Agroecomyrmecinae

Life-forms

Superregnum: Eukaryota
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Subphylum: Hexapoda
Classis: Insecta
Cladus: Dicondylia
Subclassis: Pterygota
Cladus: Metapterygota
Infraclassis: Neoptera
Cladus: Eumetabola
Cladus: Endopterygota
Superordo: Hymenopterida
Ordo: Hymenoptera
Subordo: Apocrita
Superfamilia: Formicoidea

Familia: Formicidae
Subfamilia: Agroecomyrmecinae
Genera: †Agroecomyrmex – Ankylomyrma – †Eulithomyrmex – Tatuidris
Name

Agroecomyrmecinae Carpenter, 1930
References
Carpenter, F. M. 1930. The fossil ants of North America.. Bulletin of the Museum of Comparative Zoology, 70: 1–66.

Agroecomyrmecinae is a subfamily of ants containing two extant and two fossil genera.[1] The subfamily was originally classified in 1930 by Carpenter as Agroecomyrmecini, a Myrmicinae tribe.[2] Bolton 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 from the basal polytomy for all ants.[3][4] In 2014, the subfamily was expanded to two tribes. The tribe Ankylomyrmini was moved from the subfamily Myrmicinae to Agroemyrmecinae.[5]

Tribes and genera

Agroecomyrmecinae Carpenter, 1930
Agroecomyrmecini Carpenter, 1930
†Agroecomyrmex Wheeler, 1910
†Agroecomyrmex duisburgi Wheeler, 1910
†Eulithomyrmex Carpenter, 1935
†Eulithomyrmex rugosus Carpenter, 1930
†Eulithomyrmex striatus Carpenter, 1930
Tatuidris Brown & Kempf, 1968
Tatuidris tatusia Brown & Kempf, 1968 (=T. kapasi Lacau & Groc, 2012)[6]
Ankylomyrmini
Ankylomyrma 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:[7][8]

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 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,[note 1]
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 shared between Tatuidris and the dacetines, justifying the inclusion of the genus within Myrmicinae. These characters included:[9][10]

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 and sternum, 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.[9][11]

Unlike phylogenetic 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, and gave support for the exclusion of the genus from the Myrmicinae, a subfamily located inside the 'formicoid' clade.[12] 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 forms.[13][14]

Recently, Keller (2011) challenged the phylogenetic relationships of the poneromorph subfamilies (including Tatuidris).[14][15]
Distribution

According to Brown & Kempf (1967), agroecomyrmecines were probably widespread in both hemispheres during the early Tertiary.[16] Agroecomyrmex is known from Early Eocene, Lutetian, Baltic amber dating to 44 million years (Myr) ago, and Eulithomyrmex from late Eocene, Priabonian, Florissant shale (34.1 Myr ago) in present-day Colorado, United States.

Tatuidris, rare but broadly distributed,[14] inhabits the leaf litter of Neotropical forests in Central and South America, from Mexico to French Guiana,[17] central Brazil,[18] and Amazonian Peru.[6] Ankylomyrma is known only from Western Africa.[19]
Notes

This character was described incorrectly by Bolton (l.c.); in Tatuidris the tergosternal suture of the abdominal segment IV is strong but not fused.[9]

References

"Subfamily: Agroecomyrmecinae". antweb.org. AntWeb. Retrieved 3 January 2015.
Carpenter 1930
Ward 2007
"Genus: Tatuidris". antweb.org. AntWeb. Retrieved 29 August 2013.
Ward et al. 2014
Donoso 2012, p. 61
Bolton 2003, p. 51
Donoso 2012, pp. 61–62
Donoso 2012, p. 62
Baroni Urbani & de Andrade 2007, p. 78
Baroni Urbani & de Andrade 2007, pp. 80–81
Ward 2007, pp. 555–557
Ward 2011, p. 23
Donoso 2012, p. 63
Keller 2011, p. 73
Brown & Kempf 1967, p. 186
Lacau et al. 2012, p. 4
Vasconcelos & Vilhena 2002, p. 278

Bolton 1981

Baroni Urbani, C.; de Andrade, M.L. (2007), "The ant tribe Dacetini: Limits and constituent genera, with descriptions of new species", Annali del Museo Civico di Storia Naturale "G. Doria", 99: 1–191
Bolton, B. (1981), "A revision of six minor genera of Myrmicinae (Hymenoptera: Formicidae) in the Ethiopian zoogeographical region", Bulletin of the British Museum (Natural History), Entomology, 43 (4): 245–307, doi:10.15468/uu3l6q
Bolton, B. (2003), Synopsis and classification of Formicidae, Memoirs of the American Entomological Institute, vol. 71, The American Entomological Institute, pp. 1–370
Brady, S.G.; Schultz, T.R.; Fisher, B.L.; Ward, P.S. (2006), "Evaluating alternative hypotheses for the early evolution and diversification of ants", Proceedings of the National Academy of Sciences, 103 (48): 18172–18177, doi:10.1073/pnas.0605858103, PMC 1838725, PMID 17079492
Brown, W. L., Jr.; Kempf, W. W. (1967), "Tatuidris, a remarkable new genus of Formicidae (Hymenoptera)" (PDF), Psyche, 74 (3): 183–190, doi:10.5281/zenodo.27017
Carpenter, F.M. (January 1930), "The fossil ants of North America" (PDF), Bulletin of the Museum of Comparative Zoology, 70 (1): 1–66
Donoso, D.A. (2012), "Additions to the taxonomy of the armadillo ants (Hymenoptera, Formicidae, Tatuidris)" (PDF), Zootaxa, 3503: 61–81, doi:10.11646/zootaxa.3503.1.5
Keller, R.A. (2011), "A phylogenetic analysis of ant morphology (Hymenoptera: Formicidae) with special reference to the poneromorph subfamilies" (PDF), Bulletin of the American Museum of Natural History, 355: 1–90, doi:10.1206/355.1, hdl:2246/6124, S2CID 84005152
Lacau, Sébastien; Groc, Sarah; Dejean, Alain; Oliveira, Muriel L. de; Delabie, Jacques H. C. (2012), "Tatuidris kapasi sp. nov.: a new armadillo ant from French Guiana (Formicidae: Agroecomyrmecinae)", Psyche, 2012: 1–6, doi:10.1155/2012/926089
Moreau, C.S.; Bell, C.D.; Vila, R.; Archibald, S.B.; Pierce, N.E. (2006), "Phylogeny of the ants: diversification in the age of angiosperms", Science, 312 (5770): 101–104, Bibcode:2006Sci...312..101M, doi:10.1126/science.1124891, PMID 16601190, S2CID 20729380
Rabeling, C.; Brown, J.M.; Verhaagh, M. (2008), "Newly discovered sister lineage sheds light on early ant evolution", Proceedings of the National Academy of Sciences of the United States of America, 105 (39): 14913–14917, Bibcode:2008PNAS..10514913R, doi:10.1073/pnas.0806187105, PMC 2567467, PMID 18794530
Vasconcelos, Heraldo L.; Vilhena, José M.S. (2002), "First record of the ant genus Tatuidris (Hymenoptera: Formicidae) in Brazil" (PDF), Revista de Biología Tropical, 51 (1): 278
Ward, P.S (2007), "Phylogeny, classification, and species-level taxonomy of ants (Hymenoptera: Formicidae)", Zootaxa, 1668: 549–563, doi:10.11646/zootaxa.1668.1.26
Ward, P.S (2011), "Integrating molecular phylogenetic results into ant taxonomy (Hymenoptera: Formicidae)", Myrmecological News, 15: 21–29
Ward, P.S.; Brady, S.G.; Fisher, B.L.; Schultz, T.R. (July 2014), "The evolution of myrmicine ants: phylogeny and biogeography of a hyperdiverse ant clade (Hymenoptera: Formicidae)", Systematic Entomology, 40 (1): 61–81, doi:10.1111/syen.12090, ISSN 1365-3113, S2CID 83986771

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