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Ensifera is a
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
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 that includes the various types of crickets and their allies including: true crickets, camel crickets, bush crickets or katydids, grigs, weta and
Cooloola monster The Cooloola monster (''Cooloola propator'') is a large burrowing orthopteran of the family Cooloolidae, a family erected to accommodate it because it is so dissimilar to other ensiferans. It was discovered in 1980 in the Great Sandy National P ...
s. This and the 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 and their allies) make up the order
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 – grassho ...
. Ensifera is believed to be a more ancient group than Caelifera, with its origins in the
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...
period, the split having occurred at the end of the
Permian The Permian ( ) is a geologic period and 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 period of the Paleoz ...
period. Unlike the Caelifera, the Ensifera contain numerous members that are partially carnivorous, feeding on other insects, as well as plants. ''Ensifer'' is
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 of the ...
for "sword bearer", and refers to the typically elongated and blade-like
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 typical ...
of the females.


Characteristics

Characteristics shared by the two orthopteran suborders, Caelifera and Ensifera, are the mouthparts adapted for biting and chewing, the modified prothorax, the hind legs modified for jumping, the wing shape and venation, and the sound-producing stridulatory organs. Ensiferans are distinguished from Caeliferans by their elongated, threadlike antennae, which are often longer than the length of their bodies and have over 30 segments (except in the subterranean
Cooloolidae ''Cooloola'' is a genus of ensiferan orthopterans known as Cooloola monsters. It is the only genus in the subfamily Cooloolinae and family Cooloolidae of the superfamily Stenopelmatoidea. Four species are known from this family, all endemic to ...
family). For this reason, they are sometimes referred to as "long-horned orthopterans". In the families in which the males sing, the fore wings have modifications that include toothed veins and scrapers for making the noise, and the surrounding membranous areas amplify the sound. In these groups, the sound-detecting
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 ...
s are located on the tibiae of the front legs. The tarsi have three segments and the
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 typical ...
is blade-like or needle-like. The male attaches the
spermatophore A spermatophore or sperm ampulla is a capsule or mass containing spermatozoa created by males of various animal species, especially salamanders and arthropods, and transferred in entirety to the female's ovipore during reproduction. Spermatophores ...
externally to the female's
gonopore A gonopore, sometimes called a gonadopore, is a genital pore in many invertebrates. Hexapods, including insects have a single common gonopore, except mayflies, which have a pair of gonopores. More specifically, in the unmodified female it is the ...
. The spermatophore is often surrounded by a
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
aceous
spermatophylax A spermatophylax is a gelatinous bolus which some male insects eject during copulation with females through their aedeagi together with spermatophores, and which functions as a nutritive supplement for the female. See also *Nuptial gift A nupt ...
, the function of which is to provide a nutritional
nuptial gift A nuptial gift is a nutritional gift given by one partner in some animals' sexual reproduction practices. Formally, a nuptial gift is a material presentation to a recipient by a donor during or in relation to sexual intercourse that is not simpl ...
to the female.


Taxonomy

The Orthoptera Species File database lists the following superfamilies and families. *Infraorder † Elcanidea **Superfamily † Elcanoidea ***Family †
Elcanidae Elcanidae are an extinct family of Mesozoic and early Cenozoic orthopterans. Members of the family are distinguished by the presence of spurs on the distal part of the metatibia, unique among orthopterans, these have been suggested to have been ...
(Late Triassic - Paleocene) ***Family † Permelcanidae (Early Permian - Late Triassic) **Superfamily † Permoraphidioidea ***Family † Permoraphidiidae (Permian) ***Family † Pseudelcanidae (Early Permian) ***Family † Thueringoedischiidae (Early Permian) ***Family ''
incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
'' ****Genus †'' Acridiites'' *Infraorder Gryllidea **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 ...
***Family † Baissogryllidae ***Family
Gryllidae The family ''Gryllidae'' contains the subfamilies and genera which entomologists now term true crickets. Having long, whip-like antennae, they belong to the Orthopteran suborder Ensifera, which has been greatly reduced in the last 100 years (''e ...
- true crickets ***Family
Mogoplistidae Mogoplistidae is a family of scaly crickets within the superfamily Grylloidea. Considered to be monophyletic, a sister taxon to the Gryllidae crickets. This family consists of more than 370 species worldwide; 20 species in 4 genera occur in North ...
- scaly crickets ***Family
Phalangopsidae The Phalangopsidae are a recently reconstituted family of crickets (Orthoptera: Ensifera), based on the type genus '' Phalangopsis'' Serville, 1831 from South America. Priority for family-group names based on this genus dates from Blanchard's "' ...
***Family † Protogryllidae ***Family
Trigonidiidae The Trigonidiidae are a family of crickets: Grylloidea consisting of two subfamilies: * Subfamily Nemobiinae Saussure, 1877 – wood crickets or ground crickets * Subfamily Trigonidiinae Trigonidiinae is a subfamily of insects in the order ...
**Superfamily Gryllotalpoidea ***Family
Gryllotalpidae 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 ...
– mole crickets ***Family
Myrmecophilidae The Myrmecophilidae or ant-loving crickets are rarely encountered relatives of mole crickets, and are obligate inquilines within ant nests. They are very small, wingless, and flattened, so resemble small cockroach nymphs. The few genera contain fe ...
- ant crickets *Infraorder † Oedischiidea **Superfamily † Oedischioidea ***Family † Anelcanidae ***Family † Bintoniellidae ***Family † Mesoedischiidae ***Family † Oedischiidae ***Family † Proparagryllacrididae ***Family † Pruvostitidae ***Family ''incertae sedis'' ****Genus †'' Crinoedischia'' ****Genus †'' Loxoedischia'' **Superfamily † Triassomantoidea ***Family † Adumbratomorphidae ***Family † Triassomantidae **Superfamily † Xenopteroidea ***Family † Xenopteridae **Superfamily ''incertae sedis'' ***family † Permotettigoniidae ***Family ''incertae sedis'' ****Genus †'' Permophyllum'' *Infraorder Tettigoniidea **Superfamily Hagloidea - grigs ***Family † Eospilopteronidae ***Family † Haglidae ***Family † Hagloedischiidae ***Family † Prezottophlebiidae ***Family
Prophalangopsidae The family Prophalangopsidae are insects belonging to the order Orthoptera. They are the only extant members of the superfamily Hagloidea. There is only one extant genus in North America, where they are known as grigs, four genera in Asia, an ...
***Family † Tuphellidae **Superfamily † Phasmomimoidea ***Family † Phasmomimidae **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 ...
***Family
Rhaphidophoridae 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 shortene ...
- 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 ...
***Family Schizodactylidae - dune or splay-footed 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 co ...
***Family
Anostostomatidae Anostostomatidae is a family of insects in the order Orthoptera, widely distributed in the southern hemisphere. It is named Mimnermidae or Henicidae in some taxonomies, and common names include ''king crickets'' in South Africa and ''wētā'' i ...
- wētā (except cave wētā), king crickets ***Family
Cooloolidae ''Cooloola'' is a genus of ensiferan orthopterans known as Cooloola monsters. It is the only genus in the subfamily Cooloolinae and family Cooloolidae of the superfamily Stenopelmatoidea. Four species are known from this family, all endemic to ...
- Cooloola monsters ***Family
Gryllacrididae Gryllacrididae are a family of non-jumping insects in the suborder Ensifera occurring worldwide, known commonly as leaf-rolling crickets or raspy crickets. The family historically has been broadly defined to include what are presently several oth ...
- leaf-rolling crickets ***Family Stenopelmatidae - Jerusalem crickets **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 ...
***Family † Haglotettigoniidae ***Family
Tettigoniidae 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 ...
- bush crickets, katydids, koringkrieks **Superfamily ''Incertae sedis'' ***Family ''incertae sedis'' ****Genus †'' Tettoraptor'' *Infraorder ''incertae sedis'' **Superfamily † Gryllavoidea ***Family † Gryllavidae *Superfamily ''Incertae sedis'' ** Family † Palaeorehniidae (syn " Zeuneropterinae") **Family † Vitimiidae


Phylogeny

The phylogenetic relationships of the Ensifera, summarized by Darryl Gwynne in 1995 from his own work and that of earlier authors, are shown in the following
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 d ...
, with the Orthoptera divided into two main groups, Ensifera and
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). Fossil Ensifera are found from the late
Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...
period onwards. The oldest known fossil in the
Archaeorthoptera 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, n ...
, the
crown group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...
of the Orthoptera, and also the oldest member of the
Pterygota The Pterygota ( grc, πτερυγωτός, pterugōtós, winged) are a subclass of insects that includes the winged insects. It also includes insect orders that are secondarily wingless (that is, insect groups whose ancestors once had wings b ...
(winged insects), is from the
Namurian The Namurian is a stage in the regional stratigraphy of northwest Europe with an age between roughly 326 and 313 Ma (million years ago). It is a subdivision of the Carboniferous system or period and the regional Silesian series. The Namurian is ...
(324 mya) Lower Carboniferous beds in the Upper Silesian Basin of the Czech Republic.


Notes


References


External links


The Orthopterists' Society
{{Authority control Insect suborders Cisuralian first appearances Extant Permian first appearances