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Orthoptera () is an order 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 ...
s that comprises the grasshoppers,
locust Locusts (derived from the Vulgar Latin ''locusta'', meaning grasshopper) are various species of short-horned grasshoppers in the family Acrididae that have a swarming phase. These insects are usually solitary, but under certain circumstanc ...
s, and
cricket Cricket is a bat-and-ball game played between two teams of eleven players on a field at the centre of which is a pitch with a wicket at each end, each comprising two bails balanced on three stumps. The batting side scores runs by str ...
s, including closely related insects, such as the bush crickets or katydids and
wētā Wētā (also spelt weta) is the common name for a group of about 100 insect species in the families Anostostomatidae and Rhaphidophoridae endemic to New Zealand. They are giant flightless crickets, and some are among the heaviest insects in ...
. The order is subdivided into two suborders:
Caelifera The Caelifera are a suborder of orthopteran insects. They include the grasshoppers and grasshopper-like insects, as well as other superfamilies classified with them: the ground-hoppers (Tetrigoidea) and pygmy mole crickets ( Tridactyloidea). ...
– grasshoppers, locusts, and close relatives; and
Ensifera Ensifera is a suborder of insects that includes the various types of crickets and their allies including: true crickets, camel crickets, bush crickets or katydids, grigs, weta and Cooloola monsters. This and the suborder Caelifera (grassh ...
– crickets and close relatives. More than 20,000
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
are distributed worldwide. The insects in the order have
incomplete metamorphosis Hemimetabolism or hemimetaboly, also called incomplete metamorphosis and paurometabolism,McGavin, George C. ''Essential Entomology: An Order-by-Order Introduction''. Oxford: Oxford University Press, 2001. pp. 20. is the mode of development of certa ...
, and produce
sound In physics, sound is a vibration that propagates as an acoustic wave, through a transmission medium such as a gas, liquid or solid. In human physiology and psychology, sound is the ''reception'' of such waves and their ''perception'' b ...
(known as a "
stridulation 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 ...
") by rubbing their wings against each other or their legs, the wings or legs containing rows of corrugated bumps. The tympanum, or
ear An ear is the organ that enables hearing and, in mammals, body balance using the vestibular system. In mammals, the ear is usually described as having three parts—the outer ear, the middle ear and the inner ear. The outer ear consists of ...
, is located in the front
tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
in crickets,
mole cricket Mole crickets are members of the insect family Gryllotalpidae, in the order Orthoptera (grasshoppers, locusts, and crickets). Mole crickets are cylindrical-bodied, fossorial insects about long as adults, with small eyes and shovel-like fore ...
s, and bush crickets or katydids, and on the first abdominal segment in the grasshoppers and locusts. These organisms use vibrations to locate other individuals. Grasshoppers and other orthopterans are able to fold their
wings A wing is a type of fin that produces lift while moving through air or some other fluid. Accordingly, wings have streamlined cross-sections that are subject to aerodynamic forces and act as airfoils. A wing's aerodynamic efficiency is expre ...
(i.e. they are members of
Neoptera Neoptera (Ancient Greek ''néos'' (“new”) + ''pterón'' (“wing”)) is a classification group that includes most orders of the winged insects, specifically those that can flex their wings over their abdomens. This is in contrast with the mo ...
).


Etymology

The name is derived from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
ὀρθός ''orthos'' meaning "straight" and πτερόν ''pteron'' meaning "wing".


Characteristics

Orthopterans have a generally cylindrical body, with elongated hindlegs and musculature adapted for
jumping Jumping or leaping is a form of locomotion or movement in which an organism or non-living (e.g., robotics, robotic) mechanical system propels itself through the air along a ballistic trajectory. Jumping can be distinguished from running, gallo ...
. They have
mandibulate Mandibulata, termed "mandibulates", is a clade of arthropods that comprises the extant subphyla Myriapoda (millipedes and others), Crustacea and Hexapoda (insects and others). Mandibulata is currently believed to be the sister group of the clade ...
mouthparts for biting and chewing and large
compound eye A compound eye is a visual organ found in arthropods such as insects and crustaceans. It may consist of thousands of ommatidia, which are tiny independent photoreception units that consist of a cornea, lens, and photoreceptor cells which disti ...
s, and may or may not have
ocelli A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
, depending on the species. The antennae have multiple joints and filiform type, and are of variable length. The first and third segments on the
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 cre ...
are larger, while the second segment is much smaller. They have two pairs of
wings A wing is a type of fin that produces lift while moving through air or some other fluid. Accordingly, wings have streamlined cross-sections that are subject to aerodynamic forces and act as airfoils. A wing's aerodynamic efficiency is expre ...
, which are held overlapping the
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 ...
at rest. The forewings, or
tegmina A tegmen (plural: ''tegmina'') designates the modified leathery front wing on an insect particularly in the orders Dermaptera ( earwigs), Orthoptera (grasshoppers, crickets and similar families), Mantodea (praying mantis), Phasmatodea (stick an ...
, are narrower than the hindwings and hardened at the base, while the hindwings are membranous, with straight veins and numerous cross-veins. At rest, the hindwings are held folded fan-like under the forewings. The final two to three segments of the abdomen are reduced, and have single-segmented cerci.


Life cycle

Orthopterans have a paurometabolous lifecycle or
incomplete metamorphosis Hemimetabolism or hemimetaboly, also called incomplete metamorphosis and paurometabolism,McGavin, George C. ''Essential Entomology: An Order-by-Order Introduction''. Oxford: Oxford University Press, 2001. pp. 20. is the mode of development of certa ...
. The use of sound is generally crucial in courtship, and most species have distinct songs. Most grasshoppers lay their
eggs Humans and human ancestors have scavenged and eaten animal eggs for millions of years. Humans in Southeast Asia had domesticated chickens and harvested their eggs for food by 1,500 BCE. The most widely consumed eggs are those of fowl, especial ...
in the ground or on vegetation. The eggs hatch and the young
nymphs 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 ...
resemble adults, but lack wings and at this stage are often called 'hoppers'. They may often also have a radically different coloration from the adults. Through successive moults, the nymphs develop wings until their final moult into a mature adult with fully developed wings. The number of moults varies between species; growth is also very variable and may take a few weeks to some months depending on food availability and weather conditions.


Evolution

This order evolved with a division into two suborders -
Caelifera The Caelifera are a suborder of orthopteran insects. They include the grasshoppers and grasshopper-like insects, as well as other superfamilies classified with them: the ground-hoppers (Tetrigoidea) and pygmy mole crickets ( Tridactyloidea). ...
and
Ensifera Ensifera is a suborder of insects that includes the various types of crickets and their allies including: true crickets, camel crickets, bush crickets or katydids, grigs, weta and Cooloola monsters. This and the suborder Caelifera (grassh ...
- occurring .Chang H, Qiu Z, Yuan H, Wang X, Li X, Sun H, Guo X, Lu Y, Feng X, Majid M, Huang Y (2020) Evolutionary rates of and selective constraints on the mitochondrial genomes of Orthoptera insects with different wing types. Mol Phylogenet Evol


Phylogeny

The Orthoptera are divided into two suborders,
Caelifera The Caelifera are a suborder of orthopteran insects. They include the grasshoppers and grasshopper-like insects, as well as other superfamilies classified with them: the ground-hoppers (Tetrigoidea) and pygmy mole crickets ( Tridactyloidea). ...
and
Ensifera Ensifera is a suborder of insects that includes the various types of crickets and their allies including: true crickets, camel crickets, bush crickets or katydids, grigs, weta and Cooloola monsters. This and the suborder Caelifera (grassh ...
, that have been shown to be monophyletic.Zhou Z, Ye H, Huang Y, Shi F. (2010) The phylogeny of Orthoptera inferred from mtDNA and description of ''Elimaea cheni'' (Tettigoniidae: Phaneropterinae) mitogenome. ''J. Genet. Genomics.'' 37(5):315-324


Taxonomy

Taxonomists classify members of the Caelifera and Ensifera into infraorders and superfamilies as follows: * Suborder
Caelifera The Caelifera are a suborder of orthopteran insects. They include the grasshoppers and grasshopper-like insects, as well as other superfamilies classified with them: the ground-hoppers (Tetrigoidea) and pygmy mole crickets ( Tridactyloidea). ...
– grasshoppers, pygmy mole crickets and allies ** Infraorder
Acrididea Acrididea including the Acridomorpha is an infraorder of insects that describe the grasshoppers (thus also locusts) and ground-hoppers. It contains a large majority of species in the suborder Caelifera and the taxon Acridomorpha may also be use ...
*** Superfamily
Acridoidea Acridoidea is the largest superfamily of grasshoppers in the order Orthoptera with species found on every continent except Antarctica. Classification ''Orthoptera Species File'' includes the following families: *Acrididae MacLeay, 1821 * Derico ...
– grasshoppers, locusts *** Superfamily
Eumastacoidea Eumastacoidea is a superfamily within the order Orthoptera, suborder Caelifera. The family has a mainly tropical distribution and have sometimes been called "monkey grasshoppers". Description Some of the characters of the members of the superf ...
– monkey or matchstick grasshoppers and allies *** Superfamily Locustopsoidea† *** Superfamily
Pneumoroidea The Pneumoridae are a family of nocturnal short-horned grasshoppers in the order Orthoptera, commonly known as the bladder grasshoppers and the sole representative of the superfamily Pneumoroidea. Their centre of diversity is in southern Africa, ...
– bladder grasshoppers *** Superfamily
Pyrgomorphoidea Pyrgomorphidae is a family of grasshoppers in the order Orthoptera; it is the only family in the superfamily Pyrgomorphoidea, with a pan-tropical distribution. Their name is probably derived from ''pyrgos'' (Greek: ''Πύργος'') meaning "tow ...
– gaudy grasshoppers *** Superfamily Tanaoceroidea – desert long-horned grasshoppers *** Superfamily
Tetrigoidea Tetrigidae is an ancient family in the order Orthoptera, which also includes similar families such as crickets, grasshoppers, and their allies. Species within the Tetrigidae are variously called groundhoppers, pygmy grasshoppers,Borror DJ, Trippl ...
– ground-hoppers or grouse locusts *** Superfamily
Trigonopterygoidea The Trigonopterygoidea are an insect superfamily in the Orthoptera: Caelifera.Tridactylidea *** Superfamily Dzhajloutshelloidea† *** Superfamily Regiatoidea† *** Superfamily Tridactyloidea – pygmy mole crickets and allies * Suborder
Ensifera Ensifera is a suborder of insects that includes the various types of crickets and their allies including: true crickets, camel crickets, bush crickets or katydids, grigs, weta and Cooloola monsters. This and the suborder Caelifera (grassh ...
– crickets ** Superfamily
Grylloidea Grylloidea is the superfamily of insects, in the order Orthoptera, known as crickets. It includes the " true crickets", scaly crickets, wood crickets and other families, some only known from fossils. Grylloidea dates from the Triassic period a ...
– crickets, mole crickets ** Superfamily Hagloidea – grigs and allies ** Superfamily Phasmomimoidea† ** Superfamily
Rhaphidophoroidea The orthopteran family Rhaphidophoridae of the suborder Ensifera has a worldwide distribution. Common names for these insects include cave wētā, cave crickets, camelback crickets, camel crickets, Hogan bugs, spider crickets (sometimes shorte ...
– camel crickets, cave crickets, cave wētā ** Superfamily
Schizodactyloidea Schizodactylidae is a family of orthopteran insects found in Asia and southern Africa, known as dune crickets or splay-footed crickets. They are usually found in desert and sandy areas. Species are predatory, including ''Schizodactylus inexspecta ...
– dune crickets ** Superfamily
Stenopelmatoidea Stenopelmatoidea is a superfamily of insects in the order Orthoptera; in some older classifications this group was referred to as Gryllacridoidea. Classification The classification and constituency of Stenopelmatoidea is an ongoing source of ...
– wētā and allies ** Superfamily
Tettigonioidea 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, t ...
– katydids / bush crickets *''Incertae sedis'' **Superfamily Elcanoidea
Permian The Permian ( ) is a geologic period and System (stratigraphy), stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last ...
-
Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch that lasted from about 66 to 56 million years ago (mya). It is the first epoch of the Paleogene Period in the modern Cenozoic Era. The name is a combination of the Ancient Greek ''pal ...


Relationships with humans


As pests

Several species of Orthoptera are considered pests of crops and rangelands or seeking warmth in homes by humans. The two groups of Orthoptera that cause the most damage are grasshoppers and
locust Locusts (derived from the Vulgar Latin ''locusta'', meaning grasshopper) are various species of short-horned grasshoppers in the family Acrididae that have a swarming phase. These insects are usually solitary, but under certain circumstanc ...
s. Locust are historically known for wiping out fields of crops in a day. Locust have the ability to eat up to their own body weight in a single day. Individuals gather in large groups called swarms, these swarms can range up to 80 million individuals that stretch 460 square miles. Grasshoppers can cause major agricultural damage but not to the documented extent as locust historically have. These insects mainly feed on weeds and grasses, however, during times of drought and high population density they will feed on crops. They are known pest in
soybean The soybean, soy bean, or soya bean (''Glycine max'') is a species of legume native to East Asia, widely grown for its edible bean, which has numerous uses. Traditional unfermented food uses of soybeans include soy milk, from which tofu a ...
fields and will likely feed on these crops once preferred food sources have become scarce.


As food

Most orthopterans are edible, making up 13% of all insects including some 80 species of grasshoppers being regularly consumed worldwide. In
Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Africa ...
and
Oaxaca Oaxaca ( , also , , from nci, Huāxyacac ), officially the Free and Sovereign State of Oaxaca ( es, Estado Libre y Soberano de Oaxaca), is one of the 32 states that compose the political divisions of Mexico, Federative Entities of Mexico. It is ...
, grasshoppers and locusts are usually collected early in the morning when it is cooler as the orthopterans are less mobile due to being cold-blooded. In Thailand, house crickets are commonly reared and eaten; as of 2012, around 20,000 cricket farmers had farms in 53 of their 76 provinces. In the second century BCE in
Ancient Greece Ancient Greece ( el, Ἑλλάς, Hellás) was a northeastern Mediterranean Sea, Mediterranean civilization, existing from the Greek Dark Ages of the 12th–9th centuries BC to the end of Classical Antiquity, classical antiquity ( AD 600), th ...
, Diodorus Siculus is known to have called people from
Ethiopia Ethiopia, , om, Itiyoophiyaa, so, Itoobiya, ti, ኢትዮጵያ, Ítiyop'iya, aa, Itiyoppiya officially the Federal Democratic Republic of Ethiopia, is a landlocked country in the Horn of Africa. It shares borders with Eritrea to the ...
''Acridophagi'', meaning "eaters of locusts." In
Judaism Judaism ( he, ''Yahăḏūṯ'') is an Abrahamic, monotheistic, and ethnic religion comprising the collective religious, cultural, and legal tradition and civilization of the Jewish people. It has its roots as an organized religion in t ...
, the Orthoptera include the only insects considered kosher. The list of dietary laws in the
book of Leviticus The book of Leviticus (, from grc, Λευιτικόν, ; he, וַיִּקְרָא, , "And He called") is the third book of the Torah (the Pentateuch) and of the Old Testament, also known as the Third Book of Moses. Scholars generally agree ...
forbids all flying insects that walk, but makes an exception for certain locusts. Notably, the dragonfly and
crane fly Crane fly is a common name referring to any member of the insect family Tipulidae. Cylindrotominae, Limoniinae, and Pediciinae have been ranked as subfamilies of Tipulidae by most authors, though occasionally elevated to family rank. In the m ...
are not considered kosher even though they are helpless when unable to fly. The
Torah The Torah (; hbo, ''Tōrā'', "Instruction", "Teaching" or "Law") is the compilation of the first five books of the Hebrew Bible, namely the books of Genesis, Exodus, Leviticus, Numbers and Deuteronomy. In that sense, Torah means the ...
states the only kosher flying insects with four walking legs have knees that extend above their feet so that they hop.


As creators of biofuel

With new research showing promise in locating alternative biofuel sources in the gut of insects, grasshoppers are one species of interest. The insect's ability to break down
cellulose Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell w ...
and lignin without producing
greenhouse gases A greenhouse gas (GHG or GhG) is a gas that absorbs and emits radiant energy within the thermal infrared range, causing the greenhouse effect. The primary greenhouse gases in Earth's atmosphere are water vapor (), carbon dioxide (), methane ...
has aroused scientific interest.


See also

* List of Orthoptera recorded in Britain *
Orthopterida The cohort Polyneoptera is a proposed taxonomic ranking for the Orthoptera (grasshoppers, crickets, etc.) and all other Neopteran insects believed to be more closely related to Orthoptera than to any other insect orders. These winged insects, no ...
*
Female sperm storage Female sperm storage is a biological process and often a type of sexual selection in which sperm cells transferred to a female during mating are temporarily retained within a specific part of the reproductive tract before the oocyte, or egg, is fe ...


References


External links


Orthoptera Species File OnlineOrthoptera Image Gallery (Iowa State University Entomology Department)Australian Plague Locust CommissionThe Orthopterists' SocietyAcridAfrica, les acridiens d'Afrique de l'Ouest
*
Birdwing Grasshoppers in BelizeSound recordings of Orthoptera at BioAcoustica
{{Authority control Insect orders Edible insects Carboniferous first appearances Taxa named by Pierre André Latreille Orthopterida