View source for Fossil Ants

Fossil ants include both specimens fossilised in shale or other stone (compression fossils) and ants embedded in Amber.  Copal is a general term that refers to non-fossilised resin material whatever its geological age.  Copal older than Holocene is referred to as fossil copal. 

The geologic age of the deposits are important in understanding the evolution of ants as well as their extinction and the distribution of ants worldwide.  In some cases the surrounding vegetation and insects that are preserved along with the ants allows a partial reconstruction of the habitat and in some cases the microhabitat as well.

Winged and worker ants were much larger in times past so that some ant specimens are still identifiable as ants when found in stone.  Specimens preserved in amber are in much better condition allowing for a more reliable identification.
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{{Photo Gallery
|name1=Gerontoformica cretacica.jpg
|comment1=One of the earliest known ants, ''[[Gerontoformica cretacica]]'', from [[Charentese Amber|Charentese amber]], Aquitaine Basin, France. It belongs to the subfamily Specomyrminae, the most primitive group of true ants.
|size1=500px
|name2=Azteca alphaD.jpg
|comment2= ''[[Azteca alpha]]'' from [[Dominican Amber|Dominican Republic Amber]].
|size2=500px
|name3=Baltic amber inclusions - Ant (Hymenoptera, Formicidae)10.JPG
|comment3=An inclusion from [[Baltic Amber|Baltic amber]].
|size3=500px
|name4=Protoazteca-elongataF.jpg
|comment4=''[[Protazteca elongata]]'', based on an impression fossil.
|size4=500px
}}

==Amber - Fossil Resins==
A few plant resins have the ability to fossilize and insects trapped and embeded in this matrix are known as amber fossils.  Resin that is older than one million years are designated as amber and younger aged resins are known as copal.

Amber is often designated by the type of plant exudate or given a mineral name.  Ant fossils are most often named for its geographical origin; Dominican amber, Mexican amber, Baltic amber, Saxonian amber, Rovno amber, Spanish amber, Lebanese amber, etc.

==Copal==
Subfossil resins or Copal can be Carbon-14 dated so that the resin sold as "amber" in Madagascar by gem dealers is approximately 50 years old.  The Copal from Colombia has been dated as 250 years old.

==[[Ichnofossils]]==
Trace fossils of ant nests. The evidence for these fossils representing preserved ant nest structures are based on interpretation of various elements of the ichnofossils that are analogous to modern day ant nests.

==Fossil Taxa==
*[[:Category:Extinct_subfamily|Subfamilies]]
*[[:Category:Extinct_genus|Genera]]
*[[:Category:Extinct_species|Species]]
*[[:Category:Fossil|All fossils]] (subfamilies, genera and species)

==Geologic Time Zones Containing Ants==
*[[Pliocene]] - About 5.333 to 3.6 million years ago.
*[[Miocene]] - About 23.03 to 5.332 million years ago.
*[[Oligocene]] - About 34 million to 23 million years before the present.
*[[Eocene]] - Lasted approximately from 56 to 34  million years ago.
*[[Paleocene]] - Lasted from about 65.5 to 56 million years ago.
*[[Cretaceous]] - From circa 145.5 to 65.5 million years ago.

==Taxa by Geologic Time Zones==
{| class="wikitable collapsible" style="clear:both;margin:0; font-size:95%"
! Era
! Period
! Epoch
! Age
! Start, million years age
! style="width: 15%"| Formations
! Taxa
|-
|rowspan="24" style="background:{{period color|Cenozoic}}"| Cenozoic
|rowspan="7" style="background:{{period color|Quaternary}}"| Quaternary
|rowspan="3" style="background:{{period color|Holocene}}"| Holocene
|style="background:#fcf0f2"|Meghalayan
|style="background:#fcf0f2"| {{Period start|meghalayan}}
|
|
|-
|style="background:#fcf0e8"| Northgrippian
|style="background:#fcf0e8"| {{Period start|northgrippian}}
|
|
|-
|style="background:#fcf0de"| Greenlandian
|style="background:#fcf0de"| {{Period start|greenlandian}}
|
|
|-
|rowspan="4" style="background:{{period color|Pleistocene}}"| Pleistocene
|style="background:{{period color|Upper Pleistocene}}"| Late ''('Tarantian')''
|style="background:{{period color|upper Pleistocene}}"| {{Period start|Late pleistocene}}
|
|
|-
|style="background:{{period color|Middle Pleistocene}}"| Chibanian
|style="background:{{period color|Middle Pleistocene}}"| {{Period start|middle pleistocene}}
|
|
|-
|style="background:{{period color|Calabrian}}"| Calabrian
|style="background:{{period color|Calabrian}}"| {{Period start|calabrian}}
|
|
|-
|style="background:{{period color|Gelasian}}"| Gelasian
|style="background:{{period color|Gelasian}}"| {{Period start|gelasian}}
|
|
|-
|rowspan="8" style="background:{{period color|Neogene}}"| Neogene
|rowspan="2" style="background:{{period color|Pliocene}}"| [[Pliocene]]
|style="background:{{period color|Piacenzian}}"| Piacenzian
|style="background:{{period color|Piacenzian}}"| {{Period start|piacenzian}}
|{{FossilFormationsForAgeRange|3.6|.6}}
|{{FossilSpeciesForAgeRange|3.6|.6}}
|-
|style="background:{{period color|Zanclean}}"| Zanclean
|style="background:{{period color|Zanclean}}"| {{Period start|zanclean}}
|{{FossilFormationsForAgeRange|5.32|3.6}}
|{{FossilSpeciesForAgeRange|5.32|3.6}}
|-
|rowspan="6" style="background:{{period color|Miocene}}"| [[Miocene]]
|style="background:{{period color|Messinian}}"| Messinian
|style="background:{{period color|Messinian}}"| {{Period start|messinian}}
|{{FossilFormationsForAgeRange|7.3|5.3}}
|{{FossilSpeciesForAgeRange|7.3|5.3}}
|-
|style="background:{{period color|Tortonian}}"| Tortonian
|style="background:{{period color|Tortonian}}"| {{Period start|tortonian}}
|{{FossilFormationsForAgeRange|11.7|7.2}}
|{{FossilSpeciesForAgeRange|11.7|7.2}}
|-
|style="background:{{period color|Serravallian}}"| Serravallian
|style="background:{{period color|Serravallian}}"| {{Period start|serravallian}}
|{{FossilFormationsForAgeRange|13.9|11.6}}
|{{FossilSpeciesForAgeRange|13.9|11.6}}
|-
|style="background:{{period color|Langhian}}"| Langhian
|style="background:{{period color|Langhian}}"| {{Period start|langhian}}
|{{FossilFormationsForAgeRange|16.1|13.8}}
|{{FossilSpeciesForAgeRange|16.1|13.8}}
|-
|style="background:{{period color|Burdigalian}}"| Burdigalian
|style="background:{{period color|Burdigalian}}"| {{Period start|burdigalian}}
|{{FossilFormationsForAgeRange|20.5|15.97}}
|{{FossilSpeciesForAgeRange|20.5|15.97}}
|-
|style="background:{{period color|Aquitanian}}"| Aquitanian
|style="background:{{period color|Aquitanian}}"| {{Period start|aquitanian}}
|{{FossilFormationsForAgeRange|23.1|20.44}}
|{{FossilSpeciesForAgeRange|23.1|20.44}}
|-
|rowspan="9" style="background:{{period color|Paleogene}}"| Paleogene
|rowspan="2" style="background:{{period color|Oligocene}}"| [[Oligocene]]
|style="background:{{period color|Chattian}}"| Chattian
|style="background:{{period color|Chattian}}"| {{Period start|chattian}}
|{{FossilFormationsForAgeRange|28.2|23}}
|{{FossilSpeciesForAgeRange|28.2|23}}
|-
|style="background:{{period color|Rupelian}}"| Rupelian
|style="background:{{period color|Rupelian}}"| {{Period start|rupelian}}
|{{FossilFormationsForAgeRange|34|28.1}}
|{{FossilSpeciesForAgeRange|34|28.1}}
|-
|rowspan="4" style="background:{{period color|Eocene}}"| [[Eocene]]
|style="background:{{period color|Priabonian}}"| Priabonian
|style="background:{{period color|Priabonian}}"| {{Period start|priabonian}}
|{{FossilFormationsForAgeRange|37.9|33.9}}
|{{FossilSpeciesForAgeRange|37.9|33.9}}
|-
|style="background:{{period color|Bartonian}}"| Bartonian
|style="background:{{period color|Bartonian}}"| {{Period start|bartonian}}
|{{FossilFormationsForAgeRange|41.3|37.8}}
|{{FossilSpeciesForAgeRange|41.3|37.8}}
|-
|style="background:{{period color|Lutetian}}"| Lutetian
|style="background:{{period color|Lutetian}}"| {{Period start|lutetian}}
|{{FossilFormationsForAgeRange|47.9|41.2}}
|{{FossilSpeciesForAgeRange|47.9|41.2}}
|-
|style="background:{{period color|Ypresian}}"| Ypresian
|style="background:{{period color|Ypresian}}"| {{Period start|ypresian}}
|{{FossilFormationsForAgeRange|56.1|47.8}}
|{{FossilSpeciesForAgeRange|56.1|47.8}}
|-
|rowspan="3" style="background:{{period color|Paleocene}}"| [[Paleocene]]
|style="background:{{period color|Thanetian}}"| Thanetian
|style="background:{{period color|Thanetian}}"| {{Period start|thanetian}}
|{{FossilFormationsForAgeRange|59.3|56}}
|{{FossilSpeciesForAgeRange|59.3|56}}
|-
|style="background:{{period color|Selandian}}"| Selandian
|style="background:{{period color|Selandian}}"| {{Period start|selandian}}
|{{FossilFormationsForAgeRange|61.7|59.2}}
|{{FossilSpeciesForAgeRange|61.7|59.2}}
|-
|style="background:{{period color|Danian}}"| Danian
|style="background:{{period color|Danian}}"| {{Period start|danian}}
|{{FossilFormationsForAgeRange|66.1|61.6}}
|{{FossilSpeciesForAgeRange|66.1|61.6}}
|-
|rowspan="12" style="background:{{period color|Mesozoic}}"| Mesozoic
|rowspan="12" style="background:{{period color|Cretaceous}}" | [[Cretaceous]]
|rowspan="6" style="background:{{period color|Late Cretaceous}}"| Late
|style="background:{{period color|Maastrichtian}}"| Maastrichtian
|style="background:{{period color|Maastrichtian}}"| {{Period start|maastrichtian}} ±&nbsp;0.2
|{{FossilFormationsForAgeRange|72.2|66}}
|{{FossilSpeciesForAgeRange|72.2|66}}
|-
|style="background:{{period color|Campanian}}"| Campanian
|style="background:{{period color|Campanian}}"| {{Period start|campanian}} ±&nbsp;0.2
|{{FossilFormationsForAgeRange|83.7|72.1}}
|{{FossilSpeciesForAgeRange|83.7|72.1}}
|-
|style="background:{{period color|Santonian}}"| Santonian
|style="background:{{period color|Santonian}}"| {{Period start|santonian}} ±&nbsp;0.5
|{{FossilFormationsForAgeRange|86.4|83.6}}
|{{FossilSpeciesForAgeRange|86.4|83.6}}
|-
|style="background:{{period color|Coniacian}}"| Coniacian
|style="background:{{period color|Coniacian}}"| {{Period start|coniacian}} ±&nbsp;0.3
|{{FossilFormationsForAgeRange|89.9|86.3}}
|{{FossilSpeciesForAgeRange|89.9|86.3}}
|-
|style="background:{{period color|Turonian}}"| Turonian
|style="background:{{period color|Turonian}}"| {{Period start|turonian}}
|{{FossilFormationsForAgeRange|94|89.9}}
|{{FossilSpeciesForAgeRange|94|89.8}}
|-
|style="background:{{period color|Cenomanian}}"| Cenomanian
|style="background:{{period color|Cenomanian}}"| {{Period start|cenomanian}}
|{{FossilFormationsForAgeRange|100.6|93.9}}
|{{FossilSpeciesForAgeRange|100.6|93.9}}
|-
|rowspan="6" style="background:{{period color|Early Cretaceous}}"| Early
|style="background:{{period color|Albian}}"| Albian
|style="background:{{period color|Albian}}"| ~{{Period start|albian}}
|{{FossilFormationsForAgeRange|113.1|100.5}}
|{{FossilSpeciesForAgeRange|113.1|100.5}}
|-
|style="background:{{period color|Aptian}}"| Aptian
|style="background:{{period color|Aptian}}"| ~{{Period start|aptian}}
|{{FossilFormationsForAgeRange|125.1|113}}
|{{FossilSpeciesForAgeRange|125.1|113}}
|-
|style="background:{{period color|Barremian}}"| Barremian
|style="background:{{period color|Barremian}}"| ~{{Period start|barremian}}
|{{FossilFormationsForAgeRange|129.5|125}}
|{{FossilSpeciesForAgeRange|129.5|125}}
|-
|style="background:{{period color|Hauterivian}}"| Hauterivian
|style="background:{{period color|Hauterivian}}"| ~{{Period start|hauterivian}}
|{{FossilFormationsForAgeRange|133|129.4}}
|{{FossilSpeciesForAgeRange|133|129.4}}
|-
|style="background:{{period color|Valanginian}}"| Valanginian
|style="background:{{period color|Valanginian}}"| ~{{Period start|valanginian}}
|{{FossilFormationsForAgeRange|140|132.9}}
|{{FossilSpeciesForAgeRange|140|132.9}}
|-
|style="background:{{period color|Berriasian}}"| Berriasian
|style="background:{{period color|Berriasian}}"| ~{{Period start|berriasian}}
|{{FossilFormationsForAgeRange|145.1|139.8}}
|{{FossilSpeciesForAgeRange|145.1|139.8}}
|-
|}

==Subfamily-level Phylogeny with Fossil Occurrences==
{{Photo Gallery
|noheading=yes
|name1=Barden 2017, Fig. 3.jpg
|comment1=A dated subfamily-level phylogeny with first confident fossil occurrences for each lineage, last occurrences indicated for extinct lineages. Fossil deposit numbers correspond with Table 2, Figure 4, and Figure 5. Tree topology as well as crown and divergence dates from MOREAU & BELL (2013), which is so far the largest Formicidae-wide analysis with respect to taxon sampling. No crown age included for Aneuretinae, Martialis, and Paraponerinae as these are monotypic. Agroecomyrmecinae includes no crown age due to insufficient terminal sampling. Lineages with dotted lines were not placed directly through analyses, but rather added to the molecule-derived topology of Moreau & Bell, which did not include fossils. Stem ants and Brownimecia were placed according to conservative results of BARDEN & GRIMALDI (2016) – Note that while Sphecomyrminae and stem group ants are depicted on a single lineage, this group is not necessarily monophyletic. Formiciinae placement is based on WARD (2007), although this position was not recovered in the only two phylogenetic treatments of the subfamily (BARONI URBANI & al. 1992, GRIMALDI & al. 1997), it seems plausible to this author (Barden, 2017, Fig. 3).
|size1=1103px
}}

==Fossil Formations Known to Contain Ants==
{{#cargo_query:
tables=FossilFormation, FossilOccurrence, TaxonName
|join on=FossilFormation.Formation=FossilOccurrence.Formation, FossilOccurrence._pageName=TaxonName._pageName
|fields= CONCAT("[[", FossilFormation._pageName, "]]")=Formation, count(TaxonName.Species)=No. of Species, FossilFormation.Start=Start Age, FossilFormation.End=End Age, FossilFormation.Period, FossilFormation.Epoch, FossilFormation.Age, FossilFormation.Locality, FossilFormation.Country, FossilFormation.Coordinates, FossilFormation.Paleocoordinates, FossilFormation.Source, FossilFormation.Notes
|where=TaxonName.Species <> ""
|order by=FossilFormation._pageName
|group by=FossilOccurrence.Formation
}}

==References==
*[[Media:Barden, P. 2017. Fossil ants ancient diversity and the rise of modern lineages.pdf|Barden, P. 2017. Fossil ants (Hymenoptera: Formicidae): ancient diversity and the rise of modern lineages. Myrmecological News 24: 1-30.]]
*Bjorn Berning & Sigitas Podenas 2009.  Amber: Archive of Deep Time. Denisia 26, zugleich Kataloge de Oberösterreichischen Landesmuseen N.S. 86. 294 pages.
*DuBois, M. B.; LaPolla, J. S. 1999. A preliminary review of Colombian ants (Hymenoptera: Formicidae) preserved in copal. Entomol. News 110: 162-172
*Grimaldi, D. and M. S. Engel, Evolution of the Insects Cambridge University Press, Cambridge, 2005, xv+755 pp
*Krynicki, V. E. 2013. Primitive ants (Hymenoptera: Sphecomyrminae) in the Campanian (Late Cretaceous) of North Carolina (USA). Life: The Excitement of Biology 1:156-165.
*[[Media:LaPolla et al. - 2013 - Ants and the fossil record.pdf|LaPolla, J. S.; Dlussky, G. M.; Perrichot, V. 2013. Ants and the fossil record. Annual Review of Entomology 58:609-630.]]
*[[Media:LaPolla, J.S., Greenwlalt, D.E. 2015. Fossil ants of the Middle Eocene Kishenehn Formation.pdf|LaPolla, J.S.,  Greenwalt, D.E. 2015. Fossil Ants (Hymenoptera: Formicidae) of the Middle Eocene Kishenehn Formation. Sociobiology. 62: 163-174]] ({{doi|10.13102/sociobiology.v62i2.163-174}}).
*[http://homepage.mac.com/paulselden/Sites/Website/PNAS2010.pdf] Schmidt AR, Perrichot V, Svojtka M, Anderson KB, Belete KH, Bussert R, Dörfelt H, Jancke S, Mohr B, Mohrmann E, Nascimbene PC, Nel A, Nel P, Ragazzi E, Roghi G, Saupe EE, Schmidt K, Schneider H, Selden PA, Vávra N. 2010. Cretaceous African life captured in amber. Proc Natl Acad Sci U S A. 20;107(16):7329-34.
*[http://gap.entclub.org/taxonomists/Wilson/Mesozoic%20ants%20subfamily.pdf] Wilson, E. O.; Carpenter, F. M.; Brown, W. L., Jr. 1967a. The first Mesozoic ants, with the description of a new subfamily. Psyche (Camb.) 74: 1-19