Agroecomyrmecinae is a subfamily of ants consisting of a single extant species, Tatuidris tatusia, found in Central and South America, and two fossil genera: Agroecomyrmex and Eulithomyrmex. The subfamily was originally classified in 1930 by Carpenter as Agroecomyrmecini, a Myrmicinae tribe. 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.
Tribes and genera
- Agroecomyrmecinae Carpenter, 1930
- Agroecomyrmecini Carpenter, 1930
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 that Agroecomyrmecinae might be the sister taxon to Myrmicinae. This assessment was based on the following 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 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, and
- 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 re-assessed 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:
- mandibles at rest opposing at least in part, instead of crossing,
- a mandibular-torular index < 130,
- reduction of maxillary palps from 2-jointed to 1-jointed,
- reduced male mandibles,
- presence of a two-segmented antennal club, and
- a 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.
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. 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.
According to Brown & Kempf (1968), agroecomyrmecines were probably widespread in both hemispheres during the early Tertiary. 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, inhabits the leaf litter of Neotropical forests in Central and South America, from Mexico to French Guiana, central Brazil, and Amazonian Peru.
- Carpenter, F. M. 1930. The fossil ants of North America. Bulletin of the Museum of Comparative Zoology 70:1-66. [1930-01] PDF 123533
- Ward, P. S. 2007. Phylogeny, classification, and species-level taxonomy of ants (Hymenoptera: Formicidae). Zootaxa 1668:549–563.
- "Genus: Tatuidris". antweb.org. AntWeb. Retrieved 29 August 2013.
- 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, p. 80–81
- Ward 2007, p. 555–557
- Ward 2011, p. 23
- Donoso 2012, p. 63
- Keller 2011, p. 73
- Brown & Kempf 1968, p. 186
- Lacau et al. 2012, p. 4
- Vasconcelos & Vilhena 2002, p. 278
- 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. (2003), Synopsis and classification of Formicidae, Memoirs of the American Entomological Institute 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: 18172–18177
- Brown, W. L., Jr.; Kempf, W. W. (1968 ["1967"]), "Tatuidris, a remarkable new genus of Formicidae (Hymenoptera).", Psyche 74: 183–190
- Donoso, D.A. (2012), "Additions to the taxonomy of the armadillo ants (Hymenoptera, Formicidae, Tatuidris).", Zootaxa 3503: 61–81
- Keller, R.A. (2011), "A phylogenetic analysis of ant morphology (Hymenoptera: Formicidae) with special reference to the poneromorph subfamilies.", Bulletin of the American Museum of Natural History 355: 1–90
- 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, 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: 101–104
- 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: 14913–14917
- Vasconcelos, Heraldo L.; Vilhena, José M.S. (2002), "First record of the ant genus Tatuidris (Hymenoptera: Formicidae) in Brazil.", Revista de biología tropical 51: 278
- Ward, P.S (2007), "Phylogeny, classification, and species-level taxonomy of ants (Hymenoptera: Formicidae).", Zootaxa 1668: 549–563
- Ward, P.S (2011), "Integrating molecular phylogenetic results into ant taxonomy (Hymenoptera: Formicidae).", Myrmecological News 15: 21–29