Titanoptera Temporal range: | |
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Reconstruction of Gigatitan | |
Scientific classification ![]() | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Superorder: | Orthopterida |
(unranked): | Panorthoptera |
Order: | † Titanoptera |
Families | |
†Mesotitanidae Contents |
Titanoptera is an extinct order of neopteran insects from late Carboniferous to Triassic periods. [1] Titanopterans were very large in comparison with modern insects, some having wingspans of up to 36 centimetres (14 in) or even 40 centimetres (16 in). [2] [3]
Titanopterans are related to modern grasshoppers, but were much larger, had proportionally weaker hindlegs that could not allow the animals to leap, and grasping forelegs and elongated mandibles. Another distinctive feature was the presence of prominent fluted regions on the forewings, which may have been used in stridulation. The general shape and anatomy of the titanopterans suggests that they were predators. [2]
An examination of a fossil of the oldest titanopteran genus, Theiatitan, seems to indicate that titanopterans did not utilize stridulation (unlike modern orthopterans), but rather used flashes of light from wing displays and crepitation, moving their wings to produce sound. The authors argue that stridulation, crepitation, castanet signaling or light flash alone do not fully explains the diversity of structures observed in Titanoptera, and note that both sexes seem to have the fluted region on the forewing. Theiatian is 50 Ma older than the previous oldest species of Titanoptera, and thus Theiatitan would be the oldest known insect with a wing structure specialized for communication. [1]
Some titanopterans may have been able to only glide, not fly, such as Gigatitan vulgaris. The hind wing area of it is almost the same as that of Pseudophyllanax imperialis, one of the largest modern Orthoptera, and a poor flier, but Gigatitan is larger in volume. All known hind wings of Titanoptera, whatever their sizes, have quite reduced vannus, while most extant flying Orthoptera have large ones. [1]
Other than Theiatitan, reliable records of titanopterans are known from Kyrgyzstan, Australia and South Korea. Considering some possible records from Russia as well, titanopterans possibly had a circum-Tethys distribution. [3]
There is controversy regarding the classification of Titanoptera. Titanoptera was previously thought to be related to Geraridae (including Gerarus ), but it is no longer supported. [4] Olivier (2007) considered that genera in Titanoptera should be included in Orthoptera, and divided from extinct orthopteran family Tchomanvissidae. [5] But later study considered that the relationships between Titanoptera and Tcholmanvissiidae remain controversial. [1] Three genera known from Permian, Permotitan , Deinotitan , Monstrotitan possibly not belong to Titanoptera. [1] Although the genus Jubilaeus originally belonged to Mesotitanidae, but it is later considered to belong to Tcholmanvissiidae. [5] [6] Steinhardtia was originally attributed to Titanoptera, but as fossil does not show the venational structures of the order Titanoptera, and it is even possible to be misidentification of plant fossil, possibly fern. [1]
Order Titanoptera
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 – grasshoppers, locusts, and close relatives; and Ensifera – crickets and close relatives.
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 make up the order Orthoptera. Ensifera is believed to be a more ancient group than Caelifera, with its origins in the Carboniferous period, the split having occurred at the end of the Permian period. Unlike the Caelifera, the Ensifera contain numerous members that are partially carnivorous, feeding on other insects, as well as plants.
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). The latter should not be confused with the mole crickets (Gryllotalpidae), which belong to the other Orthopteran sub-order Ensifera.
Michael S. Engel, FLS, FRES is an American paleontologist and entomologist, notable for contributions to insect evolutionary biology and classification. In connection with his studies he has undertaken field expeditions in Central Asia, Asia Minor, the Levant, Arabia, eastern Africa, the high Arctic, and South and North America, and has published more than 983 papers in scientific journals and over 925 new living and fossil species. Some of Engel's research images were included in exhibitions on the aesthetic value of scientific imagery.
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Petrolacosaurus is an extinct genus of diapsid reptile from the late Carboniferous period. It was a small, 40-centimetre (16 in) long reptile, and one of the earliest known reptile with two temporal fenestrae. This means that it was at the base of Diapsida, the largest and most successful radiation of reptiles that would eventually include all modern reptile groups, as well as dinosaurs and other famous extinct reptiles such as plesiosaurs, ichthyosaurs, and pterosaurs. However, Petrolacosaurus itself was part of Araeoscelida, a short-lived early branch of the diapsid family tree which went extinct in the mid-Permian.
Insect wings are adult outgrowths of the insect exoskeleton that enable insects to fly. They are found on the second and third thoracic segments, and the two pairs are often referred to as the forewings and hindwings, respectively, though a few insects lack hindwings, even rudiments. The wings are strengthened by a number of longitudinal veins, which often have cross-connections that form closed "cells" in the membrane. The patterns resulting from the fusion and cross-connection of the wing veins are often diagnostic for different evolutionary lineages and can be used for identification to the family or even genus level in many orders of insects.
Glosselytrodea is an extinct order of insects, containing about thirty species. Its fossil record dates from the Permian to the Upper Jurassic, and is distributed across Eurasia, the Americas, and Australia. Its classification is uncertain, but may be closely related to Neuropterida or Orthoptera.
Protophasma is an extinct genus of Protorthopteran insect from the Carboniferous of Europe and North America.
Cyclida is an extinct order of crab-like fossil arthropods that lived from the Carboniferous to the Jurassic and possibly Cretaceous. Their classification is uncertain, but they are generally interpreted as crustaceans, likely belonging to the superclass Multicrustacea.
Paleontology in Rhode Island refers to paleontological research occurring within or conducted by people from the U.S. state of Rhode Island. Rhode Island has a relatively sparse fossil record. Among its more common fossils are plant remains that are frequently associated with the state's coal beds. During the early Paleozoic, Rhode Island was at least partially submerged under a sea inhabited by trilobites. During the Carboniferous period the state became a swampy environment where lush vegetation included trees more than 50 feet high. Local animal life included arachnids and insects like cockroaches. Rift basins formed locally during the Permian. The ensuing Triassic and Jurassic periods are absent from the state's rock record. Little is known about the state's Cretaceous history. The Paleogene and Neogene periods are also missing from Rhode Island's rock record. During the Pleistocene the state was subjected to glacial activity. Notable local fossil finds have included previously unknown kinds of insect and abundant ancient amphibian trackways.
The cohort Polyneoptera is a proposed taxonomic ranking for the Orthoptera and all other Neopteran insects believed to be more closely related to Orthoptera than to any other insect orders. These winged insects, now in the Paraneoptera, were formerly grouped as the Hemimetabola or Exopterygota on the grounds that they have no metamorphosis, the wings gradually developing externally throughout the nymphal stages.
Adelophthalmidae is a family of eurypterids, an extinct group of aquatic arthropods. Adelophthalmidae is the only family classified as part of the superfamily Adelophthalmoidea, which in turn is classified within the infraorder Diploperculata in the suborder Eurypterina.
Cyphoderris strepitans, the sagebrush cricket or sagebrush grig, is a one of only a few surviving species in the family Prophalangopsidae. Three of these species are in the genus Cyphoderris and all three are endemic to North America. C. strepitans name is from the Latin word 'strepitans' which means 'making a great noise', refers to their calling song during the mating season.
Gigatitan is an extinct genus of titanopteran insect that lived in Kyrgyzstan during the Triassic period. The type species is G. vulgaris, described by Aleksandr Grigorevich Sharov in 1968. Fossils of Gigatitan have been found in the Madygen Formation. It is the type genus of the family Gigatitanidae, in which the closely related Nanotitan and Ootitan are also included.
Cretophasmomima is an extinct genus of stem group-stick insect from the Cretaceous of Eurasia and is one of the oldest and most basal stick insects known, it belongs to the Susumanioidea. The type species, Cretophasmomima vitimica, was described in 1985 from the Aptian aged Zaza Formation in Russia. A second species, Cretophasmomima burjatica was described from the same formation in 1988. A third and fourth species, Cretophasmomima clara and Cretophasmomima arkagalica were also described in 1988 in the same paper from the Ola Formation and the Arkagalinskaya Formation respectively, both formations are Lower Campanian in age. A fifth species Cretophasmomima melanogramma was described in 2014 off the basis of three specimens from the Aptian Yixian Formation in China. In 2020 a sixth species Cretophasmomima traceyae was described based on a forewing with preserved colouration from the Barremian aged Weald Clay Formation in England.
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Clatrotitan is an extinct genus of titanopteran insect, known from the Triassic of Australia. It is originally described from a species, C. andersoni, then later study considered that Mesotitan scullyi as species of Clatrotitan too. But another study synonymized Mesotitan and Clatrotitan. A study in 2021 proposed to keep the two genera Clatrotitan and Mesotitan separated. C. andersoni had a large forewing, which was 13.8 centimetres (5.4 in) long.
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