List of prehistoric insects

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Prehistoric insects are various groups of insects that lived before recorded history. Their study is the field of paleoentomology. Insects inhabited Earth since before the time of the dinosaurs. The earliest identifiable insect is the Devonian Rhyniognatha hirsti, estimated at 407 to 396 million years ago. Forms similar to many modern insects had already evolved before the dawning of the dinosaur and lived alongside them and beyond up to the present day. Like today, prehistoric insects were an important part of the food chain in their time.

The differences between modern and prehistoric varieties can be essential, and, like many other creatures of prehistory, the latter tended to be much larger than their contemporary equivalents. This size difference is thought to be due to higher atmospheric oxygen levels (allowing diffusion through spiracles over greater distances), higher temperatures (enhancing metabolism), and the absence of birds as key predators of insect life.[1]

Since insects have chitin exoskeletons rather than mineralized bones, their burial processes differ compared to the fossils of much larger vertebrates such as dinosaurs. Many insect remains are found preserved in the hardened sap of ancient trees (amber).

A weevil caught in Baltic amber

Selected taxa[edit]

Incomplete list of prehistoric insects arranged by extinction date:

Devonian[edit]

Name Authors Unit Location Notes Images

Rhyniognatha

Tillyard, 1928

Rhynie Chert

 Scotland

oldest described insect

Carboniferous[edit]

Name Authors Unit Location Notes Images

Meganeura

Brongniart, 1885

Europe

 Europe

three described species

Bohemiatupus[2]

Prokop & Nel, 2010

Kladno Formation

 Czech Republic

A griffenfly

Sinomeganeura[3]

Ren, Nel & Prokop, 2008

Tupo Formation

 China

A griffenfly

Permian[edit]

Jurassic[edit]

Name Authors Unit Location Notes Images

Formosibittacus[4]

Li, Ren & Shih, 2008

Jiulongshan Formation

 China

A hangingfly

Jurahylobittacus[4]

Li, Ren & Shih, 2008

Jiulongshan Formation

 China

A hangingfly

Mongolbittacus[5]

Petrulevicius, Huang & Ren, 2007

Jiulongshan Formation

 China

A hangingfly

Ororaphidia[6]

Engel & Ren, 2008

Jiulongshan Formation

 China

A snakefly

Cretaceous[edit]

Name Authors Unit Location Notes Images

Afromyrma[7]

Dlussky, Brothers & Rasnitsyn, 2004

 Botswana

A myrmicin ant

Afropone[7]

Dlussky, Brothers & Rasnitsyn, 2004

 Botswana

A ponerin ant, 2 species

Alavaraphidia[8]

Pérez-de la Fuente, et al, 2012

Spanish amber

 Spain

A snakefly

Amarantoraphidia[8]

Pérez-de la Fuente, et al, 2012

Spanish amber

 Spain

A snakefly

Armaniidae

Dlussky, 1983

Asia and Africa

A family of "ant like wasps"

Astreptolabis[9]

Engel, 2011

Burmese amber

 Myanmar

An earwig

Burmomyrma[10]

Dlussky, 1996

Burmese amber

 Myanmar

A possible aneuretin ant, one species

Cantabroraphidia[8]

Pérez-de la Fuente, et al, 2010

Spanish amber

 Spain

A snakefly

Deinodryinus? aptianus[11]

Olmi, Rasnitsyn & Guglielmino, 2010

Khurilt rock unit

 Mongolia

An earwig

Ektatotricha[12]

Chatzimanolis et al, 2010

Burmese amber

 Myanmar

A rove beetle

Haidomyrmex[13]

Dlussky, 1996

Burmese amber

 Myanmar

A sphecomyrmin ant, three species

Haidomyrmodes[14]

Perrichot et al, 2008

Charentese amber

 France

A sphecomyrmin ant

Haidoterminus[15]

McKellar, Glasier, & Engel, 2013

Canadian amber

 Canada

A sphecomyrmin ant

Iberoraphidia[16]

Jepson, Ansorge & Jarzembowski, 2011

Serra del Montsec

 Spain

A snakefly

Kachinus[12]

Chatzimanolis et al, 2010

Burmese amber

 Myanmar

A rove beetle

Lebanoraphidia[17]

Bechly & Wolf-Schwenninger, 2011

Lebanese amber

 Lebanon

A snakefly

Mesojassoides[18]

Oman, 1937

Fox Hills Formation

 USA

A leafhopper

Nanoraphidia[19]

Engel, 2002

Burmese amber

 Myanmar

A snakefly

Necroraphidia[8]

Pérez-de la Fuente, et al, 2010

Spanish amber

 Spain

A snakefly

Plumalexius[20]

Brothers, 2011

New Jersey Amber

 USA

A wasp of the extinct family Plumalexiidae

Proraphidia[21]

Martynova, 1941

Europe & Asia

A Jurassic - Cretaceous snakefly genus; three species

Styporaphidia[8]

Engel & Ren, 2008

Europe & Asia

A Jurassic - Cretaceous snakefly genus; two species

Tytthodiplatys[9]

Engel, 2011

Burmese amber

 Myanmar

An earwig

Undulopsychopsis[22]

Peng, Makarkin, Wang, & Ren, 2011

Yixian Formation

 China

A silky lacewing

Zigrasimecia[23]

Barden & Grimaldi, 2013

Burmese amber

 Myanmar

A sphecomyrmin ant

Eocene[edit]

Name Authors Unit Location Notes Images

Agroecomyrmex[24]

Wheeler, 1910

Baltic amber

 Europe

An agroecomyrmecin ant

Agulla protomaculata[25]

Engel, 2011

Green River Formation

 USA

A snake fly

Ainigmapsychops[26]

Makarkin & Archibald, 2014

Klondike Mountain Formation

 USA

A possible psychopsid lacewing

Anonychomyrma constricta[27]

(Mayr, 1868)

Baltic amber

 Europe

A dolichoderin ant

Anonychomyrma geinitzi[27]

(Mayr, 1868)

Baltic amber

 Europe

A dolichoderin ant

Anonychomyrma samlandica[27]

(Wheeler, 1915)

Baltic amber

 Europe

A dolichoderin ant

Anthidium exhumatum[28]

Cockerell, 1906

Florissant Formation

 USA

A mason bee

Anthidium scudderi[28]

Cockerell, 1906

Florissant Formation

 USA

A mason bee

Aphaenogaster donisthorpei[29][29]

Carpenter, 1930

Florissant Formation

 USA

A myrmicin ant

Aphaenogaster mayri[29]

Carpenter, 1930

Florissant Formation

 USA

A myrmicin ant

Aphaenogaster mersa[24]

Wheeler, 1915

Baltic amber

 Europe

A myrmicin ant

Aphaenogaster oligocenica[24]

Wheeler, 1915

Baltic amber

 Europe

A myrmicin ant

Aphaenogaster sommerfeldti[24]

Mayr, 1868

Baltic amber

 Europe

A myrmicin ant

Archiinocellia[30]

Handlirsch, 1910

Horsefly beds

 Canada

A snake fly

Archimyrmex[31]

Cockerell, 1923

Green River Formation, Messel Pit,
& Ventana Formation

 Europe USA,
 Argentina

A myrmeciine ant genus, four species described

Archiponera[29]

Carpenter, 1930

Florissant Formation

 USA

A ponerine ant

Arostropsis[32]

Yunakov & Kirejtshuk, 2011

Baltic Amber

 Europe

A broad nosed weevil

Asymphylomyrmex

Wheeler, 1915

Baltic Amber

 Europe

A dolichoderine ant.

Avitomyrmex[33]

Archibald, Cover, & Moreau, 2006

McAbee Fossil Beds

 Canada

A bulldog ant gneus; three species known

Baltimartyria[34]

Skalski, 1995

Baltic amber

 Europe

A metallic moth, two species

Brevivulva[35]

Gibson, 2009

Baltic amber

 Europe

A parasitic wasp

Carmenelectra[36]

Evenhuis, 2002

Baltic amber

 Europe

A Mythicomyiidae fly

Deinodryinus areolatus[11]

(Ponomarenko, 1975)

Baltic Amber

 Europe

A dryinid wasp

Deinodryinus velteni[11]

Guglielmino & Olmi, 2011

Baltic Amber

 Europe

A dryinid wasp

Denaeaspis[37]

Chaboo & Engel, 2009

Green River Formation

 USA

A tortoise beetle

Dinokanaga[38]

Archibald, 2005

Canada and the United States

 Canada
 USA

A scorpion fly genus, six species

Dinopanorpa[38]

Cockerell, 1924

Khutsin Formation

 Russia

A scorpion fly

Diochus electrus[39]

Chatzimanolis & Engel, 2011

Baltic amber

 Europe

a rove beetle

Dominickus[40]

Tindale, 1985

Florissant Formation

 USA

A butterfly-moth

Electrinocellia[41]

Engel, 1995

Baltic amber

 Europe

A snake fly

Electrostephanus[42]

Brues, 1933

Baltic amber

 Europe

A crown wasp, 4 species

Emiliana[43]

Shcherbakov, 2006

Green River Formation

 USA

A plant-hopper

Epiborkhausenites[44]

Skalski, 1973

Baltic amber

 Europe

A concealer moth

Eosacantha[37]

Chaboo & Engel, 2009

Green River Formation

 USA

A tortoise beetle

Eriocampa tulameenensis[45]

Rice, 1968

Allenby Formation

 Canada

A tenthredinid sawfly

Eulithomyrmex[29]

Carpenter, 1935

Florissant Formation

 USA

An agroecomyrmecin ant

Fibla carpenteri[41]

Engel, 1995

Baltic amber

 Europe

A snakefly

Formicium[46]

John O. Westwood, 1854

North America and Europe

A form genus of ants

Glisachaemus[47]

Szwedo, 2007

Baltic amber

 Europe

A plant-hopper

Halictus? savenyei[48]

Engel & Archibald, 2003

Princeton Group

 Canada

A sweat bee

Hydriomena? protrita[49]

Cockerell, 1922

Florissant Formation

 USA

A geometrid moth

Macabeemyrma[33]

Archibald, Cover, & Moreau, 2006

McAbee Fossil Beds

 Canada

A bulldog ant

Metanephrocerus[50]

Carpenter & Hull, 1939

Klondike Mountain Formation & Baltic amber

 USA
 Russia

A genus of pipunculids, 4 species

Metapelma archetypon[35]

Gibson, 2009

Baltic amber

 Europe

A parasitic wasp

Microberotha[51]

Archibald & Makarkin, 2004

Hat Creek Amber

 Canada

A lacewing

Myrmeciities[33]

Archibald, Cover, & Moreau, 2006

Canada and the United States

 Canada
 USA

A bulldog ant form genus

Nanotermes[25]

Engel & Grimaldi, 2011

Cambay Amber

 India

A termite

Neanaperiallus[35]

Gibson, 2009

Baltic amber

 Europe

A parasitic wasp

Neoephemera antiqua[52]

Sinitchenkova, 1999

Klondike Mountain Formation

 USA

A mayfly

Ordralfabetix[53]

Szwedo, 2011

Oise amber

 France

A plant-hopper

Palaeovespa[54]

Cockerell, 1906

Baltic amber
Florissant Formation

 Europe
 USA

A vespid wasp;
seven described species

Pristomyrmex rasnitsyni[55]

Dlussky & Radchenko, 2011

Scandinavian amber

 Denmark

An ant

Prodryas[56]

Scudder, 1878

Florissant Formation

 USA

A butterfly

Prostylotermes[25]

Engel & Grimaldi, 2011

Cambay Amber

 India

A termite

Protostephanus[42]

Cockerell, 1906

Florissant Formation

 USA

A crown wasp

Pseudosiobla campbelli[45]

Rice, 1968

Allenby Formation

 Canada

A tenthredinid sawfly

Raphidia funerata[48]

Engel, 2003

Florissant Formation

 USA

A snakefly

Tortrix? destructus[57]

Cockerell, 1917

Florissant Formation

 USA

A tortricid moth

Tortrix? florissantana[58]

Cockerell, 1917

Florissant Formation

 USA

A tortricid moth

Ypresiomyrma[33]

Archibald, Cover, & Moreau, 2006

McAbee Fossil Beds
Ølst and Fur Formations

 Canada
 Denmark

A bulldog ant genus

Zophotermes [25]

Engel & Singh, 2011

Cambay Amber

 India

A termite

Oligocene[edit]

Name Authors Unit Location Notes Images

Aphaenogaster longaeva[29]

Scudder, 1877

Fraser Formation?

 Canada

myrmicin ant species, generic placement uncertain

Miocene[edit]

Name Authors Unit Location Notes Images

Acanthognathus poinari[59]

Baroni Urbani & De Andrade, 1994

Dominican amber

 Dominican Republic

A trap-jaw ant

Acanthostichus hispaniolicus[60]

De Andrade, 1998

Dominican amber

 Dominican Republic

A doryline ant

Acropyga glaesaria[61]

Lapolla, 2005

Dominican amber

 Dominican Republic

Oldest record of trophophoresy in ants

Anochetus ambiguus[62]

De Andrade, 1994

Dominican amber

 Dominican Republic

A ponerin ant

Anochetus brevidentatus[63]

MacKay, 1991

Dominican amber

 Dominican Republic

A ponerin ant

Anochetus conisquamis[62]

De Andrade, 1994

Dominican amber

 Dominican Republic

A ponerin ant

Anochetus dubius[62]

De Andrade, 1994

Dominican amber

 Dominican Republic

A ponerin ant

Anochetus exstinctus[62]

De Andrade, 1994

Dominican amber

 Dominican Republic

A ponerin ant

Anochetus intermedius[62]

De Andrade, 1994

Dominican amber

 Dominican Republic

A ponerin ant

Anochetus lucidus[62]

De Andrade, 1994

Dominican amber

 Dominican Republic

A ponerin ant

Anthophorula persephone[64]

Engel, 2012

Dominican amber

 Dominican Republic

A bee

Aphaenogaster amphioceanica[65]

De Andrade, 1995

Dominican amber

 Dominican Republic

A myrmicin ant

Aphaenogaster avita[66]

Fujiyama, 1970

Chojabaru Formation

 Japan

A myrmicin ant

Aphaenogaster praerelicta[65]

De Andrade, 1995

Mexican amber

 Mexico

A myrmicin ant

Apterostigma electropilosum[67]

Schultz, 2007

Dominican amber

 Dominican Republic

A myrmicin fungus farming ant

Apterostigma eowilsoni[67]

Schultz, 2007

Dominican amber

 Dominican Republic

A myrmicin fungus farming ant

Attopsis longipennis[68]

Heer, 1849

Radoboj

 Croatia

A formicine ant.

Augochlora leptoloba[69]

Engel, 1995

Dominican amber

 Dominican Republic

A sweat bee

Dryinus grimaldii[70]

Olmi, 1995

Dominican amber

 Dominican Republic

A dryinid wasp

Dryinus rasnitsyni[70]

Olmi & Guglielmino, 2011

Dominican amber

 Dominican Republic

A dryinid wasp

Eickwortapis[71]

Michener & Poinar, 1996

Dominican amber

 Dominican Republic

A sweat bee

Electromyrmococcus[72]

Williams, 2001

Dominican amber

 Dominican Republic

A mealybug; three described species

Electrotettix[73]

Heads, Thomas, & Wang, 2014

Dominican amber

 Dominican Republic

A tetrigid grasshopper.

Hyptia deansi[74]

Henderickx, 2012

Mexican amber

 Mexico

Late Oligocene - Early Miocene; An ensign wasp

Leptofoenus pittfieldae

Engel, 2009

Dominican amber

 Dominican Republic

A pteromalid wasp

Lutzomyia adiketis[75]

Poinar, 2008

Dominican amber

 Dominican Republic

A phlebotomid fly

Mastotermes electromexicus[76]

Krishna & Emerson, 1983

Mexican amber

 Mexico

Late Oligocene - Early Miocene

Neocorynura electra[77]

Engel, 1995

Dominican amber

 Dominican Republic

A sweat bee

Nesagapostemon[78]

Engel, 2009

Dominican amber

 Dominican Republic

A sweat bee

Neurosymploca? oligocenica[79]

Fernández-Rubio & Nel, 2000

paleolake Céreste

 France

A zygaenid moth

Oligochlora[80]

Engel, 1996

Dominican amber

 Dominican Republic

A sweat bee genus, six species

Parastylotermes[25]

Snyder & Emerson, 1949

North America, Europe, India

A termite genus;
lived from the Eocene to the Miocene
five described species

Pintomyia falcaorum[81]

Brazil & Andrade Filho, 2002

Dominican amber

 Dominican Republic

A phlebotomin fly

Protosialis casca

Engel & Grimaldi, 2007

Dominican amber

 Dominican Republic

An alderfly

Syndesus ambericus[82]

Woodruff, 2009

Dominican amber

 Dominican Republic

A stag beetle

Schwenckfeldina archoica[83]

Mohrig & Solórzano Kraemer, 2007

Mexican amber

 Mexico

Late Oligocene - Early Miocene; A sciarid fly

Tainosia[84]

Szwedo, 2001

Dominican amber

 Dominican Republic

A plant-hopper

Termitaradus avitinquilinus[78]

Grimaldi & Engel, 2008

Dominican amber

 Dominican Republic

A termite bug

Termitaradus dominicanus[85]

Poinar, 2011

Dominican amber

 Dominican Republic

A termite bug

Termitaradus mitnicki[78]

Engel, 2009

Dominican amber

 Dominican Republic

A termite bug

Termitaradus protera[85]

Brongniart, 1885

Mexican amber

 Mexico

Late Oligocene - Early Miocene; A termite bug

Triatoma dominicana[86]

Poinar, 2005

Dominican amber

 Dominican Republic

An assassin bug

See also[edit]

References[edit]

  1. ^ Giant prehistoric insects shrank to escape developing birds, Bloomberg News, 4th June 2012
  2. ^ Prokop, J.; Nel, A. (2010). "New griffenfly, Bohemiatupus elegans from the Late Carboniferous of western Bohemia in the Czech Republic (Odonatoptera: Meganisoptera: Meganeuridae)". Annales de la Société Entomologique de France 46 (1-2): 183–188. doi:10.1080/00379271.2010.10697655. 
  3. ^ Ren, D.; Nel, A.; Prokop, J. (2008). "New early griffenfly, Sinomeganeura huangheensis from the Late Carboniferous of northern China (Meganisoptera: Meganeuridae)". Insect Systematic Evolution 38: 223–229. doi:10.1163/187631208788784075. ISSN 1399-560X. 
  4. ^ a b Li, Y-L.; Ren, D.; Shih, C-K (2008). "Two Middle Jurassic hanging-flies (Insecta: Mecoptera: Bittacidae) from Northeast China". Zootaxa 1929: 38–46. ISSN 1175-5334. 
  5. ^ Petrulevicius, J. F.; Huang, D-Y.; Ren, D. (2007). "A new hangingfly (Insecta: Mecoptera: Bittacidae) from the Middle Jurassic of Inner Mongolia, China". African Invertebrates 48 (1): 145–152. 
  6. ^ Engel, M.S.; Ren, D. (2008). "New Snakeflies from the Jiulongshan Formation of Inner Mongolia, China (Raphidioptera)". Journal of the Kansas Entomological Society 81: 188–193. doi:10.2317/JKES-802.19.1. 
  7. ^ a b Dlussky, G. M.; Brothers, D. J.; Rasnitsyn, A. P. (2004). "The first Late Cretaceous ants (Hymenoptera: Formicidae) from southern Africa, with comments on the origin of the Myrmicinae". Insect Systematics and Evolution 35: 1–13. doi:10.1163/187631204788964727. 
  8. ^ a b c d e Pérez-de la Fuente, R.; Peñalver, E.; Delclòs, X.; Engel, M.S. (2012). "Snakefly diversity in Early Cretaceous amber from Spain (Neuropterida, Raphidioptera)". ZooKeys 204: 1–40. doi:10.3897/zookeys.204.2740. PMC 3391719. PMID 22787417. 
  9. ^ a b Engel, M.S. (2011). "New earwigs in mid-Cretaceous amber from Myanmar (Dermaptera, Neodermaptera)". ZooKeys 130: 137–152. doi:10.3897/zookeys.130.1293. 
  10. ^ Dlussky, G. M. (1996). "Ants (Hymenoptera: Formicidae) from Burmese amber". Paleontological Journal 30 (4): 449–454. doi:10.1206/3755.2. 
  11. ^ a b c Guglielmino, A.; Olmi, M. (2011). "Revision of fossil species of Deinodryinus, with description of a new species (Hymenoptera, Dryinidae)". ZooKeys 130: 495–504. doi:10.3897/zookeys.130.1326. PMC 3260777. PMID 22259295. 
  12. ^ a b Stylianos Chatzimanolis; Michael S. Engel; Alfred F. Newton; David A. Grimaldi (2010). "New ant-like stone beetles in mid-Cretaceous amber from Myanmar (Coleoptera:Staphylinidae: Scydmaeninae)". Cretaceous Research 31 (1): 77–84. doi:10.1016/j.cretres.2009.09.009. 
  13. ^ Barden, P.; Grimaldi, D. (2012). "Rediscovery of the bizarre Cretaceous ant Haidomyrmex Dlussky (Hymenoptera: Formicidae), with two new species". American Museum Novitates 3755: 1–16. doi:10.1206/3755.2. 
  14. ^ Perrichot, V.; Nel, A.; Néraudeau, D.; Lacau, S.; Guyot, T. (2008). "New fossil ants in French Cretaceous amber (Hymenoptera: Formicidae)". Naturwissenschaften 95 (2): 91–97. doi:10.1007/s00114-007-0302-7. PMID 17828384. 
  15. ^ McKellar, R. C.; Glasier, J. R. N.; Engel, M. S. (2013). "A new trap-jawed ant (Hymenoptera: Formicidae: Haidomyrmecini) from Canadian Late Cretaceous amber". Canadian Entomologist 145: 454–465. doi:10.4039/tce.2013.23. 
  16. ^ Jepson, J.E.; Ansorge, J.; Jarzembowski, E.A. (2011). "New snakeflies (Insecta: Raphidioptera) from the Lower Cretaceous of the UK, Spain and Brazil". Palaeontology 54 (2): 385–395. doi:10.1111/j.1475-4983.2011.01038.x. 
  17. ^ Bechly, G.; Wolf-Schwenninger, K. (2011). "A new fossil genus and species of snakefly (Raphidioptera: Mesoraphidiidae) from Lower Cretaceous Lebanese amber, with a discussion of snakefly phylogeny and fossil history". Insect Systematics and Evolution 42 (2): 221–236. doi:10.1163/187631211X568164. 
  18. ^ Oman, P. W. (January 1937). Journal of Paleontology 11 (1): 37–38. 
  19. ^ Engel, M.S. (2002). "The Smallest Snakefly(Raphidioptera: Mesoraphidiidae): A New Species in Cretaceous Amber from Myanmar, with a Catalog of Fossil Snakeflies". American Museum Novitates 3363: 1–22. doi:10.1206/0003-0082(2002)363<0001:TSSRMA>2.0.CO;2. hdl:2246/2852. 
  20. ^ Brothers, Denis J. (2011). "A new Late Cretaceous family of Hymenoptera, and phylogeny of the Plumariidae and Chrysidoidea (Aculeata)". ZooKeys 130: 515–542. doi:10.3897/zookeys.130.1591. PMC 3260779. PMID 22259297. 
  21. ^ Jepson, J.E.; Jarzembowski, E.A. (2008). "Two new species of snakefly (Insecta:Raphidioptera) from the Lower Cretaceous of England and Spain with a review of other fossil raphidiopterans from the Jurassic/Cretaceous transition". Alavesia 2: 193–201. 
  22. ^ Peng, Y.; Makarkin, V.N.; Wang, X.; Ren, D. (2011). "A new fossil silky lacewing genus (Neuroptera, Psychopsidae) from the Early Cretaceous Yixian Formation of China". ZooKeys 130: 217–228. doi:10.3897/zookeys.130.1576. 
  23. ^ Barden, P.; Grimaldi, D. (2013). "A New Genus of Highly Specialized Ants in Cretaceous Burmese Amber (Hymenoptera: Formicidae)". Zootaxa 3681 (4): 405–412. doi:10.11646/zootaxa.3681.4.5. 
  24. ^ a b c d Wheeler, W. M. (1915). "The ants of the Baltic amber". Schriften der Physikalisch-Okonomischen Gesellschaft zu Konigsberg 55 (4): 56–59. 
  25. ^ a b c d e Engel, Michael S. (2011). "A new snakefly from the Eocene Green River Formation (Raphidioptera: Raphidiidae)". Transactions of the Kansas Academy of Science 114 (1-2): 77–87. doi:10.1660/062.114.0107. 
  26. ^ Makarkin, V.; Archibald, S.B. (2014). "An unusual new fossil genus probably belonging to the Psychopsidae (Neuroptera) from the Eocene Okanagan Highlands, western North America". Zootaxa 3838 (3): 385–391. doi:10.11646/zootaxa.3838.3.8. 
  27. ^ a b c Heterick, B. E.; Shattuck, S. (2011). "Revision of the ant genus Iridomyrmex (Hymenoptera: Formicidae)". Zootaxa 2845: 169. 
  28. ^ a b Cockerell, T.D.A. (1906). "Fossil Hymenoptera from Florissant, Colorado". Bulletin of the Museum of Comparative Zoology L (2). 
  29. ^ a b c d e f Carpenter, F. M. (1930). "The fossil ants of North America.". Bulletin of the Museum of Comparative Zoology 70: 1–66. 
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