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An ergatoid (from
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
'' ergat-'', "worker" + ''
-oid In linguistics, a suffix is an affix which is placed after the stem of a word. Common examples are case endings, which indicate the grammatical case of nouns, adjectives, and verb endings, which form the conjugation of verbs. Suffixes can carry ...
'', "like") is a permanently wingless reproductive adult
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 22 ...
or
termite Termites are small insects that live in colonies and have distinct castes (eusocial) and feed on wood or other dead plant matter. Termites comprise the infraorder Isoptera, or alternatively the epifamily Termitoidae, within the order Blattode ...
. The similar but somewhat ambiguous term ergatogyne refers to any intermediate form between workers and standard
gynes The gyne (, from Greek γυνή, "woman") is the primary reproductive female caste of social insects (especially ants, wasps, and bees of order Hymenoptera, as well as termites). Gynes are those destined to become queens, whereas female workers a ...
. 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 Caste is a form of social stratification characterised by endogamy, hereditary transmission of a style of life which often includes an occupation, ritual status in a hierarchy, and customary social interaction and exclusion based on cultura ...
individuals, another class of ergatogynes, often are not. 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. 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. 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. Without wings, almost all species of ants that solely produce ergatoid queens establish new colonies by fission.


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. 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.


Morphology

The defining morphological features of all ergatoids include functional reproductive organs, which include
spermatheca The spermatheca (pronounced plural: spermathecae ), also called receptaculum seminis (plural: receptacula seminis), is an organ of the female reproductive tract in insects, e.g. ants, bees, some molluscs, oligochaeta worms and certain other ...
e and numerous
ovariole An ovariole is a tubular component of the insect ovary, and the basic unit of egg production. Each ovariole is composed of a germarium (the germline stem cell niche) at the anterior tip, a set of developing oocytes contained within follicles, and ...
s in females and a lack of wings at any point in their life history. Whereas standard queens shed their wings after mating, ergatoids never develop wings, and thus, they must disperse on foot. 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.


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 ''Megalomyrmex foreli'' (named after Auguste Forel) is a Neotropical species of ants in the subfamily Myrmicinae. Habitat and distribution The species is known from Costa Rica south to Andean regions of Colombia, Ecuador, and northern Peru. ''M ...
'' are distinct from workers, with an enlarged gaster, while the ergatoid queens of closely related ''
Megalomyrmex wallacei ''Megalomyrmex wallacei'' is a Neotropical species of ants in the subfamily Myrmicinae. ''Megalomyrmex wallacei'' can be found in Brazil (Amazonas (Brazilian state), Amazonas, Rondônia, Pará, Tocantins), Guyana, Colombia, Costa Rica. This spec ...
'' are very similar in size to their workers.
Army ant The name army ant (or legionary ant or ''marabunta'') is applied to over 200 ant species in different lineages. Because of their aggressive predatory foraging groups, known as "raids", a huge number of ants forage simultaneously over a limit ...
ergatoids are much larger than workers, with large gasters that help them to maintain millions of ants that can make up army ant colonies. The ergatoid queens of the species ''
Blepharidatta conops ''Blepharidatta'' is a rare Neotropical genus of ant in the subfamily Myrmicinae. The genus, formed by predatory species whose small colonies nest in soil or leaf-litter, has up to seven species, but most of them are waiting for a formal taxono ...
'' are quite morphologically distinct as well, with an enlarged head used to wall off the nest entrance in the case of invasion by predators.


Ergatoid males

The species ''
Cardiocondyla obscurior ''Cardiocondyla'' is an Old World genus of ants in the subfamily Myrmicinae. Distribution Approximately 70 species are currently recognized as belonging to this genus, most of which are distributed in the Old World tropics and subtropics, but a ...
'' produces both regular, winged males and ergatoid males. In addition to the loss of wings, male ''C. obscurior'' have altered eye structure and pigmentation, larger bodies, and lifelong spermatogensis. In addition to their morphological changes, male ''C. obscurior'' ergatoids exhibit high levels of aggression toward each other, so much so that they've been dubbed local fighter ants. Similar morphology and behavior has been noted in ''
Cardiocondyla nuda ''Cardiocondyla nuda'' is a species of ant in the subfamily Myrmicinae. It is a widespread ant species, and not invasive in nature. Subspecies * ''Cardiocondyla nuda fajumensis'' Forel, 1913 * ''Cardiocondyla nuda nuda'' Mayr, 1866 - See below ...
'', which only produces ergatoid males.


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. In other species, such as the army ant species, ergatoids live solely to reproduce. Colony formation behavior in most ergatoid queens is starkly different than in most winged queens.


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. Only 3 species' ergatoid queens, all from the genus ''
Pogonomyrmex ''Pogonomyrmex'' is a genus of harvester ants, occurring primarily in the deserts of North, Central, and South America, with a single endemic species from Haiti.Cole, A.C. 1968. Pogonomyrmex Harvester Ants: a study of the genus in North America. ...
'' (harvester ants), have been shown to practice independent colony formation, or ICF.


Evolution

The production of ergatoid queens has developed across at least 16 subfamilies and 55 genera. It has been suggested that this
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
toward ergatoids stems from the DCF behavior that most ergatoid-producing species exhibit. 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.


See also

*
Alate Alate (Latin ''ālātus'', from ''āla'' (“wing”)) is an adjective and noun used in entomology and botany to refer to something that has wings or winglike structures. In entomology In entomology, "alate" usually refers to the winged form o ...
*
Gamergate (ant) A gamergate ( ) is a mated worker ant that can reproduce sexually, i.e., lay fertilized eggs that will develop as females. In the vast majority of ant species, workers are sterile and gamergates are restricted to taxa where the workers have a f ...
*
Gyne The gyne (, from Greek γυνή, "woman") is the primary reproductive female caste of social insects (especially ants, wasps, and bees of order Hymenoptera, as well as termites). Gynes are those destined to become queens, whereas female workers ...
*
Queen ant A queen ant (formally known as a gyne) is an adult, reproducing female ant in an ant colony; generally she will be the mother of all the other ants in that colony. Some female ants, such as the '' Cataglyphis'', do not need to mate to produce offs ...


References

{{Reflist Ants Insect reproduction