Grasshopper
Grasshoppers | |
---|---|
American grasshopper (Schistocerca americana) | |
Scientific classification | |
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
Grasshoppers are typically ground-dwelling insects with powerful hind legs which allow them to escape from threats by leaping vigorously. Their front leg is shorter and used for grasping food. As
Grasshoppers are plant-eaters, with a few species at times becoming serious
Grasshoppers have had a long relationship with humans. Swarms of locusts can have devastating effects and cause famine, having done so since
Phylogeny
Grasshoppers belong to the suborder Caelifera. Although "grasshopper" has been used as a common name for the suborder in general,
Orthoptera |
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In evolutionary terms, the split between the Caelifera and the Ensifera is no more recent than the
The Caelifera includes some 2,400 valid genera and about 11,000 known species. Many undescribed species probably exist, especially in
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,
The thorax and abdomen are segmented and have a rigid
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.[17]
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.[18]
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.[19] 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.[18]
Biology
Diet and digestion
Most grasshoppers are
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
Circulation and respiration
Like other insects, grasshoppers have an
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.[31] 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.[32]
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.[33]
The jump is a three-stage process.[34] 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.[35] 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.[36] 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.[37]
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.[38] In male grasshoppers of the family Pneumoridae, the enlarged abdomen amplifies stridulation.[15]
Life cycle
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.[39]
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.[40] 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.[41]
Grasshoppers undergo
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.[42] 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.[43]
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,[44] and one estimate puts the number of locusts involved at 3.5 trillion.[45] 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.[46]
Predators, parasites, and pathogens
Grasshoppers have a wide range of
The eggs and nymphs are under attack by
The grasshopper nematode (Mermis nigrescens) is a long slender worm that infects grasshoppers, living in the insect's hemocoel. Adult worms lay eggs on plants and the host becomes infected when the foliage is eaten.[49] 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.[50][51]
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 insect's temperature above a threshold tolerated by the fungus.[53] 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.[54]
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.[53] 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.[53]
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.[55]
Several species such as the hooded leaf grasshopper
Some species are genuinely
-
Gaudy grasshopper, Atractomorpha lata, evades predators with camouflage.
-
Lubber grasshopper,deimaticallycoloured wings, used to startle predators.
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Leaf grasshopper, Phyllochoreia ramakrishnai, mimics a green leaf.
-
Painted grasshopper,warning coloration.
-
Spotted grasshopper,Aularches miliaris, defends itself with toxic foam and warning colours.[60]
Relationship with humans
In art and media
Grasshoppers are occasionally depicted in artworks, such as the
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.[62][63]
Grasshoppers are also featured in cinema. The 1957 film
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
Grasshoppers are sometimes used as symbols.
Another symbolic use of the grasshopper is Sir
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".[70]
As food
In some countries, grasshoppers are used as food.[71] In southern Mexico, grasshoppers, known as chapulines, are eaten in a variety of dishes, such as in tortillas with chilli sauce.[72] Grasshoppers are served on skewers in some Chinese food markets, like the Donghuamen Night Market.[73] Fried grasshoppers (walang goreng) are eaten in the Gunung Kidul Regency, Yogyakarta, Java in Indonesia.[74] Grasshoppers are a beloved delicacy in Uganda; they are usually eaten fried (most commonly in November and May after the rains).[75] In America, the Ohlone burned grassland to herd grasshoppers into pits where they could be collected as food.[76]
It is recorded in the Bible that
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.[80]
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.[81] 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.[80]
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.[82]
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.[46] 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.[83] Besides conventional control using contact insecticides,[83] biological pest control using the entomopathogenic fungus Metarhizium acridum, which specifically infects grasshoppers, has been used with some success.[84]
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.[85][86]
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."[87]
One of
In mechanical engineering
The name "Grasshopper" was given to the
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.[94][95][96]
Notes
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- Cott, Hugh (1940). Adaptive Coloration in Animals. Oxford University Press.
- Pfadt, Robert E. (1994). Field Guide to Common Western Grasshoppers (2nd ed.). Wyoming Agricultural Experiment Station.
External links
- Media related to Caelifera at Wikimedia Commons
- Quotations related to Grasshoppers at Wikiquote
- Data related to Caelifera at Wikispecies
- The dictionary definition of grasshopper at Wiktionary