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''Titanomyrma'' is a genus of prehistoric giant
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of ...
. The latest species to be discovered, ''T. lubei'', was described in 2011, when a 49.5-million-year-old
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
ized winged
queen ant A queen ant (formally known as a gyne) is an adult, reproducing female ant in an ant colony; generally she will be the mother of all the other ants in that colony. Some female ants, such as the ''Cataglyphis'', do not need to mate to produce offs ...
, comparable in size to hummingbirds, was found in
Wyoming Wyoming () is a state in the Mountain West subregion of the Western United States. It is bordered by Montana to the north and northwest, South Dakota and Nebraska to the east, Idaho to the west, Utah to the southwest, and Colorado to the s ...
,
United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territori ...
. 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 A thermophile is an organism—a type of extremophile—that thrives at relatively high temperatures, between . Many thermophiles are archaea, though they can be bacteria or fungi. Thermophilic eubacteria are suggested to have been among the earl ...
insect group". Another fossil species of this genus, ''T. gigantea'', is the largest-known fossil or extant species of giant ant in the world.


Discovery

The fossil ant ''Titanomyrma lubei'' was discovered in ancient lake sediments in the DMNS locality 784, Farson Fish Beds, Laney Member of the
Green River Formation The Green River Formation is an Eocene geologic formation that records the sedimentation in a group of intermountain lakes in three basins along the present-day Green River in Colorado, Wyoming, and Utah. The sediments are deposited in very fine ...
; Sweetwater County,
Wyoming Wyoming () is a state in the Mountain West subregion of the Western United States. It is bordered by Montana to the north and northwest, South Dakota and Nebraska to the east, Idaho to the west, Utah to the southwest, and Colorado to the s ...
, USA by Louis Lube, the collector and donor of the
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of sever ...
specimen. The fossil is part of the collection of
Denver Museum of Nature and Science The Denver Museum of Nature & Science is a municipal natural history and science museum in Denver, Colorado. It is a resource for informal science education in the Rocky Mountain region. A variety of exhibitions, programs, and activities help mus ...
and was discovered by Bruce Archibald and co-author and museum chief curator Kirk R. Johnson while casually going through storage drawers. The fossil depicts the queen ant only and no fossils of workers have been found. With the discovery of ''Titanomyrma lubei'' and the description of the genus, two other species of giant ants, ''Formicium giganteum'' and ''Formicium simillima'', were reclassified into this genus. Well preserved fossils of ''T. gigantea'' and ''T. simillima'' were discovered in the
Messel shales The Messel pit (german: Grube Messel) is a disused quarry near the village of Messel ( Landkreis Darmstadt-Dieburg, Hesse) about southeast of Frankfurt am Main, Germany. Bituminous shale was mined there. Because of its abundance of well-preser ...
, near the village of Messel, in the state of
Hessen Hesse (, , ) or Hessia (, ; german: Hessen ), officially the State of Hessen (german: links=no, Land Hessen), is a state in Germany. Its capital city is Wiesbaden, and the largest urban area is Frankfurt. Two other major historic cities are Darms ...
, 30 km south of
Frankfurt am Main Frankfurt, officially Frankfurt am Main (; Hessian: , "Frank ford on the Main"), is the most populous city in the German state of Hesse. Its 791,000 inhabitants as of 2022 make it the fifth-most populous city in Germany. Located on its na ...
(Darmstadt Administrative District), in
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
. A related undescribed species has been found in the nearby Eckfeld Maar.


Etymology

The name of the genus is a derivative of the Greek ('), meaning 'one of prodigious size, strength, or achievement', and alluding to the Titans of Greek mythology; and the Greek word (') meaning 'ant', gender feminine. The specific epithet of the new species is formed from the surname of the collector of the holotype, Louis Lube.


Taxonomy

Archibald ''et al.'' in 2011 erected the genus ''Titanomyrma'', described the species ''Titanomyrma lubei'' and proposed two new combinations, ''T. gigantea'' (formerly ''Formicium giganteum'' Lutz, 1986) and ''T. simillima'' (formerly ''F. simillimum'' Lutz, 1986). ''T. gigantea'' has been designated the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specime ...
for the genus. The genus ''Titanomyrma'' is differentiated from others in the family by the shape of the gaster which is variable. In the three included species the gaster ranges from ovate to more slender or cylindrical. The A5 abdominal segment width relative to other gaster segments is variable and the relative lengths of A3–A7 are also variable. The queens of the three ''Titanomyrma'' can be distinguished from those of other genera most easily by gaster characters. The gasters are more slender in ''Titanomyrma''. Amongst the three species of ''Titanomyrma'', the ratio between length to width of the queens of the three species is as follows, ''T. lubei'' – 2.14, ''T. gigantea'' – 1.40, and ''T. simillima'' – 1.50. The middle half of ''T. lubei'' is roughly cylindrical while for other ''Titanomyrma'' species, it is ovate with the segment A5 being the widest. The segment A3 has a length about a quarter of the width while for other species it is about a third. The segment A4 is thrice as long as the length of segment A3 while that of A4 segments of other species is less than twice. The segment A5–6 of ''T. lubei'' is approximately half as long as it is wide while for other species it is about a third. A3 is not curved around the petiole at the junction. ''T. gigantea'' is the largest giant ant ever found, larger than the biggest extant giant ants, which are the driver ants of the genus ''
Dorylus ''Dorylus'', also known as driver ants, safari ants, or siafu, is a large genus of army ants found primarily in central and east Africa, although the range also extends to southern Africa and tropical Asia. The term siafu is a loanword from Swah ...
'', found in Central and East Africa. The fossils indicate that the males grew up to and the
queens Queens is a borough of New York City, coextensive with Queens County, in the U.S. state of New York. Located on Long Island, it is the largest New York City borough by area. It is bordered by the borough of Brooklyn at the western tip of Long ...
grew to . It had a wingspan of about .


Paleo-ecological implications

The fossils of ''Titanomyrma gigantea'', the first of the genus to be discovered, are very well preserved. They show that ''T. gigantea'' did not possess a sting and did not have a closing mechanism on the
crop A crop is a plant that can be grown and harvested extensively for profit or subsistence. When the plants of the same kind are cultivated at one place on a large scale, it is called a crop. Most crops are cultivated in agriculture or hydropon ...
. It is surmised that it must have sprayed formic acid as a defence, and either ate fresh food, in the manner of
leafcutter ant Leafcutter ants, a non-generic name, are any of 47 species of leaf-chewing ants belonging to the two genera '' Atta'' and ''Acromyrmex''. These species of tropical, fungus-growing ants are all endemic to South and Central America, Mexico, and par ...
s (which eat only the fungi they personally cultivate in their nests), or was carnivorous. Modern relatives include
driver ant ''Dorylus'', also known as driver ants, safari ants, or siafu, is a large genus of army ants found primarily in central and east Africa, although the range also extends to southern Africa and tropical Asia. The term siafu is a loanword from Swah ...
s. ''Titanomyrma'' may have been a precursor species, possibly following a raiding lifestyle and butchering large animals. ''T. lubei'' is related to fossil giant ants previously found in
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
and in the
Isle of Wight The Isle of Wight ( ) is a Counties of England, county in the English Channel, off the coast of Hampshire, from which it is separated by the Solent. It is the List of islands of England#Largest islands, largest and List of islands of England#Mo ...
in southern
England England is a country that is part of the United Kingdom. It shares land borders with Wales to its west and Scotland to its north. The Irish Sea lies northwest and the Celtic Sea to the southwest. It is separated from continental Europe b ...
dating from the same period. It is the first complete giant ant fossil from North America; previous evidence that big ants lived in North America during the
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', " ...
included an ancient ant wing from
Tennessee Tennessee ( , ), officially the State of Tennessee, is a landlocked state in the Southeastern region of the United States. Tennessee is the 36th-largest by area and the 15th-most populous of the 50 states. It is bordered by Kentucky to th ...
. It is a further example, in the northern temperate regions, of tropical fauna and flora such as
hippopotamus The hippopotamus ( ; : hippopotamuses or hippopotami; ''Hippopotamus amphibius''), also called the hippo, common hippopotamus, or river hippopotamus, is a large semiaquatic mammal native to sub-Saharan Africa. It is one of only two extan ...
precursors, tropical
plankton Plankton are the diverse collection of organisms found in water (or air) that are unable to propel themselves against a current (or wind). The individual organisms constituting plankton are called plankters. In the ocean, they provide a crucia ...
and pollen from tropical palms. ''T. lubei'' supports the idea that during the Eocene (), conditions such as
land bridge In biogeography, a land bridge is an isthmus or wider land connection between otherwise separate areas, over which animals and plants are able to cross and colonize new lands. A land bridge can be created by marine regression, in which sea leve ...
s and hot spells existed which permitted the spread of ancient warmth-loving insects and other forms of life from
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a subcontinent of Eurasia and it is located entirel ...
to North America or vice versa, which would not have occurred had the temperature been uniformly cool throughout.


References


External links

* * {{Taxonbar, from=Q1949915 Eocene insects of North America Formiciinae Eocene insects Fossil taxa described in 2011 Fossil ant genera