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Grasshopper

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.

Phylogeny
Grasshoppers belong to the suborder Caelifera. Although "grasshopper" has been used as a common name for the suborder in general, modern sources restrict it to the more "evolved" families. They may be placed in the infraorder Acrididea and have been referred to as "short-horned grasshoppers" in older texts to distinguish them from the also-obsolete term "long-horned grasshoppers" (now bush-crickets or katydids) with their much longer antennae. The phylogeny of Caelifera, based on mitochondrial ribosomal RNA of thirty-two taxa in six out of seven superfamilies, is shown as a cladogram. Ensifera, 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, 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 palaeontological taxonomy is founded principally on the venation of the hindwings. 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. ==Characteristics==
Characteristics
s, with spines along the tibia of the hind legs 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. They have two strong hind legs that propel them in the air to escape predators. The downward-directed mouthparts are modified for chewing and there are two sensory palps in front of the jaws. 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. 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. 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. Others, like the Crackling Forest Grasshopper, produce distinctive clicks while flying; this behavior is called crepitation. 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==
Biology
Diet and digestion Most grasshoppers are polyphagous, eating vegetation from multiple plant sources e.g pea plant leaves , but some are omnivorous and also eat animal tissue and animal meat. They also like to eat other insects. In general their preference is for grasses, including many cereals 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, which by softening plant cell walls makes plant cell contents accessible to other digestive enzymes. Grasshoppers can also be cannibalistic when swarming. Sensory organs '') 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. 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. 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. 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. Jumping for a jump, overcoming the limitations of muscle which cannot contract powerfully and quickly at the same time. Representations of structure are diagrammatic. 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 (20 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 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. 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, 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, 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. Female grasshoppers of the species Chorthippus biguttulus appear to be able to integrate information from male calling songs. An unattractive song subunit far outweighs an attractive song subunit, and this asymmetrical integration is consistent with theories of sexual selection because it helps females avoid potentially costly interaction with unsuitable mating partners if the song belongs to another species or indicates a low-quality male. 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. 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. 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 in 1875; the swarm was long and wide, and one estimate puts the number of locusts involved at 3.5 trillion. An adult desert locust 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==
Predators, parasites, and pathogens
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, robber flies and sphecid wasps, by spiders, and by many birds and small mammals including dogs and cats. 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. s 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. 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. s 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. 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. 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. 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. 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. 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. 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. Atractomorpha lata, Burdwan, West Bengal, India 27 10 2012.jpg|Gaudy grasshopper, Atractomorpha lata, evades predators with camouflage. Titanacris Albipes Vol.jpg|Lubber grasshopper, Titanacris albipes, has deimatically coloured wings, used to startle predators. LeafGrasshopper.jpg|Leaf grasshopper, Phyllochoreia ramakrishnai, mimics a green leaf. Dactylotum bicolor.jpg|Painted grasshopper, Dactylotum bicolor, deters predators with warning coloration. 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==
Relationship with humans
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. 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 featured in cinema. The 1957 film Beginning of the End portrayed giant grasshoppers attacking Chicago. 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. File:Balthasar van der Ast - Flowers in a Vase with Shells and Insects - WGA1042.jpg|Balthasar van der Ast's oil painting Flowers in a Vase with Shells and Insects, c. 1630 File: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 File:Akragas, tetradracma, 410 ac. ca.JPG|Ancient Greek tetradrachm coin, 410 BC, with grasshopper at right Symbolism 's gilded grasshopper symbol, Lombard Street, London, 1563 Grasshoppers are sometimes used as symbols. possibly because they were among the most common insects on the dry plains of Attica. 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, but the company declined to use the symbol for fear of confusion with the locust. 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". As food , Indonesia '') In some countries, grasshoppers are used as food. In southern Mexico, grasshoppers, known as chapulines, are eaten in dishes such as in tortillas with chilli sauce. Grasshoppers are served on skewers in some Chinese food markets. Fried grasshoppers (walang goreng) are eaten in the Gunung Kidul Regency in Indonesia. Grasshoppers are a delicacy in Uganda; they are eaten fried. In America, the Ohlone burned grassland to herd grasshoppers into pits where they could be collected as food. Among ancient Jews, grasshoppers were considered a popular and even luxurious food, as attested by rabbinic sources. John the Baptist is recorded as having eaten locusts while living in the wilderness. 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. 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. 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, Detection of explosives In February 2020, researchers from Washington University in St. Louis announced they had engineered "cyborg grasshoppers" capable of accurately detecting explosives. Grasshoppers were fitted with lightweight sensor backpacks that recorded and transmitted the electrical activity of their antennal lobes to a computer. 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 compared the armies of the Hittites 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. and in Jerry Paris's 1969 film The Grasshopper. In mechanical engineering , 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. 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==
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