Ensifera
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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 pa ...
s 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 (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 – grassh ...
. 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 ''Carboniferou ...
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 Paleo ...
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 languages, 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 ...
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. Spermatophore ...
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 t ...
. 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, res ...
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 nu ...
, 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 (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, uncertain ...
'' ****Genus †'' Acridiites'' *Infraorder
Gryllidea GryllideaKevan DKM (1982) In Parker d. ''Synopsis and Classification of Living Organisms'' 2: 361. is an infraorder that includes crickets and similar insects in the order Orthoptera. There are two superfamilies, and more than 6,000 described sp ...
**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 ...
***Family †
Baissogryllidae Baissogryllidae is an extinct family of crickets Crickets are orthopteran insects which are related to bush crickets, and, more distantly, to grasshoppers. In older literature, such as Imms,Imms AD, rev. Richards OW & Davies RG (1970) '' ...
***Family Gryllidae - 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 Protogryllidae is an extinct family of crickets Crickets are orthopteran insects which are related to bush crickets, and, more distantly, to grasshoppers. In older literature, such as Imms,Imms AD, rev. Richards OW & Davies RG (1970) ''A ...
***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 Saussure, 1874 – sword-tail crickets References ...
**Superfamily
Gryllotalpoidea The Gryllotalpoidea are a superfamily (taxonomy), superfamily of insects that includes the mole crickets and the ant crickets. The type genus is ''Gryllotalpa''. Recent (2015) molecular phylogenetic studies support the monophyly of the cricket c ...
***Family Gryllotalpidae – 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 Tettigoniidea is an infraorder of the order Orthoptera, with six extant families. Families The ''Orthoptera Species File'' lists: *superfamily Hagloidea Handlirsch, 1906 ** † Eospilopteronidae Cockerell, 1916 ** † Haglidae Handlirsch, 1906 ...
**Superfamily
Hagloidea The superfamily Hagloidea are insects belonging to the order Orthoptera: 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 k ...
- grigs ***Family † Eospilopteronidae ***Family † Haglidae ***Family † Hagloedischiidae ***Family † Prezottophlebiidae ***Family Prophalangopsidae ***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 shorten ...
- camel crickets, cave crickets, cave wētā **Superfamily Schizodactyloidea ***Family Schizodactylidae - dune or splay-footed crickets **Superfamily Stenopelmatoidea ***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'' in ...
- 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 - leaf-rolling crickets ***Family
Stenopelmatidae The family Stenopelmatidae is composed of large, mostly flightless insects resembling crickets (the family Gryllidae). Two genera: '' Ammopelmatus'' and the type genus '' Stenopelmatus'' are found in the New World. '' Oryctopus'' and ''Sia'' are O ...
- Jerusalem crickets **Superfamily Tettigonioidea ***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, ...
- 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 ...
, with the Orthoptera divided into two main groups, Ensifera and Caelifera (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 ''Carboniferou ...
period onwards. The oldest known fossil in the Archaeorthoptera, 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 ...
(winged insects), is from the Namurian (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