Formicium Temporal range: Lutetian, | |
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Formicium mirabile type illustration | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Hymenoptera |
Family: | Formicidae |
Subfamily: | † Formiciinae |
Genus: | † Formicium Westwood, 1854 |
Type species | |
Formicium brodiei Westwood, 1854 | |
Other species | |
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Synonyms | |
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Formicium is an extinct collective genus of giant ants in the Formicidae subfamily Formiciinae. The genus currently contains three species, Formicium berryi, Formicium brodiei, and Formicium mirabile. All three species were described from Eocene aged sediments. [1]
The collective genus Formicium was first established by English entomologist and archaeologist John O. Westwood in 1854. [2] and originally was only described from isolated fossil forewings, with full queens, drones, and workers being described from Germany later. From 1854 until 2010, the genus was expanded to include five species, however the two German species were subsequently removed and placed in the related genus Titanomyrma as T. giganteum and T. simillimum. The species Formicium mirabile, named by Theodore D. A. Cockerell in 1920, and Formicium brodiei, named by Westwood in 1854, are both known from fore-wings recovered from middle Eocene Bagshot Formation of Bournemouth, Dorset, England. [1] The third species named, Formicium berryi was described by Frank M. Carpenter in 1929 from the middle Eocene Claiborne Formation in Puryear, Tennessee, USA, though he misidentified the formation as the Wilcox Formation. F. berryi was the first described occurrence of the genus and, until 2011, the subfamily, in North America. [1]
As the wing structure of Formicidae is very plastic and can vary greatly, even within a species and size between males and females can be notably different, the description of fossil species from wings alone is problematic. With the removal of the two German species described from full body fossils in 2011, Dr. Bruce Archibald and coauthors changed Formicium from a nominal genus to collective genus. They suggested it be used to contain species described from wings which do not have enough detail to place into a nominal genus such as Titanomyrma. As a collective genus, it does not contain a type species per the International Code of Zoological Nomenclature, but is still retained as the type genus for the subfamily Formiciinae. [1]
Formicium berryi was originally described as Eoponera berryi by Frank Carpenter and placed in the extant subfamily Ponerinae. This was based on the idea that the new species was related to the modern genus Dinoponera . [3] When initially described by Theodore D. A. Cockerell, Formicium mirabilis was placed in the monotypic genus Megapterites. At that time he considered the species to be part of the family Pseudosiricidae. This placement was retained in the Treatise on Invertebrate Paleontology Hymenoptera section written by Frank Carpenter. [4] This placement, however did not reflect the changes made by German paleoentomologist Herbert Lutz who synonymized Eoponera into Formicium in 1986 while describing the subfamily Formiciinae and the two German species. His 1990 synonymy of Megapterites into Formicium was also not reflected in the Treatise. Currently both genus names, Megapterites and Eoponera are accepted as junior synonyms of Formicium. [1] [4]
F. berryi is only known from a forewing 26 millimetres (1.0 in) long and 7 millimetres (0.28 in) wide. It was collected by professor E.W. Berry of the Johns Hopkins University. Owing to the wings size, Carpenter believed that the ant may have been 57 millimetres (2.2 in) long, making it one of the largest ants to ever live. [5] It has a long and narrow stigma (small, colored thick area near the wing-tip), and the discoidal cell is triangular. The apex is absent on the wing, but a complete shape of the wing may resemble that of Myrmecia . The wings have similar dimensions to Camponotus gigas , a giant ant found in Sumatra and the Malay Peninsula. [5]
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The F. mirabile holotype is an incomplete forewing of 45 mm (1.8 in) preserved length, and at least a 50 mm (2.0 in) estimated length in life. The specimen and its counterpart were collected by J. S. Gardner and added to the British Museum paleontology collections as specimen "I.2596".
Formiciinae is an extinct subfamily of ants known from Eocene deposits in Europe and North America.
Sphecomyrma is an extinct genus of ants which existed in the Cretaceous approximately 79 to 92 million years ago. The first specimens were collected in 1966, found embedded in amber which had been exposed in the cliffs of Cliffwood, New Jersey, by Edmund Frey and his wife. In 1967, zoologists E. O. Wilson, Frank Carpenter and William L. Brown, Jr. published a paper describing and naming Sphecomyrma freyi. They described an ant with a mosaic of features—a mix of characteristics from modern ants and aculeate wasps. It possessed a metapleural gland, a feature unique to ants. Furthermore, it was wingless and had a petiole which was ant-like in form. The mandibles were short and wasp-like with only two teeth, the gaster was constricted, and the middle and hind legs had double tibial spurs. The antennae were, in form, midway between the wasps and ants, having a short first segment but a long flexible funiculus. Three additional species, S. canadensis, S. mesaki and S. nexa, were described in 1985, 2005, and 2024, respectively.
Prionomyrmex is an extinct genus of bulldog ants in the subfamily Myrmeciinae of the family Formicidae. It was first described by Gustav Mayr in 1868, after he collected a holotype worker of P. longiceps in Baltic amber. Three species are currently described, characterised by their long mandibles, slender bodies and large size. These ants are known from the Eocene and Late Oligocene, with fossil specimens only found around Europe. It is suggested that these ants preferred to live in jungles, with one species assumed to be an arboreal nesting species. These ants had a powerful stinger that was used to subdue prey. In 2000, it was suggested by Cesare Baroni Urbani that the living species Nothomyrmecia macrops and a species he described both belonged to Prionomyrmex, but this proposal has not been widely accepted by the entomological community. Instead, scientists still classify the two genera distinctive from each other, making Nothomyrmecia a valid genus.
Archiinocellia is an extinct genus of snakefly in the family Raphidiidae known from Eocene fossils found in western North America. The genus contains two species, the older Archiinocellia oligoneura and the younger Archiinocellia protomaculata. The type species is of Ypresian age and from the Horsefly Shales of British Columbia, while the younger species from the Lutetian Green River Formation in Colorado. Archiinocellia protomaculata was first described as Agulla protomaculata, and later moved to Archiinocellia.
Palaeovespa is an extinct genus of wasp in the Vespidae subfamily Vespinae. The genus currently contains eight species: five from the Priabonian stage Florissant Formation in Colorado, United States, two from the middle Eocene Baltic amber deposits of Europe, and one species from the late Paleocene of France.
Titanomyrma is a genus of extinct giant ants which lived during the Eocene. The type species Titanomyrma gigantea and the smaller Titanomyrma simillima are known from the Eocene of Germany, while the third species Titanomyrma lubei, is known from Wyoming, United States. The presence of Titanomyrma in North America was considered to indicate "the first reported cross-Arctic dispersal by a thermophilic insect group". However a queen reported from Upland temperate shales in British Columbia raised questions on the exact thermophilic nature of the genus. The type species of this genus, T. gigantea, is the largest-known fossil or extant species of ant in the world.
Paleolepidopterites is a collective genus of fossil moths which can not be placed in any defined family. The included species were formerly placed in the leaf-roller family Tortricidae and are known from fossils found in Russia and the United States. The collective genus contains three species: Paleolepidopterites destructus, Paleolepidopterites florissantanus, and Paleolepidopterites sadilenkoi, formerly placed within the genera Tortrix and Tortricites respectively. The three species were formally redescribed and moved to the new collective genus by Heikkilä et al. (2018).
Ypresiomyrma is an extinct genus of ants in the subfamily Myrmeciinae that was described in 2006. There are four species described; one species is from the Isle of Fur in Denmark, two are from the McAbee Fossil Beds in British Columbia, Canada, and the fourth from the Bol’shaya Svetlovodnaya fossil site in Russia. The queens of this genus are large, the mandibles are elongated and the eyes are well developed; a stinger is also present. The behaviour of these ants would have been similar to that of extant Myrmeciinae ants, such as solitary foraging for arthropod prey and never leaving pheromone trails. The alates were poor flyers due to their size, and birds and animals most likely preyed on these ants. Ypresiomyrma is not assigned to any tribe, and is instead generally regarded as incertae sedis within Myrmeciinae. However, some authors believe Ypresiomyrma should be assigned as incertae sedis within Formicidae.
Avitomyrmex is an extinct genus of bulldog ants in the subfamily Myrmeciinae which contains three described species. The genus was described in 2006 from Ypresian stage deposits of British Columbia, Canada. Almost all the specimens collected are queens, with an exception of a single fossilised worker. These ants are large, and the eyes are also large and well-developed; a sting is present in one species. The behaviour of these ants may have been similar to extant Myrmeciinae ants, such as foraging solitarily for arthropod prey and never leaving pheromone trails to food sources. Avitomyrmex has not been assigned to any tribe, instead generally being regarded as incertae sedis within Myrmeciinae. However, its identity as an ant has been challenged, although it is undoubtedly a hymenopteran insect.
Macabeemyrma is an extinct genus of bulldog ants in the subfamily Myrmeciinae containing the single species Macabeemyrma ovata, described in 2006 from Ypresian stage deposits of British Columbia, Canada. Only a single specimen is known; a holotype queen found preserved as a compression fossil. The specimen had no wings and small portions of its legs and eyes were faintly preserved. It was a large ant, reaching 25 millimetres (0.98 in) in length. This ants' behaviour would have been similar to that of extant Myrmeciinae ants, such as foraging singly in search for arthropod prey and nesting in soil or in trees. Macabeemyrma shows similarities to extinct ants in the genus Ypresiomyrma, and to the living Nothomyrmecia macrops, but has not been conclusively assigned to any tribe, instead generally regarded as incertae sedis within Myrmeciinae. However, the sole specimen lacks definitive traits, and its classification in Myrmeciinae, and even its identity as an ant, has been challenged.
Myrmeciites is an extinct form genus of bulldog ants in the subfamily Myrmeciinae of the family Formicidae, which contains three described species and two fossils not placed beyond the genus level. Described in 2006 from Ypresian stage deposits, all three of the described species and one unplaced fossil are from British Columbia, Canada, while the second unplaced fossil is from Washington State, USA. These ants were large, with the largest specimens collected reaching 3 centimetres (1.2 in). The behaviour of these ants would have been similar to extant Myrmeciinae ants, such as solitary foraging, nesting either in the soil or trees, and leaving no pheromone trail to food sources. Due to the poor preservation of these ants, their phylogenetic position among Myrmeciinae is unclear, and no type species has been designated. These ants are classified as incertae sedis in Myrmeciinae, but some writers have classified it as incertae sedis within the insect order Hymenoptera. This reclassification, however, has not been accepted; instead, Myrmeciites remains in Myrmeciinae.
Protostephanus is an extinct genus of crown wasp in the Hymenoptera family Stephanidae known from an Eocene fossil found in the United States of America. The genus contains a single described species, Protostephanus ashmeadi placed in the stephanid subfamily Stephaninae.
Prionomyrmecini is an ant tribe belonging to the subfamily Myrmeciinae established by William Morton Wheeler in 1915. Two members are a part of this tribe, the extant Nothomyrmecia and the extinct Prionomyrmex. The tribe was once considered a subfamily due to the similarities between Nothomyrmecia and Prionomyrmex, but such reclassification was not widely accepted by the scientific community. These ants can be identified by their long slender bodies, powerful stingers and elongated mandibles. Fossil Prionomyrmecini ants were once found throughout Europe, possibly nesting in trees and preferring jungle habitats. Today, Prionomyrmecini is only found in Australia, preferring old-growth mallee woodland surrounded by Eucalyptus trees. Nothomyrmecia workers feed on nectar and arthropods, using their compound eyes for prey and navigational purposes. Owing to their primitive nature, they do not recruit others to food sources or create pheromone trails. Nothomyrmecia colonies are small, consisting of 50 to 100 individuals.
Eulithomyrmex is an extinct genus of ant in the formicid subfamily Agroecomyrmecinae. The genus contains two described species, Eulithomyrmex rugosus and Eulithomyrmex striatus. Eulithomyrmex is known from a group of Late Eocene fossils which were found in North America.
Burmomyrma is an extinct genus of aculeate hymenopteran, suggested to be an ant. The genus contains a single described species, Burmomyrma rossi. Burmomyrma is known from a single Middle Cretaceous fossil which was found in Asia.
Aphaenogaster donisthorpei is an extinct species of ant in formicid subfamily Myrmicinae known from a Late Eocene fossil from North America. A. donisthorpei was one of two Aphaenogaster species described in the 1930 paper.
Aphaenogaster longaeva is an extinct species of ant in formicid subfamily Myrmicinae known from a solitary Eocene or Oligocene fossil found in North America. A. longaeva was one of five insect species described by the paleoentomologist Samuel Hubbard Scudder in an 1877 paper.
Archimyrmex is an extinct genus of ant in the formicid subfamily Myrmeciinae, described by palaeoentomologist Theodore Cockerell in 1923. The genus contains four described species, Archimyrmex rostratus, Archimyrmex piatnitzkyi, Archimyrmex smekali and Archimyrmex wedmannae. Archimyrmex is known from a group of Middle Eocene fossils which were found in North America, South America, and Europe. The genus was initially placed in the subfamily Ponerinae, but it was later placed in Myrmeciinae; it is now believed to be the ancestor of the extant primitive genus Myrmecia from Australia. Despite this, Archimyrmex is not a member to any tribe and is regarded as incertae sedis within Myrmeciinae. However, some authors believe Archimyrmex should be assigned as incertae sedis within Formicidae. These ants can be characterised by their large mandibles and body length, ranging from 13.2 to 30 mm. They also have long, thin legs and an elongated mesosoma (thorax) and petiole.
Protopone is an extinct genus of ants in the formicid subfamily Ponerinae described from fossils found in Europe and Asia. There are seven described species placed into the genus, Protopone? dubia, Protopone germanica, Protopone magna, Protopone oculata, Protopone primigena, Protopone sepulta, and Protopone vetula. Protopone is one several Lutetian Ponerinae genera.
Aneuretellus is an extinct genus of ant in the formicid subfamily Aneuretinae, and is one of eight genera of the subfamily. The genus contains a single described species Aneuretellus deformis and is known from one Middle Eocene fossil which was found in Sakhalin in the Russian Far East.