Ants (

family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...

Formicidae in the

order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...

Hymenoptera) are the most species-rich of all social insects, with more than 12,000 described

species In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...

and many others awaiting description. Formicidae is divided into 21 subfamilies, of which 17 contain extant

taxa In biology, a taxon ( back-formation from '' taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular n ...

, while four are exclusively

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 ...

. Ants have come to occupy virtually all major terrestrial habitats, with the exception of tundra and cold ever-wet forests. They display a wide range of social behaviors, foraging habits and associations with other organisms, which has generated scientific and public interest.

Clades

Beginning in the 1990s, molecular ( DNA sequence) data have come to play a central role in attempts to reconstruct the ant "

tree of life The tree of life is a fundamental archetype in many of the world's mythological, religious, and philosophical traditions. It is closely related to the concept of the sacred tree.Giovino, Mariana (2007). ''The Assyrian Sacred Tree: A Histo ...

Molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...

analyses based on multiple

nuclear gene A nuclear gene is a gene whose physical DNA nucleotide sequence is located in the cell nucleus of a eukaryote. The term is used to distinguish nuclear genes from genes found in mitochondria or chloroplasts. The vast majority of genes in eukaryot ...

s have yielded robust results that reinforce some preexisting views but overturn others – and suggest that there has been considerable morphological convergence among some ant lineages. Molecular data provide very strong support for a novel group, the "formicoid clade", not revealed by previous morphological work. This clade comprises 9 of the 16 extant ant subfamilies and about 90% of all described ant species. Formicoids include such widespread and species-rich subfamilies as

Myrmicinae Myrmicinae is a subfamily of ants, with about 140 extant genera; their distribution is cosmopolitan. The pupae lack cocoons. Some species retain a functional sting. The petioles of Myrmicinae consist of two nodes. The nests are permanent an ...

, Formicinae and Dolichoderinae, as well as the

army ant The name army ant (or legionary ant or ''marabunta'') is applied to over 200 ant species in different lineages. Because of their aggressive predatory foraging groups, known as "raids", a huge number of ants forage simultaneously over a limi ...

s ( Dorylinae). Non-formicoids comprise five "poneroid" subfamilies ( Agroecomyrmecinae, Amblyoponinae, Paraponerinae, Ponerinae, and Proceratiinae), Leptanillinae, about which little is known, and Martialinae, the most recently discovered subfamily. Relationships among these remaining seven subfamilies are less well resolved. A recent study (2011) places Leptanillinae as a

sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...

to all other ants, with Martialinae, the poneroids and formicoids forming a clade.

Evolution of ants

Ants first arose during the mid-Cretaceous, more than 100 million years ago, associated with the rise of flowering plants and an increase in forest ground litter. The earliest known ants evolved from a lineage within the aculeate wasps, and a recent study suggests that they are a sister group of

Apoidea The superfamily Apoidea is a major group within the Hymenoptera, which includes two traditionally recognized lineages, the "sphecoid" wasps, and the bees. Molecular phylogeny demonstrates that the bees arose from within the traditional "Crabronida ...

. During the Cretaceous ants were confined to the northern

Laurasia Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pan ...

n supercontinent, with only a few widespread primitive species. By the middle Eocene, around 50 million years ago, ants had diversified and become ecologically dominant as predators and scavengers. Ant species are less than 2% of the total number of insect species but make up one third of the insect biomass.

History of classification

In volume 1 of ''

Systema Naturae ' (originally in Latin written ' with the ligature æ) is one of the major works of the Swedish botanist, zoologist and physician Carl Linnaeus (1707–1778) and introduced the Linnaean taxonomy. Although the system, now known as binomial nom ...

'',

Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, ...

(1758) described seventeen species of ants, all of which he placed in the single

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial n ...

'' Formica''. Within a few decades additional genera had been recognized, and this trend continued in the ensuing years, together with the development of a more complex hierarchical classification in which genera were apportioned among subfamilies and

tribes The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English is in the discipline of anthropology. This definition is contested, in part due to confl ...

. The ant species described by Linnaeus are now dispersed in eleven different genera, belonging to four subfamilies. For much of the twentieth century the number of recognized ant subfamilies varied from seven to ten, with the Aneuretinae, Cerapachyinae, Leptanillinae, Myrmeciinae and Pseudomyrmecinae being variously treated as separate subfamilies or (at different times) subsumed within Dolichoderinae, Ponerinae, Dorylinae, Ponerinae, and Myrmicinae, respectively. In 2014, Brady et al. synonymized the army ant subfamilies and their closest relatives under Dorylinae; this clade, the dorylomorph subfamilies, previously also contained Aenictinae, Aenictogitoninae, Cerapachyinae, Ecitoninae and Leptanilloidinae. The last three decades have seen a proliferation of subfamily names, as a result of three factors: (1) the realization that some subfamilies were assemblages of unrelated taxa; (2) abandonment of

paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...

taxa, and (3) the discovery of novel fossil taxa. Seventeen extant subfamilies of ants are currently recognized, along with four extinct subfamilies. One of the fossil taxa, Armaniinae, is often given

rank within the superfamily Formicoidea. About 13 genera are ''

incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertain ...

'' (of uncertain placement), and are not assigned to any subfamily.

Subfamilies

Extinct

are indicated by a

† A dagger, obelisk, or obelus is a typographical mark that usually indicates a footnote if an asterisk has already been used. The symbol is also used to indicate death (of people) or extinction (of species). It is one of the modern descendan ...

Notes

References

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Ants Ants are Eusociality, eusocial insects of the Family (biology), family Formicidae and, along with the related wasps and bees, belong to the Taxonomy (biology), order Hymenoptera. Ants evolved from Vespoidea, vespoid wasp ancestors in the Creta ...

Subfamilies Subfamilies