a hearing organ in insects, consisting of a membrane ( tympanum) stretched across a frame backed by an air sac and associated sensory neurons. Sounds vibrate the membrane, and the vibrations are sensed by ...
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. The name of the suborder "Caelifera" comes from the Latin and means ''chisel-bearing'', referring to the shape of the ovipositor. 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. 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. 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.


Biology


Diet and digestion

Most grasshoppers are
polyphagous Feeding is the process by which organisms, typically animals, obtain food. Terminology often uses either the suffixes -vore, -vory, or -vorous from Latin ''vorare'', meaning "to devour", or -phage, -phagy, or -phagous from Greek φαγε ...
, eating vegetation from multiple plant sources, but some are omnivorous and also eat animal tissue and animal faeces. In general their preference is for grasses, including many
cereal A cereal is any grass cultivated for the edible components of its grain (botanically, a type of fruit called a caryopsis), composed of the endosperm, germ, and bran. Cereal grain crops are grown in greater quantities and provide more food ...
s grown as crops. 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 Cellulase (EC 3.2.1.4; systematic name 4-β-D-glucan 4-glucanohydrolase) is any of several enzymes produced chiefly by fungi, bacteria, and protozoans that catalyze cellulolysis, the decomposition of cellulose and of some related polysaccha ...
, which by softening plant cell walls makes plant cell contents accessible to other digestive enzymes.


Sensory organs

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. At the front end of the abdomen there is a pair of tympanal organs for sound reception. There are numerous fine hairs (
setae In biology, setae (singular seta ; from the Latin word for "bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. ...
) 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.Chapman, 2013. pp. 745–755 There are special receptors ( campaniform sensillae) embedded in the cuticle of the legs that sense pressure and cuticle distortion. 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.


Circulation and respiration

Like other insects, grasshoppers have an
open circulatory system The blood circulatory system is a system of organs that includes the heart, blood vessels, and blood which is circulated throughout the entire body of a human or other vertebrate. It includes the cardiovascular system, or vascular system, t ...
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 waste Metabolic wastes or excrements are substances left over from metabolic processes (such as cellular respiration) which cannot be used by the organism (they are surplus or toxic), and must therefore be excreted. This includes nitrogen compounds, ...
s 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. Respiration is performed using
trachea The trachea, also known as the windpipe, is a cartilaginous tube that connects the larynx to the bronchi of the lungs, allowing the passage of air, and so is present in almost all air- breathing animals with lungs. The trachea extends from t ...
e, 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.


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. A large grasshopper, such as a locust, can jump about a metre (twenty body lengths) without using its wings; the acceleration peaks at about 20 g. 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 Skeletal muscles (commonly referred to as muscles) are organs of the vertebrate muscular system and typically are attached by tendons to bones of a skeleton. The muscle cells of skeletal muscles are much longer than in the other types of mus ...
is that it cannot contract with high force and high velocity at the same time. Grasshoppers overcome this by using a
catapult A catapult is a ballistic device used to launch a projectile a great distance without the aid of gunpowder or other propellants – particularly various types of ancient and medieval siege engines. A catapult uses the sudden release of stor ...
mechanism to amplify the mechanical power produced by their muscles. The jump is a three-stage process. 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. 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. 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 A catapult is a ballistic device used to launch a projectile a great distance without the aid of gunpowder or other propellants – particularly various types of ancient and medieval siege engines. A catapult uses the sudden release of stor ...
, 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.


Stridulation

Male grasshoppers spend much of the day
stridulating Stridulation is the act of producing sound by rubbing together certain body parts. This behavior is mostly associated with insects, but other animals are known to do this as well, such as a number of species of fish, snakes and spiders. The mech ...
, 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. In male grasshoppers of the family Pneumoridae, the enlarged abdomen amplifies stridulation.


Life cycle

In most grasshopper species, conflicts between males over females rarely escalate beyond ritualistic displays. Some exceptions include the chameleon grasshopper (''
Kosciuscola tristis ''Kosciuscola tristis'', known generally as the chameleon skyhopper, is a species of short-horned grasshopper in the family Acrididae The AcrididaeMacLeay WS (1821) ''Horae Entomologicae or Essays on the Annulose Animals'' 2 are the predomina ...
''), where males may fight on top of ovipositing females; engaging in leg grappling, biting, kicking and mounting. 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. Some, like the semi-aquatic ''
Cornops aquaticum ''Cornops aquaticum'' is a semiaquatic species of grasshopper native to the Neotropics, from southern Mexico south to central Argentina and Uruguay. It feeds and breeds exclusively on members of the aquatic plant family Pontederiaceae, especially ...
'', deposit the pod directly into plant tissue. 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 In animal dormancy, diapause is the delay in development in response to regular and recurring periods of adverse environmental conditions.Tauber, M.J., Tauber, C.A., Masaki, S. (1986) ''Seasonal Adaptations of Insects''. Oxford University Press I ...
, 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 An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ...
nymphs can then jump away from predators. Pfadt, 1994. pp. 11–16
Diagrams
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.


Swarming

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 Serotonin () or 5-hydroxytryptamine (5-HT) is a monoamine neurotransmitter. Its biological function is complex and multifaceted, modulating mood, cognition, reward, learning, memory, and numerous physiological processes such as vomiting and va ...
. 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. 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 The Rocky Mountain locust (''Melanoplus spretus'') is an extinct species of grasshopper that ranged through the western half of the United States and some western portions of Canada with large numbers seen until the end of the 19th century. Si ...
in 1875; the swarm was long and wide, and one estimate puts the number of locusts involved at 3.5 trillion.ffol An adult
desert locust The desert locust (''Schistocerca gregaria'') 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 sou ...
can eat about 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.


Predators, parasites, and pathogens

Grasshoppers have a wide range of
predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill t ...
s at different stages of their lives; eggs are eaten by
bee-flies The Bombyliidae are a family of flies, commonly known as bee flies. Adults generally feed on nectar and pollen, some being important pollinators. Larvae are mostly parasitoids of other insects. Overview The Bombyliidae are a large family of fl ...
,
ground beetle Ground beetles are a large, cosmopolitan family of beetles, the Carabidae, with more than 40,000 species worldwide, around 2,000 of which are found in North America and 2,700 in Europe. As of 2015, it is one of the 10 most species-rich animal f ...
s and
blister beetle Blister beetles are beetles of the family Meloidae, so called for their defensive secretion of a blistering agent, cantharidin. About 7,500 species are known worldwide. Many are conspicuous and some are aposematically colored, announcing their ...
s; hoppers and adults are taken by other insects such as ants,
robber flies The Asilidae are the robber fly family, also called assassin flies. They are powerfully built, bristly flies with a short, stout proboscis enclosing the sharp, sucking hypopharynx. The name "robber flies" reflects their notoriously aggressive pr ...
and
sphecid wasp The Sphecidae are a cosmopolitan family of wasps of the suborder Apocrita that includes sand wasps, mud daubers, and other thread-waisted wasps. The name Sphecidae was formerly given to a much larger grouping of wasps. This was found to be ...
s, by
spider Spiders (order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...
s, and by many birds and small mammals including dogs and cats.Capinera, 2008. pp. 1709–1710 The eggs and nymphs are under attack by
parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...
s including
blow flies The Calliphoridae (commonly known as blow flies, blow-flies, carrion flies, bluebottles, greenbottles, or cluster flies) are a family of insects in the order Diptera, with almost 1,900 known species. The maggot larvae, often used as fishing ba ...
, flesh flies, and tachinid flies. External parasites of adults and nymphs include mites. Female grasshoppers parasitised by mites produce fewer eggs and thus have fewer offspring than unaffected individuals. The grasshopper nematode (''Mermis nigrescens'') is a long slender worm that infects grasshoppers, living in the insect's
hemocoel The blood circulatory system is a system of organs that includes the heart, blood vessels, and blood which is circulated throughout the entire body of a human or other vertebrate. It includes the cardiovascular system, or vascular system, t ...
. Adult worms lay eggs on plants and the host becomes infected when the foliage is eaten. ''
Spinochordodes tellinii ''Spinochordodes tellinii'' is a parasitic nematomorph hairworm whose larvae develop in grasshoppers and crickets. This parasite is able to influence its host's behavior: once the parasite is grown, it causes its grasshopper host to jump in ...
'' 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. Grasshoppers are affected by diseases caused by
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were am ...
,
virus A virus is a submicroscopic infectious agent that replicates only inside the living cells of an organism. Viruses infect all life forms, from animals and plants to microorganisms, including bacteria and archaea. Since Dmitri Ivanovsk ...
es,
fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately fr ...
and
protozoa Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Histo ...
. The bacteria ''
Serratia marcescens ''Serratia marcescens'' () is a species of rod-shaped, Gram-negative bacteria in the family Yersiniaceae. It is a facultative anaerobe and an opportunistic pathogen in humans. It was discovered in 1819 by Bartolomeo Bizio in Padua, Italy.Serr ...
'' and ''
Pseudomonas aeruginosa ''Pseudomonas aeruginosa'' is a common encapsulated, gram-negative, aerobic– facultatively anaerobic, rod-shaped bacterium that can cause disease in plants and animals, including humans. A species of considerable medical importance, ''P. a ...
'' 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 insect's temperature above a threshold tolerated by the fungus.Capinera, 2008. pp. 1229–1230 The fungal
pathogen In biology, a pathogen ( el, πάθος, "suffering", "passion" and , "producer of") in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a g ...
''
Entomophaga grylli ''Entomophaga grylli'' is a fungal pathogen which infects and kills grasshoppers. It is the causal agent of one of the most widespread diseases affecting grasshoppers. This is sometimes known as summit disease because infected insects climb to ...
'' 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. The fungal pathogen ''
Metarhizium acridum ''Metarhizium acridum'' is the new name given to a group of fungal isolates that are known to be virulent and specific to the Acrididea (grasshoppers). Previously, this species has had variety status in ''Metarhizium anisopliae'' (var. ''acrid ...
'' 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. The
microsporidia Microsporidia are a group of spore-forming unicellular parasites. These spores contain an extrusion apparatus that has a coiled polar tube ending in an anchoring disc at the apical part of the spore. They were once considered protozoans or pr ...
n 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.


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
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
d to avoid detection by predators that hunt by sight; some species can change their coloration to suit their surroundings. Several species such as the hooded leaf grasshopper ''
Phyllochoreia ramakrishnai ''Phyllochoreia'' is a genus of grasshopper endemic to the Western Ghats of India and Sri Lanka. It is in the family Chorotypidae of the superfamily Eumastacoidea. The genus comprises four species: * ''Phyllochoreia equa'' Burr, 1899 * '' Phyl ...
'' (Eumastacoidea) are detailed mimics of leaves. Stick grasshoppers (Proscopiidae) mimic wooden sticks in form and coloration. Grasshoppers often have
deimatic Deimatic behaviour or startle display means any pattern of bluffing behaviour in an animal that lacks strong defences, such as suddenly displaying conspicuous eyespots, to scare off or momentarily distract a predator, thus giving the prey anima ...
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.Cott, p. 378 Some species are genuinely aposematic, having both bright warning coloration and sufficient toxicity to dissuade predators. ''
Dictyophorus productus ''Dictyophorus griseus'' is a species of African grasshopper in the family Pyrgomorphidae Pyrgomorphidae is a family of grasshoppers in the order Orthoptera; it is the only family in the superfamily Pyrgomorphoidea, with a pan-tropical distrib ...
'' (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. Another species, the rainbow or painted grasshopper of Arizona, ''
Dactylotum bicolor ''Dactylotum bicolor'', also known as the rainbow grasshopper, painted grasshopper, or the barber pole grasshopper, is a species of grasshopper in the family Acrididae. It is native to the United States, Canada and northern Mexico and exhibits ...
'' (Acridoidea), has been shown by experiment with a natural predator, the
little striped whiptail The little striped whiptail (''Aspidoscelis inornatus'') is a species of lizard found in the southwestern United States (in Arizona, New Mexico and Texas) and in northern Mexico (in Chihuahua, Coahuila, Durango, Zacatecas, San Luis Potosí, and ...
lizard, to be aposematic. Atractomorpha lata, Burdwan, West Bengal, India 27 10 2012.jpg, Gaudy grasshopper, ''
Atractomorpha lata ''Atractomorpha'' may refer to: * ''Atractomorpha'' (alga), an algae genus in the family Sphaeropleaceae * ''Atractomorpha'' (grasshopper), a grasshopper genus in the family Pyrgomorphidae {{genus disambiguation ...
'', evades predators with
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
. Titanacris Albipes Vol.jpg, Lubber grasshopper, ''
Titanacris albipes Titanacris albipes is a species of grasshopper in the family Romaleidae The Romaleidae or lubber grasshoppers are a family of grasshoppers, based on the type genus '' Romalea''. The species in this family can be found in the Americas ...
'', has
deimatic Deimatic behaviour or startle display means any pattern of bluffing behaviour in an animal that lacks strong defences, such as suddenly displaying conspicuous eyespots, to scare off or momentarily distract a predator, thus giving the prey anima ...
ally coloured wings, used to startle predators. LeafGrasshopper.jpg, Leaf grasshopper, ''
Phyllochoreia ramakrishnai ''Phyllochoreia'' is a genus of grasshopper endemic to the Western Ghats of India and Sri Lanka. It is in the family Chorotypidae of the superfamily Eumastacoidea. The genus comprises four species: * ''Phyllochoreia equa'' Burr, 1899 * '' Phyl ...
'', mimics a green leaf. Dactylotum bicolor.jpg, Painted grasshopper, ''
Dactylotum bicolor ''Dactylotum bicolor'', also known as the rainbow grasshopper, painted grasshopper, or the barber pole grasshopper, is a species of grasshopper in the family Acrididae. It is native to the United States, Canada and northern Mexico and exhibits ...
'', deters predators with
warning coloration Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defences which make the prey difficult to kill and eat, such as toxicity, venom, foul taste ...
. Aularches miliaris at Mangunan Orchard, Dlingo, Bantul, Yogyakarta 07.jpg, Spotted grasshopper, '' Aularches miliaris'', defends itself with toxic foam and warning colours.


Relationship with humans


In art and media

Grasshoppers are occasionally depicted in artworks, such as the
Dutch Golden Age The Dutch Golden Age ( nl, Gouden Eeuw ) was a period in the history of the Netherlands, roughly spanning the era from 1588 (the birth of the Dutch Republic) to 1672 (the Rampjaar, "Disaster Year"), in which Dutch trade, science, and art and ...
painter
Balthasar van der Ast Balthasar van der Ast (1593/94 – 7 March 1657) was a Dutch Golden Age painter who specialized in still lifes of flowers and fruit, as well as painting a number of remarkable shell still lifes; he is considered to be a pioneer in the genr ...
's
still life A still life (plural: still lifes) is a work of art depicting mostly inanimate subject matter, typically commonplace objects which are either natural (food, flowers, dead animals, plants, rocks, shells, etc.) or man-made (drinking glasses, bo ...
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. 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. Grasshoppers are also featured in cinema. The 1957 film '' Beginning of the End'' portrayed giant grasshoppers attacking
Chicago (''City in a Garden''); I Will , image_map = , map_caption = Interactive Map of Chicago , coordinates = , coordinates_footnotes = , subdivision_type = List of sovereign states, Count ...
. In the 1998
Disney The Walt Disney Company, commonly known as Disney (), is an American multinational mass media and entertainment conglomerate headquartered at the Walt Disney Studios complex in Burbank, California. Disney was originally founded on October ...
/ Pixar animated film '' A Bug's Life'', the antagonists are a gang of grasshoppers, with their leader Hopper serving as the main villain.


Symbolism

Grasshoppers are sometimes used as symbols. During the Greek Archaic Era, the grasshopper was the symbol of the ''
polis ''Polis'' (, ; grc-gre, πόλις, ), plural ''poleis'' (, , ), literally means "city" in Greek. In Ancient Greece, it originally referred to an administrative and religious city center, as distinct from the rest of the city. Later, it also ...
'' of
Athens Athens ( ; el, Αθήνα, Athína ; grc, Ἀθῆναι, Athênai (pl.) ) is both the capital and largest city of Greece. With a population close to four million, it is also the seventh largest city in the European Union. Athens dominates a ...
, possibly because they were among the most common insects on the dry plains of
Attica Attica ( el, Αττική, Ancient Greek ''Attikḗ'' or , or ), or the Attic Peninsula, is a historical region that encompasses the city of Athens, the capital of Greece and its countryside. It is a peninsula projecting into the Aegean ...
. Native Athenians for a while wore golden grasshopper brooches to symbolise that they were of pure Athenian lineage with no foreign ancestors. In addition, Peisistratus hung the figure of a kind of grasshopper before the Acropolis of Athens as apotropaic magic.A Dictionary of Greek and Roman Antiquities (1890), Fascinum
/ref> Another symbolic use of the grasshopper is Sir Thomas Gresham's gilded grasshopper in Lombard Street, London, dating from 1563; the building was for a while the headquarters of the
Guardian Royal Exchange Guardian Royal Exchange Assurance plc was a large British insurance company. It was listed on the London Stock Exchange and was a constituent of the FTSE 100 Index. History The company was established through the merger of the Guardian Assurance ...
, but the company declined to use the symbol for fear of confusion with the locust. When grasshoppers appear in dreams, these 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".


As food

In some countries, grasshoppers are used as food. In southern
Mexico Mexico (Spanish language, Spanish: México), officially the United Mexican States, is a List of sovereign states, country in the southern portion of North America. It is borders of Mexico, bordered to the north by the United States; to the so ...
, grasshoppers, known as '' chapulines'', are eaten in a variety of dishes, such as in tortillas with chilli sauce. Grasshoppers are served on skewers in some Chinese food markets, like the
Donghuamen Night Market Donghuamen Night Market was a night market located in the northern end of Wangfujing in Beijing, China. Dong Hua Men is written 东华门, the name of the East Gate of the Forbidden City. Description One can find a row of unusual food stalls. An ...
. Fried grasshoppers (''walang goreng'') are eaten in the
Gunung Kidul Regency Gunungkidul Regency is a regency in the southeast part of the province of Yogyakarta Special Region, Indonesia. It is located on the island of Java. The regency (the name of which means South Mountains in Javanese) is bordered by the Sleman Reg ...
,
Yogyakarta Yogyakarta (; jv, ꦔꦪꦺꦴꦒꦾꦏꦂꦠ ; pey, Jogjakarta) is the capital city of Special Region of Yogyakarta in Indonesia, in the south-central part of the island of Java. As the only Indonesian royal city still ruled by a monarchy, ...
,
Java Java (; id, Jawa, ; jv, ꦗꦮ; su, ) is one of the Greater Sunda Islands in Indonesia. It is bordered by the Indian Ocean to the south and the Java Sea to the north. With a population of 151.6 million people, Java is the world's mo ...
in Indonesia. Grasshoppers are a beloved delicacy in Uganda; they are usually eaten fried (most commonly in November and May after the rains). In America, the
Ohlone The Ohlone, formerly known as Costanoans (from Spanish meaning 'coast dweller'), are a Native American people of the Northern California coast. When Spanish explorers and missionaries arrived in the late 18th century, the Ohlone inhabited the ...
burned grassland to herd grasshoppers into pits where they could be collected as food. It is recorded in the Bible that
John the Baptist John the Baptist or , , or , ;Wetterau, Bruce. ''World history''. New York: Henry Holt and Company. 1994. syc, ܝܘܿܚܲܢܵܢ ܡܲܥܡܕ݂ܵܢܵܐ, Yoḥanān Maʿmḏānā; he, יוחנן המטביל, Yohanān HaMatbil; la, Ioannes Bapti ...
ate locusts and wild honey (Greek: ἀκρίδες καὶ μέλι ἄγριον, ''akrídes kaì méli ágrion'') while living in the wilderness. However, because of a tradition of depicting him as an
ascetic Asceticism (; from the el, ἄσκησις, áskesis, exercise', 'training) is a lifestyle characterized by abstinence from sensual pleasures, often for the purpose of pursuing spiritual goals. Ascetics may withdraw from the world for their p ...
, attempts have been made to explain that the ''locusts'' were in fact a suitably ascetic
vegetarian Vegetarianism is the practice of abstaining from the consumption of meat (red meat, poultry, seafood, insects, and the flesh of any other animal). It may also include abstaining from eating all by-products of animal slaughter. Vegetariani ...
food such as
carob The carob ( ; ''Ceratonia siliqua'') is a flowering evergreen tree or shrub in the Caesalpinioideae sub-family of the legume family, Fabaceae. It is widely cultivated for its edible fruit pods, and as an ornamental tree in gardens and lands ...
beans, notwithstanding the fact that the word ''ἀκρίδες'' means plainly ''grasshoppers''. 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

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.Capinera, 2008. pp. 1710–1712 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. 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. Some grasshopper species, like the Chinese rice grasshopper, are a pest in
rice Rice is the seed of the grass species '' Oryza sativa'' (Asian rice) or less commonly ''Oryza glaberrima'' (African rice). The name wild rice is usually used for species of the genera '' Zizania'' and '' Porteresia'', both wild and domesticat ...
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. Locust plagues can have devastating effects on human populations, causing
famine A famine is a widespread scarcity of food, caused by several factors including war, natural disasters, crop failure, population imbalance, widespread poverty, an economic catastrophe or government policies. This phenomenon is usually accompan ...
s and population upheavals. They are mentioned in both the
Koran The Quran (, ; Standard Arabic: , Quranic Arabic: , , 'the recitation'), also romanized Qur'an or Koran, is the central religious text of Islam, believed by Muslims to be a revelation from God. It is organized in 114 chapters (pl.: , si ...
and the
Bible The Bible (from Koine Greek , , 'the books') is a collection of religious texts or scriptures that are held to be sacred in Christianity Christianity is an Abrahamic monotheistic religion based on the life and teachings of Jesus ...
and have also been held responsible for
cholera Cholera is an infection of the small intestine by some strains of the bacterium '' Vibrio cholerae''. Symptoms may range from none, to mild, to severe. The classic symptom is large amounts of watery diarrhea that lasts a few days. Vomiting an ...
epidemics, resulting from the corpses of locusts drowned in the Mediterranean Sea and decomposing on beaches.Capinera, 2008. pp 1181–1183 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. Besides conventional control using contact insecticides, biological pest control using the entomopathogenic fungus ''
Metarhizium acridum ''Metarhizium acridum'' is the new name given to a group of fungal isolates that are known to be virulent and specific to the Acrididea (grasshoppers). Previously, this species has had variety status in ''Metarhizium anisopliae'' (var. ''acrid ...
'', which specifically infects grasshoppers, has been used with some success.


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.


In literature

The Egyptian word for locust or grasshopper was written ''snḥm'' in the consonantal hieroglyphic writing system. The pharaoh
Ramesses II Ramesses II ( egy, rꜥ-ms-sw ''Rīʿa-məsī-sū'', , meaning "Ra is the one who bore him"; ), commonly known as Ramesses the Great, was the third pharaoh of the Nineteenth Dynasty of Egypt. Along with Thutmose III he is often regarded a ...
compared the armies of the
Hittites The Hittites () were an Anatolian people who played an important role in establishing first a kingdom in Kussara (before 1750 BC), then the Kanesh or Nesha kingdom (c. 1750–1650 BC), and next an empire centered on Hattusa in north-cent ...
to locusts: "They covered the mountains and valleys and were like locusts in their multitude." 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. Other human weaknesses besides improvidence have become identified with the grasshopper's behaviour. So an unfaithful woman (hopping from man to man) is "a grasshopper" in "Poprygunya", an 1892 short story by Anton Chekhov, and in Jerry Paris's 1969 film '' The Grasshopper''.


In mechanical engineering

The name "Grasshopper" was given to the Aeronca L-3 and Piper L-4 light aircraft, both used for
reconnaissance In military operations, reconnaissance or scouting is the exploration of an area by military forces to obtain information about enemy forces, terrain, and other activities. Examples of reconnaissance include patrolling by troops ( skirmishe ...
and other support duties in
World War II World War II or the Second World War, often abbreviated as WWII or WW2, was a world war that lasted from 1939 to 1945. It involved the World War II by country, vast majority of the world's countries—including all of the great power ...
. 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. 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.


Notes


References


Sources

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External links

* * * * {{Authority control Extant Triassic first appearances Herbivorous insects Insects in culture Insect common names