Myrmeciites

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Myrmeciites
Temporal range: Ypresian, 51  Ma
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Myrmeciites SR 05-03-01 01.jpg
Myrmeciites " incertae sedis " SR 05-03-01
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Myrmeciinae
Tribe: incertae sedis
Genus:† Myrmeciites
Archibald, Cover, & Moreau, 2006 [1]
Species
  • see text

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 (Early Eocene) 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.

Contents

History and classification

Fossils of Myrmeciites were first studied and described by Bruce Archibald, Stefan Cover and Corrie Moreau of the Museum of Comparative Zoology in Cambridge, Massachusetts. They published their 2006 description of the form genus (which is a collection of species formally described but cannot be identified to genus level) in an Annals of the Entomological Society of America journal article. [2] The genus name is a combination of the ant subfamily name "Myrmeciinae" and the Latin suffix "ites", meaning "having the nature of", which is commonly used in the naming of fossil taxa. [3] Its phylogenetic position within Myrmeciinae is unclear, due to the incomplete and poor preservation of the collected specimens. [4] The paper that described the form genus included the description of three species, including Myrmeciites herculeanus, Myrmeciites(?) goliath, and Myrmeciites(?) tabanifluviensis. [2]

Archibald and colleagues classified Myrmeciites as incertae sedis (Latin for "of uncertain placement") within the ant subfamily Myrmeciinae, as the specimens are too poorly preserved to be assigned to any tribe. [5] However, in a 2008 paper, Cesare Baroni Urbani of the University of Basel, Switzerland, classified Myrmeciites as incertae sedis within the insect order Hymenoptera (which comprises the sawflies, wasps, bees and ants) because the critical characters used to identify Myrmeciinae ants, or even the family Formicidae cannot be carried out on Myrmeciites. [6] Despite these comments, a 2012 report by Russian palaeoentomologist Gennady M. Dlussky does not make any comment in regard to Baroni Urbani's views; instead, he accepts the classification of Archibald and colleagues. [7]

Description

Archibald, Cover, and Moreau erected the form genus as an encompassing category for all fossil ants which, while belonging to the subfamily Myrmeciinae, lack details needed for placement in other described genera. [4] This may be due to preservation quality or positioning of the individual ant resulting in obscured details. As Myrmeciites is a form genus it does not have a designated type species per the International Code of Zoological Nomenclature. [4]

M. herculeanus

M. herculeanus was described from a single side of a compression fossil found at the Middle Ypresian McAbee Fossil Beds, Kamloops Group, near Cache Creek, British Columbia. The incomplete specimen numbered UCCIPR L-18 F-974, is currently preserved in the paleontology collections housed at the Thompson Rivers University, Kamloops, British Columbia. Archibald, Cover, and Moreau coined the specific epithet "herculeanus" from the Latin name "Hercules" in reference to the notably sturdy and large morphology of the type specimen, and after the divine hero Heracles in Greek mythology, who was the son of Zeus and Alcmene. [8] The species is discernible from the other species of Myrmeciinae by its notably larger size, the worker caste ant is estimated at over 20 millimetres (0.79 in) in life. Its size is comparable to ants in the genus Ypresiomyrma . The shape of the petiole, a narrow waist that is located between the mesosoma and gaster is distinct from other species and is similar in structure to the genus Prionomyrmex though the shape and size of the mandibles are distinct. Overall, M. herculeanus was assigned to Myrmeciinae due to its mandible length, despite them being poorly preserved, and due to the appearance of its petiole and propodeum. The legs are long in comparison to its body length, and the gaster is robust. It is unknown if the species had a sting, due to the preservation of the specimen. The size of the single known worker is larger than any of the studied queens; as Myrmeciinae queens are slightly larger than the other castes in a species, this indicates M. herculeanus is most likely a distinct species. Due to the incomplete nature of the type specimen, the species was placed in Myrmeciites. [8]

M.(?) goliath

The second species described from the McAbee Fossil Beds is M.(?) goliath and as with M. herculeanus, it is known from one specimen, though both the part and counterpart are known for M.(?) goliath. The holotype is housed in the Thompson Rivers University, Kamloops, collections as UCCIPR L-18 F-999 and UCCIPRL-18 F-1000 for each side respectively. [9] The species name "goliath" was chosen by Archibald, Cover and Moreau in reference to the mythological Goliath, due to the notable size of the ant. [9] The holotype specimen is a partial worker or queen which is incomplete, with a notable portion of the gaster missing. The species can be separated from other ants by the large size, the holotype approaching 3 centimetres (1.2 in). The only other Eocene ants in this size range are members of the genus Titanomyrma , that were formerly placed in Formicium . [9] [10] M.(?) goliath is distinguishable from Titanomyrma by the shape and structure of the antennae and length of the legs, which are notably shorter in Titanomyrma. The head is very round and flattened, compound eyes are present and the mandibles are small, less than half the length of the head. The head is also small in proportion to the mesosoma. The legs are large and long, and a poorly preserved petiole is known. [9] Most of the specimen is preserved in fine detail, but some characters are so poorly preserved that the exact placement of M.(?) goliath within Myrmeciinae cannot be confirmed. Only future fossils that are better preserved can clarify the position of M.(?) goliath. [11]

M.(?) tabanifluviensis

Unlike the other described Myrmeciites species, M.(?) tabanifluviensis is the only species not from the McAbee Fossil Beds. The holotype, currently deposited in the Courtenay and District Museum paleontology collections as 2003.2.10 CDM 034 was recovered from the "Horsefly shale", part of an unnamed formation, outcropping near the town of Horsefly, British Columbia. The specimen is incomplete with both the head and portions of the mesosoma and gaster missing. The adult had an estimated length in life of over 1.5 centimetres (0.59 in). [12] The right portions of the legs are well preserved while the left legs are missing with two metatibia spurs, one of which is long and the other is short and narrow. Due to the incomplete nature of the specimen and position of the preserved portions, the gender is not identifiable, though the presence of wings indicates it to be a reproductive adult. The small size of the specimen, with a fore-wing length of approximately 12 millimetres (0.47 in), separates this species from other members of the British Columbia Myrmeciinae. The type locality for the species, near the Horsefly River, was the basis for Archibald, Cover and Moreau choosing the specific epithet tabanifluviensis, combined with the horsefly genus name Tabanus , the Latin word fluvius meaning "river" and the sufix -ensis meaning "origin" or "place". [12] The justification of placing this ant in Myrmeciites is due to incomplete preservation. [13]

Unplaced specimens

Two additional specimens were placed within the form genus but not into a specific species. The Courtenay and District Museum specimen 2003.2.9 CDM 03 a&b is a 2 centimetres (20 mm) long partial male recovered from the Falkland site, near Falkland, British Columbia. The generally robust nature of the specimen excludes it from the genus Avitomyrmex , but the details needed for placement between the genera Ypresiomyrma and Macabeemyrma are not present. The Stonerose Interpretive Center specimen "SR 05-03-01" is a possible queen or worker which would have been an estimated 1.5 centimetres (15 mm) long in life. Due to the lateral preservation of the specimen, most characters for placing the specimen into one of the defined species are obscured or absent. The specimen was recovered from outcrops of the Klondike Mountain Formation north west of Republic, Washington. [14]

Ecology

Archibald and colleagues suggested that the behavioural habits for Myrmeciites ants, like that of other extinct Myrmeciinae taxa, may have been similar to its extant relatives. Workers would forage onto trees or vegetation to capture arthropod prey or to feed on nectar, soiling either in the ground or in the trees, making them an arboreal nesting insect. Like other Myrmeciine ants, Myrmeciites most likely did not lay pheromone trails or recruit nestmates to food sources; these ants may have been solitary foragers, relying on their eyes to hunt for prey and for navigational purposes. [15] The nuptial flight of M. (?) tabanifluviensis probably occurred during the late spring or summer, as in extant relatives. [16]

Related Research Articles

Myrmeciinae Subfamily of ants

Myrmeciinae is a subfamily of the Formicidae, ants once found worldwide but now restricted to Australia and New Caledonia. This subfamily is one of several ant subfamilies which possess gamergates, female worker ants which are able to mate and reproduce, thus sustaining the colony after the loss of the queen. The Myrmeciinae subfamily was formerly composed of only one genus, Myrmecia, but the subfamily was redescribed by Ward & Brady in 2003 to include two tribes and four genera: An additional three genera, one form genus, and 9 species were described in 2006 from the Early Eocene of Denmark, Canada, and Washington.

<i>Nothomyrmecia</i> Genus of ants

Nothomyrmecia, also known as the dinosaur ant or dawn ant, is a rare genus of ants consisting of a single species, Nothomyrmecia macrops. These ants live in South Australia, nesting in old-growth mallee woodland and Eucalyptus woodland. The full distribution of Nothomyrmecia has never been assessed, and it is unknown how widespread the species truly is; its potential range may be wider if it does favour old-growth mallee woodland. Possible threats to its survival include habitat destruction and climate change. Nothomyrmecia is most active when it is cold because workers encounter fewer competitors and predators such as Camponotus and Iridomyrmex, and it also increases hunting success. Thus, the increase of temperature may prevent them from foraging and very few areas would be suitable for the ant to live in. As a result, the IUCN lists the ant as Critically Endangered.

<i>Sphecomyrma</i> Extinct genus of ants

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. Two additional species, S. canadensis and S. mesaki, were described in 1985 and 2005, respectively.

<i>Prionomyrmex</i> Extinct genus of ants

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.

<i>Titanomyrma</i> Extinct genus of ants

Titanomyrma is a genus of prehistoric giant ant. The latest species to be discovered, T. lubei, was described in 2011, when a 49.5-million-year-old fossilized winged queen ant, comparable in size to hummingbirds, was found in Wyoming, United States. This fossil is the first fossil of a giant ant found in the Western Hemisphere. The presence of Titanomyrma in North America is considered to indicate "the first reported cross-Arctic dispersal by a thermophilic insect group". Another fossil species of this genus, T. gigantea, is the largest-known fossil or extant species of giant ant in the world.

<i>Ypresiomyrma</i> Extinct genus of ants

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.

<i>Macabeemyrma</i> Extinct genus of ants

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.

Prionomyrmecini Tribe of ants

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.

Sphecomyrminae Extinct subfamily of ants

Sphecomyrminae is an extinct subfamily of ants in family Formicidae known from a series of Cretaceous fossils found in North America, Europe, and Asia. Sphecomyrminae contains eight genera, divided into two tribes Sphecomyrmini and Zigrasimeciini. The tribe Sphecomyrmini contains the six genera Armania, Cretomyrma, Gerontoformica, Orapia, Pseudarmania and Sphecomyrma; while Zigrasimeciini contains Boltonimecia and Zigrasimecia. A number of taxa have been removed from the subfamily and placed either in other subfamilies or are now treated as incertae sedis in Formicidae.

<i>Tatuidris</i> Genus of ants

Tatuidris, or armadillo ant, is a rare genus of ants consisting of a single species, Tatuidris tatusia. The ants are small in size and inhabit the leaf litter of Neotropical forests in Central and South America, from Mexico to Brazil. Workers are ferruginous-colored to dark red and present a distinctive morphology, consisting of a shield-like head with a broad vertex, ventrally-turned heavy mandibles which do not overlap at full closure, and unique among ants – an antenna socket apparatus sitting upside-down. Little is known about the biology of the ants, but they are likely nocturnal and specialist predators.

<i>Brownimecia</i> Cretaceous ant genus described from amber fossils

Brownimecia is an extinct genus of ants, the only genus in the tribe Brownimeciini and subfamily Brownimeciinae of the Formicidae. Fossils of the single identified species, Brownimecia clavata, are known from the Middle Cretaceous of North America. The genus is one of several ants described from Middle Cretaceous ambers of New Jersey. Brownimecia was initially placed in the subfamily Ponerinae, until it was transferred to its own subfamily in 2003; it can be distinguished from other ants due to its unusual sickle-like mandibles and other morphological features that makes this ant unique among the Formicidae. The ant is also small, measuring 3.43 millimetres (0.135 in), and a stinger is present in almost all of the specimens collected. The morphology of the mandibles suggest a high level of feeding specialization.

<i>Burmomyrma</i> Genus of ants

Burmomyrma is an extinct genus of ant-mimic wasp in the extinct family Falsiformicidae. The genus contains a single described species, Burmomyrma rossi. Burmomyrma is known from a single Middle Cretaceous fossil which was found in Asia.

<i>Archimyrmex</i> Extinct genus of ants

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.

<i>Yantaromyrmex</i> Genus of ants

Yantaromyrmex is an extinct genus of ants first described in 2013. Members of this genus are in the subfamily Dolichoderinae of the family Formicidae, known from Middle Eocene to Early Oligocene fossils found in Europe. The genus currently contains five described species, Y. constrictus, Y. geinitzi, Y. intermedius, Y. mayrianum and Y. samlandicus. The first specimens were collected in 1868 and studied by Austrian entomologist Gustav Mayr, who originally placed the fossils in other ant genera until the fossils were reviewed and subsequently placed into their own genus. These ants are small, measuring from 4 to 6 mm in length and can be characterized by their trapezoidal shaped head-capsules and oval compound eyes that are located slightly to the rear of the capsules midpoint, with no known ocelli present.

<i>Myanmyrma</i> Extinct genus of ants

Myanmyrma is an extinct genus of ants not placed into any Formicidae subfamily. Fossils of the single known species, Myanmyrma gracilis, are known from the Middle Cretaceous of Asia. The genus is one of several ants described from Middle Cretaceous ambers of Myanmar.

<i>Camelomecia</i> Extinct genus of ants

Camelomecia is an extinct genus of stem-group ants not placed into any Formicidae subfamily. Fossils of the single known species, Camelomecia janovitzi, are known from the Middle Cretaceous of Asia. The genus is one of several ants described from Middle Cretaceous ambers of Myanmar.

<i>Pachycondyla succinea</i> Extinct species of ant

Pachycondyla succinea is an extinct species of ant in the formicid subfamily Ponerinae described from fossils found in Europe. P. petrosa is one of three middle Eocene Pachycondyla species found in Baltic amber.

Klondikia is an extinct hymenopteran genus in the ant family Formicidae with a single described species Klondikia whiteae. The species is solely known from the Early Eocene sediments exposed in northeast Washington state, United States. The genus is currently not placed into any ant subfamily, being treated as incertae sedis.

References

  1. ↑ Johnson, Norman F. (19 December 2007). "Myrmeciites Archibald, Cover & Moreau". Hymenoptera Name Server version 1.5. Columbus, Ohio, USA: Ohio State University . Retrieved 1 April 2015.
  2. 1 2 Archibald, Cover & Moreau 2006, p. 487.
  3. ↑ Archibald, Cover & Moreau 2006, p. 500.
  4. 1 2 3 Archibald, Cover & Moreau 2006, p. 490.
  5. ↑ AntWeb. "Genus: †Myrmeciites Archibald, Cover & Moreau, 2006". The California Academy of Sciences. Retrieved 16 August 2015.
  6. ↑ Baroni Urbani, Cesare (2008). "Orthotaxonomy and parataxonomy of true and presumed bulldog ants (Hymenoptera, Formicidae)" (PDF). Doriana (Suppl. To Annali del Museo Civico di Storia Naturale Giacomo Doria). 8 (358): 1–10. ISSN   0417-9927.
  7. ↑ Dlussky, G. M. (May 2012). "New fossil ants of the subfamily Myrmeciinae (Hymenoptera, Formicidae) from Germany" (PDF). Paleontological Journal. 46 (3): 288–292. doi:10.1134/S0031030111050054. S2CID   83891156.
  8. 1 2 Archibald, Cover & Moreau 2006, pp. 500–501.
  9. 1 2 3 4 Archibald, Cover & Moreau 2006, p. 502.
  10. ↑ Archibald, S. Bruce; Johnson, Kirk R.; Mathewes, Rolf W.; Greenwood, David R. (2011). "Intercontinental dispersal of giant thermophilic ants across the Arctic during early Eocene hyperthermals". Proceedings of the Royal Society B . 278 (1725): 3679–86. doi:10.1098/rspb.2011.0729. PMC   3203508 . PMID   21543354.
  11. ↑ Archibald, Cover & Moreau 2006, pp. 508–509.
  12. 1 2 Archibald, Cover & Moreau 2006, pp. 502–503.
  13. ↑ Archibald, Cover & Moreau 2006, p. 509.
  14. ↑ Archibald, Cover & Moreau 2006, pp. 501–502.
  15. ↑ Archibald, Cover & Moreau 2006, p. 513.
  16. ↑ Archibald, Cover & Moreau 2006, p. 505.

Cited text

Archibald, S.B.; Cover, S. P.; Moreau, C. S. (2006). "Bulldog Ants of the Eocene Okanagan Highlands and History of the Subfamily (Hymenoptera: Formicidae: Myrmeciinae)" (PDF). Annals of the Entomological Society of America. 99 (3): 487–523. doi: 10.1603/0013-8746(2006)99[487:BAOTEO]2.0.CO;2 .

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