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 ...
of
orthoptera
Orthoptera () is an order of insects that comprises the grasshoppers, locusts, and crickets, including closely related insects, such as the bush crickets or katydids and wētā. The order is subdivided into two suborders: Caelifera – grass ...
n
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 pairs ...
s. They include the grasshoppers and grasshopper-like insects, as well as other superfamilies classified with them: the ground-hoppers ( Tetrigoidea) and pygmy mole crickets ( Tridactyloidea). The latter should not be confused with the mole crickets ( Gryllotalpidae), which belong to the other Orthopteran sub-order Ensifera.
The name of this suborder comes from
Latin
Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power ...
meaning ''
chisel
A chisel is a tool with a characteristically shaped cutting edge (such that wood chisels have lent part of their name to a particular grind) of blade on its end, for carving or cutting a hard material such as wood, stone, or metal by hand, s ...
-bearing'' ("chisel" in Latin: ''caelum''), referring to the "stout" shape of its species'
ovipositor
The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typica ...
s.
Subdivisions and their distribution
The Caelifera include some 2,400 valid genera and about 12,000 known species. Many undescribed species probably exist, especially in
tropical forests
Tropical forests (a.k.a. jungle) are forested landscapes in tropical regions: ''i.e.'' land areas approximately bounded by the tropic of Cancer and Capricorn, but possibly affected by other factors such as prevailing winds.
Some tropical fore ...
. The Caelifera have a predominantly tropical distribution (as with most Orthoptera) with fewer species known from
temperate climate
In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout ...
zones. Caelifera are divided into two infraorders: the more basal Tridactylidea and the Acrididea or grasshopper-like species. This latter name is derived from older sources, such as
Imms
The Institute for Marine Mammal Studies ("IMMS") is a research organization located in Gulfport, Mississippi and dedicated to education, conservation, and research on marine mammals in the wild and in captivity. It was founded in 1984 as a researc ...
,Imms AD, rev. Richards OW & Davies RG (1970) ''A General Textbook of Entomology'' 9th Ed. Methuen 886 pp. which placed the "short-horned grasshoppers" and locusts at the family level ( Acrididae).
* Infraorder Tridactylidea
** Tridactyloidea Brullé, 1835: pygmy mole crickets - all continents except Antarctica
** Dzhajloutshelloidea Gorochov, 1994 †
* Infraorder Acrididea
** Tetrigoidea (
monotypic
In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...
) Serville, 1838: groundhoppers or "grouse locusts" - all continents except Antarctica
** Informal superfamily group Acridomorpha - the grasshoppers
*** Acridoidea MacLeay, 1821: most grasshoppers – World-wide - approx. 10,000 species in the Acrididae alone
*** Eumastacoidea Burr, 1899: "monkey grasshoppers" - Americas,
Africa
Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
,
Australasia
Australasia is a region that comprises Australia, New Zealand and some neighbouring islands in the Pacific Ocean. The term is used in a number of different contexts, including geopolitically, physiogeographically, philologically, and ecolo ...
Africa
Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
south America
South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the souther ...
monotypic
In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...
) B. von Wattenwyl, 1882: "gaudy grasshoppers" - all tropical/subtropical continents
*** Tanaoceroidea (
monotypic
In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...
) Rehn, 1948: "desert long-horned grasshoppers" - north America
***
Trigonopterygoidea
The Trigonopterygoidea are an insect superfamily in the Orthoptera: Caelifera.
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 spe ...
of the Caelifera, is described in detail for
grasshoppers
Grasshoppers are a group of insects belonging to the suborder Caelifera. They are among what is possibly the most ancient living group of chewing herbivorous insects, dating back to the early Triassic around 250 million years ago.
Grasshop ...
, with 6 out of 8 extant superfamilies shown here as a
cladogram
A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
. Like the Ensifera, Caelifera and all of its superfamilies appear to be
monophyletic
In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic ...
.
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 spe ...
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 ribosomal ...
of thirty-two taxa in six out of seven superfamilies, is shown as a
cladogram
A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
. The Ensifera, Caelifera and all the superfamilies of grasshoppers except Pamphagoidea appear to be
monophyletic
In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. 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 '' Eolocustopsis'' from the latest Permian (
Changhsingian
In the geologic time scale, the Changhsingian or Changxingian is the latest age or uppermost stage of the Permian. It is also the upper or latest of two subdivisions of the Lopingian Epoch or Series. The Changhsingian lasted from to 251.902 m ...
) of the
Beaufort Group
The Beaufort Group is the third of the main subdivisions of the Karoo Supergroup in South Africa. It is composed of a lower Adelaide Subgroup and an upper Tarkastad Subgroup. It follows conformably after the Ecca Group and unconformably underli ...
, South Africa and Locustavidae from the early Triassic, roughly 250 million years ago. The oldest known member of Caelifera is 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
The Early Jurassic Epoch (in chronostratigraphy corresponding to the Lower Jurassic Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic-Jurassic extinction event, 201.3 Ma  ...
. 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 s ...
(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.
Economic significance and terminology
A number of species, especially in the Acridoidea, are significant
agricultural pest
A pest is any animal or plant harmful to humans or human concerns. The term is particularly used for creatures that damage crops, livestock, and forestry or cause a nuisance to people, especially in their homes. Humans have modified the environ ...
s, but not all of them are
locusts
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 circumst ...
: a non-taxonomic term referring to species whose populations which may change morphologically when crowded and show
swarming behaviour
Swarm behaviour, or swarming, is a collective behaviour exhibited by entities, particularly animals, of similar size which aggregate together, perhaps milling about the same spot or perhaps moving ''en masse'' or migrating in some direction. ...
.Uvarov BP (1966) ''Grasshoppers & Locusts. A Handbook of General Acridology'' Cambridge University Press, London 1:481 pp. Examples of agricultural grasshopper pests that are not called locusts include the Senegalese grasshopper and certain species in the
Pyrgomorphidae
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 "to ...
, notably the variegated grasshopper (''
Zonocerus variegatus
''Zonocerus variegatus'', the painted grasshopper or variegated grasshopper, is a species of insect belonging to the family Pyrgomorphidae. It is native to tropical Africa, and is considered a crop pest in much of Western and Central Africa.
Des ...