Prionomyrmex

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Prionomyrmex
Temporal range: Eocene to Late Oligocene, 56-29  Ma
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Prionomyrmex longiceps dorsal view Mbi5827d1.jpg
P. longiceps
Scientific classification Red Pencil Icon.png
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Myrmeciinae
Tribe: Prionomyrmecini
Genus:† Prionomyrmex
Mayr, 1868 [1]
Type species
Prionomyrmex longiceps
Species
  • P. gusakoviRadchenko & Perkovsky, 2020
  • P. janzeniBaroni Urbani, 2000
  • P. longicepsMayr, 1868
  • P. wappleriDlussky, 2012

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.

Contents

Discovery and classification

The holotype worker for P. longiceps was collected by Austrian entomologist Gustav Mayr in 1868. [2] The fossil, which was preserved in Baltic amber from the Eocene, was formally described in Mayr's journal article Die Ameisen des baltischen Bernsteins, designating it as the type species by monotypy (the condition of a taxonomic group having only a single taxon described) for the newly established genus Prionomyrmex. Originally, the genus was placed in the subfamily Ponerinae by Mayr, but in 1877, Italian entomologist Carlo Emery classified the genus into the subfamily Myrmeciidae (now known as Myrmeciinae), the same year Emery established the subfamily. [3] In 1915, the tribe Prionomyrmecini was erected by American entomologist William Morton Wheeler, who had placed Prionomyrmex in it. [4] In that year, Wheeler placed the genus back into Ponerinae without any means of justifying his decision. British myrmecologist Horace Donisthorpe would also retain the genus in Ponerinae without explanation, but William Brown Jr. would return it to Myrmeciinae in 1954. [5] [6]

In 2000, Baroni Urbani described a new Baltic fossil species, of which he named it Prionomyrmex janzeni. [7] After examining specimens of the new species and Nothomyrmecia macrops , Baroni Urbani stated that the newly described species and Nothomyrmecia macrops belong to the same genus (Prionomyrmex), in which he synonymised Nothomyrmecia as a genus and treated the tribe Prionomyrmecini as a subfamily, known as Prionomyrmecinae. [7] Prior to this, John S. Clark, the original author who described Nothomyrmecia, noted that the genus was similar in appearance to Prionomyrmex; both the heads and mandibles were identical, but the nodes were different. [8] As the mandibles of Prionomyrmex are similar to that of Nothomyrmecia, this suggests that they are intermediate to each other. [9] This classification was short-lived, as Nothomyrmecia was separated and treated as a valid genus from Prionomyrmex by Dlussky & Perfilieva in 2003, on the base of the fusion of an abdominal segment. [10] Other studies published in the same year came to the same conclusions of Dlussky & Perfilieva, and the subfamily Prionomyrmecinae would later be treated as a tribe in Myrmeciinae. [11] However, Baroni Urbani would treat the tribe as a subfamily again in both his 2005 and 2008 publications, suggesting additional evidence in favor of his former interpretation as opposed to that of Ward and Brady's arguments. [12] [13] In 2012, P. wappleri was described by Gennady M. Dlussky, based on a fossilised worker from the Late Oligocene, Aquitanian stage. This subsequent report that described new fossil myrmecines accepted the classification of Archibald et al. and Ward & Brady without comment on the views of Baroni Urbani. [14]

The generic name is a combination of two words; priono derives from Greek word priōn, meaning "a saw", and myrmex, another Greek word, means "ant". [15] [16]

The following cladogram generated by Archibald and colleagues shows the possible phylogenetic position of Prionomyrmex among some ants of the subfamily Myrmeciinae; note that P. wappleri is absent, as the generated cladogram below was created in 2006 while the species itself was described in 2012. [17]

P. longiceps casent label Prionomyrmex longiceps MBI5827 with museum tags.jpg
P. longiceps casent label
Myrmeciinae

Archimyrmex

Myrmecia

Prionomyrmex janzeni

Prionomyrmex longiceps

Macabeemyrma

Avitomyrmex

Ypresiomyrma

Nothomyrmecia

Description

The genus is characterised by large, slender workers with elongated mandibles, which are narrow and triangular in shape. The mesosoma and appendages are also long. [14] These ants are similar in appearance to Nothomyrmecia, but can be distinguished from the shape of their node. [8] They also had a powerful sting located in the abdomen. [6] Two of the three species are from the Eocene while the third species is from the Late Oligocene. [14] [18]

P. janzeni

P. janzeni Prionomyrmex janzeni GPIH-BE1301 profile.jpg
P. janzeni

P. janzeni was described by Cesare Baroni Urbani of the University of Basel, Switzerland in 2000, based on two specimens preserved in Baltic amber from Kaliningrad, Russia. [7] [19] The species is from the Eocene, Lutetian to Priabonian stage. [17] Both specimens are preserved very well, with specimen number two being larger and more visible. Specimen number one is presumed to be a worker; type material includes a holotype worker and a paratype ergatogyne, donated to the Geological-Palaecontological Institute and Museum, the University of Hamburg by palaeoentomologist Jens-Wilhelm Janzen. The ant was named after Janzen by Baroni Urbani. [7] The estimated body length is 13 millimetres (0.51 in) long with an elongated head, and large oval-shaped eyes are present. [19] The antennae are long and consist of 12 segments with a bent scape. The mandibles are very long and curved, being three-quarters the length of the total size of the head. Both the legs and mesosoma are long and slender; unlike modern ants, P. janzeni has two spurs on the tibiae instead of one. The petiole is high and domed shape while postpetiole is bell-shaped. The gaster is long with round sides, divided into five segments. The whole body and some portions of the legs were covered by weakly curved hairs, erect and suberect. The holotype specimen is brown in colour while the paratype is black. While P. janzeni looks similar to P. longiceps, the pubescence on the scapes of P. janzeni is absent. [7]

P. longiceps

P. longiceps illustrated by Wheeler, 1915 Prionomyrmex longiceps Wheeler 1915.jpg
P. longiceps illustrated by Wheeler, 1915

P. longiceps was described by Gustav Mayr in 1868, based on a holotype worker collected in Baltic amber from the Eocene. [2] The original specimen collected by Mayr, however, has been lost. The estimated body length of P. longiceps is 12 to 14 millimetres (0.47 to 0.55 in) long, with a thick petiole and large propodeal teeth. Unlike P. janzeni, P. longiceps has erect and suberect hairs on the scape. These hairs are also longer and thicker on the legs and on other body parts. [19] The mandibles are very long and curved, and the head is longer than its total width (2.2 to 2.64 millimetres (0.087 to 0.104 in) long and 1.68 to 2.08 millimetres (0.066 to 0.082 in) wide). [2] [19] The legs are very long with strong claws, and a stinger is present in the abdomen. [4] [6] Wheeler (1915) described a male P. longiceps, commenting that the head is short but broad with very large eyes while the mandibles are small and far apart. The body is dark brown or blackish in colour, and the wings are somewhat yellowish. [4]

P. wappleri

P. wappleri worker Prionomyrmex wappleri Dlussky.jpg
P. wappleri worker

P. wappleri was described in 2012 by Russian palaeoentomologist Gennady M. Dlussky of the Moscow State University, from a fossilised holotype worker found in Germany from the Aquitanian stage 29 to 30 million years ago. The specimen is currently housed in the Institut für Paläontologie at the University of Bonn, North Rhine-Westphalia. Dlussky coined the specific epithet wappleri from the surname "Wappler", as he named the ant after German palaeoentomologist Torsten Wappler. [14] The estimated body length of P. wappleri is 14 millimetres (0.55 in) long, and the head is 1.35 times longer than the total width of it. The eyes are small and oval shaped, located in the upper part of the head, which is four times as long as the eyes. The mandibles are nearly three-quarters the length of the head; P. wappleri differs from P. longiceps and P. janzeni due to the apex of the clypeal lobe being pointed instead of round, and the first segment of the flagellum is only half the length of the second segment. Before the discovery of P. wappleri, extinct Myrmeciinae ants were only found from Eocene deposits. This suggests that the subfamily was still present in Europe during the Late Oligocene. [14]

Ecology

Archibald and colleagues suggested the life habits of extinct Myrmeciinae ants including Prionomyrmex may have been similar to extant ants within the subfamily. These ants foraged on the ground and possibly onto trees and low vegetation while preying on arthropods. These ants may have collected plant nectar, as Myrmecia species use this as a food source. [17] Workers may have not recruited nest mates to food sources or lay down pheromone trails, as these ants were solitary hunters. Workers would have relied on their vision to hunt for prey and help themselves navigate. [17] [20]

William Morton Wheeler comments that P. longiceps were possibly an arboreal nesting species. This means the ant did not live in the soil and nested in trees instead. He assumed this due to its long legs, strong claws and long mandibles; Prionomyrmex was also assumed to be predacious, equipped with a well developed and powerful sting that was most likely used to kill prey. Prionomyrmex may have preferred a jungle habitat at low elevations, and is even more primitive in its body structure than Myrmecia . [4] [6] Female stylopids were known to parasite Prionomyrmex ants. [21]

Related Research Articles

<span class="mw-page-title-main">Myrmeciinae</span> 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. Subsequently an additional fossil genus was moved from the family Rhopalosomatidae in 2018, and a new genus was described in 2021.

<i>Nothomyrmecia</i> Genus of ants

Nothomyrmecia, also known as the dinosaur ant or dawn ant, is an extremely 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>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.

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

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.

<span class="mw-page-title-main">Prionomyrmecini</span> 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.

Pristomyrmex rasnitsyni is an extinct species of ant in the genus Pristomyrmex. The species is known from a single Late Eocene fossil which was found in Europe.

Acanthognathus poinari is an extinct species of ant in the subfamily Myrmicinae known from a single possibly Miocene fossil found on Hispaniola. A. poinari is the first species of the ant genus Acanthognathus to have been described from fossils found in Dominican amber and is one of several species of Acanthognathus found in the Greater Antillas.

<i>Anochetus corayi</i> Extinct species of ant

Anochetus corayi is an extinct species of ant in the subfamily Ponerinae known from one possibly Miocene fossil found on Hispaniola. A. corayi is one of eight species in the ant genus Anochetus to have been described from fossils found in Dominican amber and is one of a number of Anochetus species found in the Greater Antillies.

<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>Emplastus</i> Genus of ants

Emplastus is an extinct morphogenus of ants in the subfamily Dolichoderinae, known from fossils found in Asia and Europe. The genus contains twelve species described from sites in England, Eastern Europe and Far Eastern Russia.

<i>Casaleia</i> Genus of ants

Casaleia is an extinct genus of ants in the formicid subfamily Amblyoponinae described by Pagliano & Scaramozzino in 1990 from fossils found in Europe. The genus contains four species dating from the Eocene to Miocene, Casaleia eocenica, Casaleia inversa, Casaleia longiventris, Casaleia orientalis.

<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>Zherichinius</i> Genus of ants

Zherichinius is an extinct genus of ants in the subfamily Dolichoderinae known from fossils found in amber from the Middle Eocene of Sakhalin island Far eastern Russia and Bitterfeld, Germany. At the time of description the species Zherichinius horribilis and Zherichinius rapax were two of eight ant species known from Sakhalin fossils.

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

Pachycondyla petrosa is an extinct species of ant in the formicid subfamily Ponerinae described from a fossil found in Europe. P. petrosa is one of six Lutetian Pachycondyla species.

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

Bradoponera is an extinct genus of ant in the Formicidae subfamily Proceratiinae, and is one of four genera of the subfamily. The genus contains four described species Bradoponera electrina, Bradoponera meieri, Bradoponera similis, and Bradoponera wunderlichi. The species are known from several Middle Eocene amber fossils which were found in Europe.

<i>Nylanderia pygmaea</i> Extinct species of ant

Nylanderia pygmaea is an extinct species of formicid in the ant subfamily Formicinae known from fossils found in the Baltic region.

References

  1. ↑ Johnson, Norman F. (19 December 2007). "Prionomyrmex Mayr, 1868". Hymenoptera Name Server version 1.5. Columbus, Ohio, USA: Ohio State University . Retrieved 4 August 2015.
  2. 1 2 3 Mayr, Gustav (1868). "Die Ameisen des baltischen Bernsteins" (PDF). Beiträge zur Naturkunde Preussens Königlichen Physikalisch-Oekonomischen Gesellschaft zu Königsberg. 1: 1–102. doi:10.5281/zenodo.25852.
  3. ↑ Emery, Carlo (1877). "Saggio di un ordinamento naturale dei Mirmicidei e considerazioni sulla filogenesi delle formiche" (PDF). Bullettino della Società Entomologica Italiana. 9: 67–83. doi:10.5281/zenodo.25408.
  4. 1 2 3 4 Wheeler, W. M. (1915). "The ants of the Baltic amber" (PDF). Schriften der Physikalisch-Okonomischen Gesellschaft zu Konigsberg. 55 (4): 56–59. doi:10.5962/bhl.title.14207.
  5. ↑ Donisthorpe, Horace (1943). "A list of the type-species of the genera and subgenera of Formicidae". Annals and Magazine of Natural History. 10 (11): 617–688. doi:10.5281/zenodo.26489.
  6. 1 2 3 4 Brown, W. L. (1954). "Remarks on the internal phylogeny and subfamily classification of the family Formicidae". Insectes Sociaux. 1 (1): 21–31. doi:10.1007/BF02223148. S2CID   33824626.
  7. 1 2 3 4 5 Baroni Urbani, Cesare (2000). "Rediscovery of the Baltic amber ant genus Prionomyrmex (Hymenoptera, Formicidae) and its taxonomic consequences". Eclogae Geologicae Helveticae. 93 (3): 471–480.
  8. 1 2 Clark, John S. (1934). "Notes on Australian ants, with descriptions of new species and a new genus" (PDF). Memoirs of the National Museum of Victoria. 8: 5–20. doi:10.24199/j.mmv.1934.8.01.
  9. ↑ Brown, William Brown Jr.; Wilson, E.O. (1959). "The search for Nothomyrmecia" (PDF). Western Australian Naturalist. 7 (2): 25–30. Archived from the original (PDF) on 2015-11-21.
  10. ↑ Dlussky, G.M; Perfilieva, K.S. (2003). "Paleogene ants of the genus Archimyrmex Cockerell, 1923 (Hymenoptera, Formicidae, Myrmeciinae)". Paleontological Journal. 37: 39–47.
  11. ↑ Ward, Philip S.; Brady, Seán G. (2003). "Phylogeny and biogeography of the ant subfamily Myrmeciinae (Hymenoptera : Formicidae)" (PDF). Invertebrate Systematics. 17 (3): 361–386. doi:10.1071/IS02046.
  12. ↑ Baroni Urbani, Cesare (2005). "Phylogeny and biogeography of the ant subfamily Prionomyrmecinae (Hymenoptera: Formicidae)" (PDF). Annali del Museo Civico di Storia Naturale di Genova. 96: 581–595. Archived from the original (PDF) on 2015-11-20.
  13. ↑ 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.
  14. 1 2 3 4 5 Dlussky, G.M. (2012). "New Fossil Ants of the Subfamily Myrmeciinae (Hymenoptera, Formicidae) from Germany". Paleontological Journal. 46 (3): 288–292. doi:10.1134/s0031030111050054. S2CID   83891156.
  15. ↑ Harper, Douglas. "Prion". Online Etymology Dictionary .
  16. ↑ Harper, Douglas. "Formica". Online Etymology Dictionary .
  17. 1 2 3 4 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. S2CID   4845957.
  18. ↑ Wilson, E. O.; Hölldobler, B. (17 May 2005). "The rise of the ants: A phylogenetic and ecological explanation". Proceedings of the National Academy of Sciences. 102 (21): 7411–7414. Bibcode:2005PNAS..102.7411W. doi: 10.1073/pnas.0502264102 . PMC   1140440 . PMID   15899976.
  19. 1 2 3 4 Baroni Urbani, Cesare (2002). "The Baltic amber species of Prionomyrmex (Hymenoptera, Formicidae)". Mitteilungen aus dem Geologisch-Paläontologischen Institut der Universität Hamburg. 87: 141–146.
  20. ↑ Veeresh, G.K.; Mallik, B.; Viraktamath, C.A. (1990). Social insects and the environment. Leiden: E.J. Brill. p. 314. ISBN   978-90-04-09316-4.
  21. ↑ Kinzelbach, Ragnar; Pohl, Hans (2001). "First record of a female stylopid (Strepsiptera: ?Myrmecolacidae) parasite of a prionomyrmecine ant (Hymenoptera: Formicidae) in Baltic amber". Insect Systematics & Evolution. 32 (2): 143–146. doi:10.1163/187631201X00092. ISSN   1399-560X.
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