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Agroecomyrmecinae is a
subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zo ...
of
ant 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 Cre ...
s containing two extant and two fossil genera. The subfamily was originally classified in 1930 by Frank M. Carpenter as Agroecomyrmecini, a
Myrmicinae Myrmicinae is a subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily ...
tribe.
Bolton Bolton ( , locally ) is a town in Greater Manchester in England. In the foothills of the West Pennine Moors, Bolton is between Manchester, Blackburn, Wigan, Bury, Greater Manchester, Bury and Salford. It is surrounded by several towns and vill ...
raised the tribe to subfamily status in 2003, suggesting that Agroecomyrmecinae might be the sister taxon to Myrmicinae. It has since been discovered to be one of the earliest lineages of ants, a
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
from the basal polytomy for all ants. In 2014, the subfamily was expanded to two tribes. The tribe
Ankylomyrmini ''Ankylomyrma'', from Ancient Greek ἀγκύλος (''ankúlos''), meaning "curved", and μύρμηξ (''múrmēx''), meaning "ant", is a genus of large arboreal ants in the subfamily Agroecomyrmecinae. It contains the single species ''Ankylomyr ...
was moved from the subfamily
Myrmicinae Myrmicinae is a subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily ...
to Agroemyrmecinae.


Tribes and genera

*Agroecomyrmecinae Carpenter, 1930 ** Agroecomyrmecini Carpenter, 1930 *** †''
Agroecomyrmex ''Agroecomyrmex'' is an extinct genus of ants in the formicid subfamily Agroecomyrmecinae, for which it is the type genus. The genus contains a single described species, ''Agroecomyrmex duisburgi''. ''Agroecomyrmex'' is known from a group of M ...
'' Wheeler, 1910 **** †'' Agroecomyrmex duisburgi'' Wheeler, 1910 *** †''
Eulithomyrmex ''Eulithomyrmex'' is an extinct genus of ant in the formicid subfamily Agroecomyrmecinae. The genus contains two described species, ''Eulithomyrmex rugosus'' and ''Eulithomyrmex striatus''. ''Eulithomyrmex'' is known from a group of Late Eocene ...
'' Carpenter, 1935 ****†''Eulithomyrmex rugosus'' Carpenter, 1930 ****†''Eulithomyrmex striatus'' Carpenter, 1930 *** ''
Tatuidris ''Tatuidris'', or armadillo ant, is a rare genus of ants consisting of a single species, ''Tatuidris tatusia''. The ants are small in size and inhabit the leaf litter of Neotropical forests in Central and South America, from Mexico to Brazil ...
'' Brown & Kempf, 1968 **** ''
Tatuidris tatusia ''Tatuidris'', or armadillo ant, is a rare genus of ants consisting of a single species, ''Tatuidris tatusia''. The ants are small in size and inhabit the leaf litter of Neotropical forests in Central and South America, from Mexico to Brazil ...
'' Brown & Kempf, 1968 (=''T. kapasi'' Lacau & Groc, 2012) **
Ankylomyrmini ''Ankylomyrma'', from Ancient Greek ἀγκύλος (''ankúlos''), meaning "curved", and μύρμηξ (''múrmēx''), meaning "ant", is a genus of large arboreal ants in the subfamily Agroecomyrmecinae. It contains the single species ''Ankylomyr ...
***''
Ankylomyrma ''Ankylomyrma'', from Ancient Greek ἀγκύλος (''ankúlos''), meaning "curved", and μύρμηξ (''múrmēx''), meaning "ant", is a genus of large arboreal ants in the subfamily Agroecomyrmecinae. It contains the single species ''Ankylomyr ...
'' Bolton, 1973 ****'' Ankylomyrma coronacantha'' Bolton, 1973


Taxonomy

Since the original description, the systematic status of the Agroecomyrmecini tribe has been the focus of intense debate. Bolton (2003) was the first to suggest the taxonomic instability of ''Tatuidris'' within Myrmicinae and raised the genus to the level of a new subfamily, the Agroecomyrmecinae, suggesting the Agroecomyrmecinae might be the sister taxon to Myrmicinae. This assessment was based on these diagnostic characters: # large mandibles with mandibular masticatory margins that oppose at full closure but do not overlap # eyes at extreme posterior apex of deep antennal scrobes # clypeus very broadly triangular, broadly inserted between the frontal lobes # antennal sockets and frontal lobes strongly migrated laterally, far apart and close to lateral margins of the head # mesotibia and metatibia with pectinate spurs # short and compact mesosoma # a sessile petiole, in posterior view the
tergite A ''tergum'' (Latin for "the back"; : ''terga'', associated adjective tergal) is the Anatomical terms of location#Dorsal and ventral, dorsal ('upper') portion of an arthropod segment other than the head. The Anatomical terms of location#Anterior ...
and sternite not equally convex # an abdominal segment III (postpetiole) without tergosternal fusion, segment large and very broadly articulated to segment IV, # a helcium in frontal view with the sternite bulging ventrally and overlapped by the tergite # an abdominal segment IV with a complete tergosternal fusion, # abdominal segment IV with a stridulitrum on the pretergite # the sternite of abdominal segment IV is reduced, the tergite is much larger than the sternite and strongly vaulted The subfamily rank of the armadillo ants was reassessed by Baroni Urbani & de Andrade (2007) in their last systematic assessment of the dacetines. They analyzed a morphological dataset that included former dacetines, basicerotines, phalacromyrmecines, and ''Tatuidris'', as well as other non-Myrmicinae taxa such as the Australian genus '' Myrmecia'' and the Neotropical genus '' Pseudomyrmex''. This work was the first attempt to include ''Tatuidris'' as a terminal taxon in a morphological cladistic analysis. In their study, Baroni Urbani & de Andrade (2007) identified six morphological
synapomorphies In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to ...
shared between ''Tatuidris'' and the dacetines, justifying the inclusion of the genus within Myrmicinae. These characters included: # mandibles at rest opposing at least in part, instead of crossing # a mandibular-torular index < 130 # reduction of maxillary palps from double-jointed to single-jointed # reduced male mandibles # presence of a two-segmented antennal club # reduced number of antennal joints In addition, two autapomorphies (a differently shaped petiolar
tergum A ''tergum'' (Latin for "the back"; : ''terga'', associated adjective tergal) is the dorsal ('upper') portion of an arthropod Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton wi ...
and
sternum The sternum (: sternums or sterna) or breastbone is a long flat bone located in the central part of the chest. It connects to the ribs via cartilage and forms the front of the rib cage, thus helping to protect the heart, lungs, and major bl ...
, and the eyes at or close to the apex of the antennal scrobe) separated ''Tatuidris'' from all other extant ant genera included in their study. Unlike
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
studies based on morphological traits, molecular analyses of the internal phylogeny of the ants have given strong evidence that the armadillo ants are neither closely related to nor nested within the Myrmicinae. Brady ''et al''. (2006), Moreau ''et al''. (2006) and Rabeling ''et al''. (2008) reconstructed phylogenetic trees with the agroecomyrmecines inside the 'poneroid' group of subfamilies, close to the
Paraponerinae ''Paraponera'' is a genus of ants and the only genus in the subfamily Paraponerinae. The name means "near-'' Ponera''". It consists of two species: the extant '' Paraponera clavata'', also known as a bullet ant, found in the Neotropics, and the ...
, and gave support for the exclusion of the genus from the Myrmicinae, a subfamily located inside the 'formicoid' clade. Given the early appearance of the Agroecomyrmecinae in the geologic record, the similarities of armadillo ants to Myrmicinae were hypothesized to represent convergence and/or retention of
plesiomorphic In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorphy, apomorphy, an ...
forms. Recently, Keller (2011) challenged the phylogenetic relationships of the poneromorph subfamilies (including ''Tatuidris'').


Distribution

According to Brown & Kempf (1967), agroecomyrmecines were probably widespread in both hemispheres during the early
Tertiary Tertiary (from Latin, meaning 'third' or 'of the third degree/order..') may refer to: * Tertiary period, an obsolete geologic period spanning from 66 to 2.6 million years ago * Tertiary (chemistry), a term describing bonding patterns in organic ch ...
. ''Agroecomyrmex'' is known from Early Eocene,
Lutetian The Lutetian is, in the geologic timescale, a stage (stratigraphy), stage or age (geology), age in the Eocene. It spans the time between . The Lutetian is preceded by the Ypresian and is followed by the Bartonian. Together with the Bartonian it ...
,
Baltic amber Baltic amber or succinite is amber from the Baltic region, home of its largest known deposits. It was produced sometime during the Eocene epoch, but exactly when is controversial. It has been estimated that this forested region provided the re ...
dating to 44 million years (
Myr Million years ago, abbreviated as Mya, Myr (megayear) or Ma (megaannum), is a unit of time equal to (i.e. years), or approximately 31.6 teraseconds. Usage Myr is in common use in fields such as Earth science and cosmology. Myr is also used w ...
) ago, and ''Eulithomyrmex'' from
late Eocene The Priabonian is, in the International Commission on Stratigraphy, ICS's geologic timescale, the latest age (geology), age or the upper stage (stratigraphy), stage of the Eocene epoch (geology), Epoch or series (stratigraphy), Series. It spans ...
,
Priabonian The Priabonian is, in the ICS's geologic timescale, the latest age or the upper stage of the Eocene Epoch or Series. It spans the time between . The Priabonian is preceded by the Bartonian and is followed by the Rupelian, the lowest stage ...
, Florissant shale (34.1 Myr ago) in present-day
Colorado Colorado is a U.S. state, state in the Western United States. It is one of the Mountain states, sharing the Four Corners region with Arizona, New Mexico, and Utah. It is also bordered by Wyoming to the north, Nebraska to the northeast, Kansas ...
,
United States The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...
. ''Tatuidris'', rare but broadly distributed, inhabits the leaf litter of
Neotropical The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropical terrestrial ecoregions of the Americas and the entire South American temperate zone. Definition In biogeogra ...
forests in Central and South America, from
Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin America, and borders the United States to the north, and Guatemala and Belize to the southeast; while having maritime boundar ...
to
French Guiana French Guiana, or Guyane in French, is an Overseas departments and regions of France, overseas department and region of France located on the northern coast of South America in the Guianas and the West Indies. Bordered by Suriname to the west ...
, central
Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...
, and Amazonian Peru. ''Ankylomyrma'' is known only from Western Africa.


Notes


References

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External links

* * {{Taxonbar, from=Q4033595 Ant subfamilies Extant Lutetian first appearances