Ergatoid

An ergatoid queen of the species Myrmecia esuriens

An ergatoid (from Greek ergat-, "worker" + -oid, "like") is a permanently wingless reproductive adult ant or termite.^[1][2] The similar but somewhat ambiguous term ergatogyne refers to any intermediate form between workers and standard gynes.^[1][3] Ergatoid queens are distinct from other ergatogyne individuals in that they are morphologically consistent within a species and are always capable of mating, whereas inter caste individuals, another class of ergatogynes, often are not.^[1][3] Ergatoids can exhibit wide morphological differences between species, sometimes appearing almost identical to normal workers and other times being quite distinct from both workers and standard queens.^[1][4][5][6] In addition to morphological features, ergatoids among different species can exhibit a wide range of behaviors, with some ergatoids acting only as reproductives and others actively foraging.^[1][7] Ergatoid queens have developed among a large number of ant species, and their presence within colonies can often provide clues on the social structures of colonies and as to how new colonies are founded.^[4] Without wings, almost all species of ants that solely produce ergatoid queens establish new colonies by fission.^[6][7][8]

Nomenclature

The term ergatoid has been used to denote wingless reproductive ants since Margaret Holliday's 1903 paper, "A study of a few ergatogynic ants", although its current usage was suggested and more completely defined in the 1991 paper, "Ergatoid queens and intercastes in ants: two distinct adult forms which look morphologically intermediate between workers and winged queens", by C.P. Peeters.^[1] The paper, "Intercastes, intermorphs, and ergatoid queens: who is who in ant reproduction?" provided further definition and discussion of how certain "intermorphic queens" should be denoted in 1998.^[3]

Morphology

The defining morphological features of all ergatoids include functional reproductive organs, which include spermathecae and numerous ovarioles in females and a lack of wings at any point in their life history.^[1] Whereas standard queens shed their wings after mating, ergatoids never develop wings, and thus, they must disperse on foot.^[4][7] Other than these key, defining traits, ergatoid queen morphology can vary widely among species. Some species' ergatoids look quite similar to workers of the species, whereas other species' ergatoids can exhibit extreme morphologies that make them highly distinct.^[5][6][7]

This top-down view of an ergatoid queen of Blepharidatta conops is a good example of the extreme morphologies that ergatoids can have.^[6]

Variation among species

Ergatoid morphology can vary greatly among different species, even within the same genus. For example, one paper found that the ergatoid queens of Megalomyrmex foreli are distinct from workers, with an enlarged gaster, while the ergatoid queens of closely related Megalomyrmex wallacei are very similar in size to their workers.^[6] Army ant ergatoids are much larger than workers, with large gasters that help them to maintain millions of ants that can make up army ant colonies.^[1] The ergatoid queens of the species Blepharidatta conops are quite morphologically distinct as well, with an enlarged head used to wall off the nest entrance in the case of invasion by predators.^[5]

Ergatoid males

The species Cardiocondyla obscurior produces both regular, winged males and ergatoid males.^[9] In addition to the loss of wings, male C. obscurior have altered eye structure and pigmentation, larger bodies, and lifelong spermatogensis.^[9][10] In addition to their morphological changes, male C. obscurior ergatoids exhibit high levels of aggression toward each other, so much so that they have been dubbed local fighter ants.^[9][10] Similar morphology and behavior has been noted in Cardiocondyla nuda, which only produces ergatoid males.^[11]

Behavior

As with morphology, ergatoid behavior varies greatly among species. In many species, ergatoids are quite numerous, and un-mated ergatoids exhibit the same foraging behaviors that workers do.^[1][7] In other species, such as the army ant species, ergatoids live solely to reproduce.^[1] Colony formation behavior in most ergatoid queens is starkly different than in most winged queens.^[6][7]

Colony formation

Without wings, ergatoid queens founding new colonies disperse on foot. In the vast majority ergatoid-producing species, new colonies are founded by fission when a mated ergatoid and a group of workers disperse to a new nesting site; this is commonly known as dependent colony formation, or DCF.^{[6][7][12][13][14]} Only 3 species' ergatoid queens, all from the genus Pogonomyrmex (harvester ants), have been shown to practice independent colony formation, or ICF.^[8]

Evolution

The production of ergatoid queens has developed across at least 16 subfamilies and 55 genera.^[6][14] It has been suggested that this convergent evolution toward ergatoids stems from the DCF behavior that most ergatoid-producing species exhibit.^[6][13][14] It has also been suggested that the production of ergatoid queens is advantageous as it is less costly, and ergatoid queens may have higher survival rates than winged, independent colony forming queens.^[6][15]

See also

• Alate
• Gamergate (ant)
• Gyne
• Queen ant

References

1. Peeters, C. P. (March 1991). "Ergatoid queens and intercastes in ants: Two distinct adult forms which look morphologically intermediate between workers and winged queens". Insectes Sociaux. 38 (1): 1–15. doi:10.1007/BF01242708. ISSN 0020-1812. S2CID 19188793.
2. da Silva, Iago Bueno; Haifig, Ives; Vargo, Edward L.; Casarin, Fabiana Elaine; da Mota, Marcelo Luiz; Lima, Juliana Toledo; Costa-Leonardo, Ana Maria (2019-10-23). "Ergatoid reproductives in the Neotropical termite Nasutitermes aquilinus (Holmgren) (Blattaria: Isoptera: Termitidae): developmental origin, fecundity, and genetics". Insect Science. 27 (6): 1744–7917.12727. doi:10.1111/1744-7917.12727. ISSN 1672-9609. PMID 31553524. S2CID 202760301.
3. Heinze, J. (1998-05-01). "Intercastes, intermorphs, and ergatoid queens: who is who in ant reproduction?". Insectes Sociaux. 45 (2): 113–124. doi:10.1007/s000400050073. ISSN 0020-1812. S2CID 20786749.
4. Ito, Fuminori (1996-07-01). "Colony Characteristics of the Indonesian Myrmicine Ant Myrmecina sp. (Hymenoptera, Formicidae, Myrmicinae): Polygynous Reproduction by Ergatoid Queens". Annals of the Entomological Society of America. 89 (4): 550–554. doi:10.1093/aesa/89.4.550. ISSN 1938-2901.
5. Brandão, C. R. F.; Diniz, J. L. M.; Silva, P. R.; Albuquerque, N. L.; Silvestre, R. (September 2001). "The first case of intranidal phragmosis in ants. The ergatoid queen of Blepharidatta conops (Formicidae, Myrmicinae) blocks the entrance of the brood chamber". Insectes Sociaux. 48 (3): 251–258. doi:10.1007/PL00001774. ISSN 0020-1812. S2CID 25923049.
6. Peeters, Christian; Adams, Rachelle M. M. (2016). "Uncoupling Flight and Reproduction in Ants: Evolution of Ergatoid Queens in Two Lineages ofMegalomyrmex(Hymenoptera: Formicidae)". Journal of Insect Science. 16 (1): 85. doi:10.1093/jisesa/iew068. ISSN 1536-2442. PMC 5019021. PMID 27620557.
7. Molet, Mathieu; Fisher, Brian L.; Ito, Fuminori; Peeters, Christian (2009-08-25). "Shift from independent to dependent colony foundation and evolution of 'multi-purpose' ergatoid queens in Mystrium ants (subfamily Amblyoponinae): REPLACEMENT BY ERGATOID QUEENS". Biological Journal of the Linnean Society. 98 (1): 198–207. doi:10.1111/j.1095-8312.2009.01257.x.
8. Johnson, R. A. (May 2010). "Independent colony founding by ergatoid queens in the ant genus Pogonomyrmex: queen foraging provides an alternative to dependent colony founding". Insectes Sociaux. 57 (2): 169–176. doi:10.1007/s00040-010-0065-6. ISSN 0020-1812. S2CID 320189.
9. Cremer, S.; Lautenschläger, B.; Heinze, J. (2002-08-01). "A transitional stage between the ergatoid and winged male morph in the ant Cardiocondyla obscurior". Insectes Sociaux. 49 (3): 221–228. doi:10.1007/s00040-002-8305-z. ISSN 0020-1812. S2CID 24415257.
10. Heinze, Jürgen; Trindl, Andreas; Seifert, Bernhard; Yamauchi, Katsusuke (October 2005). "Evolution of male morphology in the ant genus Cardiocondyla". Molecular Phylogenetics and Evolution. 37 (1): 278–288. Bibcode:2005MolPE..37..278H. doi:10.1016/j.ympev.2005.04.005. ISSN 1055-7903. PMID 15922629.
11. Heinze, J.; Kühnholz, S.; Schilder, K.; Hölldobler, B. (September 1993). "Behavior of ergatoid males in the ant,Cardiocondyla nuda" (PDF). Insectes Sociaux. 40 (3): 273–282. doi:10.1007/BF01242363. ISSN 0020-1812. S2CID 9140326.
12. Amor, Fernando; Ortega, Patrocinio; Jowers, Michael J.; Cerdá, Xim; Billen, Johan; Lenoir, Alain; Boulay, Raphaël R. (2011-03-04). "The evolution of worker–queen polymorphism in Cataglyphis ants: interplay between individual- and colony-level selections". Behavioral Ecology and Sociobiology. 65 (7): 1473–1482. Bibcode:2011BEcoS..65.1473A. doi:10.1007/s00265-011-1157-7. hdl:10261/58665. ISSN 0340-5443. S2CID 2784030.
13. Cronin, Adam L.; Molet, Mathieu; Doums, Claudie; Monnin, Thibaud; Peeters, Christian (2013-01-07). "Recurrent Evolution of Dependent Colony Foundation Across Eusocial Insects". Annual Review of Entomology. 58 (1): 37–55. doi:10.1146/annurev-ento-120811-153643. ISSN 0066-4170. PMID 22934981. S2CID 3066504.
14. Peeters, Christian (2012-01-10). "Convergent evolution of wingless reproductives across all subfamilies of ants, and sporadic loss of winged queens (Hymenoptera: Formicidae)". Myrmecological News. 16: 75–91. doi:10.5281/zenodo.844241.
15. Molet, M.; Peeters, C. (May 2006). "Evolution of wingless reproductives in ants: weakly specialized ergatoid queen instead of gamergates in Platythyrea conradti". Insectes Sociaux. 53 (2): 177–182. doi:10.1007/s00040-005-0856-3. ISSN 0020-1812. S2CID 33971950.