Armaniidae was a name formerly given to a group of extinct

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 ...

-like

hymenopteran Hymenoptera is a large order of insects, comprising the sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are parasitic. Females typica ...

s known from a series of

Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...

fossils found in Asia and Africa. Armaniidae has been suggested by several authors to belong to the family Formicidae as one of the

stem-group In phylogenetics, the crown group or crown assemblage is a collection of species composed of the living representatives of the collection, the most recent common ancestor of the collection, and all descendants of the most recent common ancestor. ...

subfamilies, Armaniinae; however, this position has not been taken up by myrmecologists. The family contained seven described genera and thirteen described species. An analysis of fossil ants based on antenna structure in 2017 resulted in three of the family being broken up with some genera being moved to

Sphecomyrminae Sphecomyrminae is an extinct subfamily of ants in family Formicidae known from a series of Cretaceous fossils found in North America, Europe, and Asia. Sphecomyrminae contains eight genera, divided into two tribes Sphecomyrmini and Zigrasimec ...

and other genera being relegated to ''

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 ...

'' in Formicidae or

Aculeata Aculeata is a subclade of Hymenoptera containing ants, bees, and stinging wasps. The name is a reference to the defining feature of the group, which is the modification of the ovipositor into a stinger. However, many members of the group canno ...

History and classification

Armaniidae was sometimes treated as the most basal of the Formicidae subfamilies, and classed as a stem-group which is more distant in relation to modern ants than the next stem group,

. More often the group, treated as "ant-like wasps", was elevated to the rank of family, and considered as a possible sister group to Formicidae. It has been suggested by Engel and Grimaldi that the group may be paraphyletic. This position is in contrast to the original hypothesis of Russian paleoentomologist Gennady Dlussky, who first described the family. Dlussky considered the group, when erected in 1983, have been an intermediate family bridging the families

Scoliidae The Scoliidae, the scoliid wasps, are a family of about 560 species found worldwide. They tend to be black, often marked with yellow or orange, and their wing tips are distinctively corrugated. Males are more slender and elongated than females, wi ...

and the true formicids. In contrast to both the treatment as a separate family and as a distinct subfamily, entomologist

E. O. Wilson Edward Osborne Wilson (June 10, 1929 – December 26, 2021) was an American biologist, naturalist, entomologist and writer. According to David Attenborough, Wilson was the world's leading expert in his specialty of myrmecology, the study of an ...

, in a 1987 paper, suggested that the then known armaniids and '' Sphecomyrma'' represented a single species. Wilson, in synonymizing the groups, made the hypothesis that the different described genera were actually fossils of different castes of the same species, with ''Sphecomyrma freyi'' being workers, ''Armania robusta'' being queens, and '' "Paleomyrmex" zherichini'' as the winged males. This view was rejected as new fossils and species were described. The group is known exclusively from impression fossils, which have a limited preservation quality, leading to the uncertainty of what features are present in the described species. Overall armaniids have a poorly developed petiole which is broadly attached to the

thorax The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the cre ...

, short scapes on the antennae, and queen-like looking females. The mandibles are

vespid The Vespidae are a large (nearly 5000 species), diverse, cosmopolitan family of wasps, including nearly all the known eusocial wasps (such as ''Polistes fuscatus'', ''Vespa orientalis'', and ''Vespula germanica'') and many solitary wasps. Each ...

like, with possibly only one or two teeth, though this may be an artifact of preservation. The short scape is a feature that is also seen in Sphecomyrminae members, and does not exclude armaniids from Formicidae. Similarly the petiole is a feature that is seen in both the true formicids, and in the extinct chrysidoid wasp family Falsiformicidae, which is not related to formicids at all. The two defining features of the true formicids are considered to be the presence of females which are divided into adult workers and queens. Currently no worker like armaniid specimens are known for the described species. The presence of

metapleural gland Metapleural glands (also called metasternal or metathoracic glands) are secretory glands that are unique to ants and basal in the evolutionary history of ants. They are responsible for the production of an antibiotic fluid that then collects in a ...

s in some fossils has been reported by Dlussky,± but the veracity of the presence is uncertain. A review of the Cretaceous ant groups was performed by Borysenko in 2017 leading to the breakup of Armaniidae, with the three genera '' Armania'', '' Orapia'', and '' Pseudarmania'' being moved to

. The genera '' Archaeopone'' and '' Poneropterus'' were considered as ''incertae sedis'' in Formicidae, and the genera '' Dolichomyrma'' and '' Khetania'' were removed from the family entirely as ''incertae sedis'' in

Genera and species

*'' Archaeopone'' Dlussky, 1975 **''Archaeopone kzylzharica'' Dlussky, 1975 **''Archaeopone taylori'' Dlussky, 1983 *'' Armania'' Dlussky, 1983 (jr synonym = "''Armaniella''" Dlussky, 1983) **''Armania capitata'' Dlussky, 1983 **''Armania curiosa'' (Dlussky, 1983) **''Armania pristina'' Dlussky, 1983 **''Armania robusta'' Dlussky, 1983 *'' Dolichomyrma'' Dlussky, 1975 **''Dolichomyrma latipes'' Dlussky, 1975 **''Dolichomyrma longiceps'' Dlussky, 1975 *'' Khetania'' Dlussky, 1999 **''Khetania mandibulata'' Dlussky, 1999 *'' Orapia'' Dlussky, Brothers & Rasnitsyn, 2004 **''Orapia minor'' Dlussky, Brothers & Rasnitsyn, 2004 **''Orapia rayneri'' Dlussky, Brothers & Rasnitsyn, 2004 *'' Poneropterus'' Dlussky, 1983 **''Poneropterus sphecoides'' Dlussky, 1983 *'' Pseudarmania'' Dlussky, 1983 **''Pseudarmania aberrans'' Dlussky, 1983 **''Pseudarmania rasnitsyni'' Dlussky, 1983

References

{{Taxonbar, from=Q4034379 † Cretaceous insects Fossil ant taxa Cenomanian first appearances Turonian extinctions