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{{short description|Reproductively viable female worker ant}}
{{about|a type of ant|the 2014 video game culture controversy|GamerGate}}
{{Use dmy dates|date=May 2016}}
⚫ ]'' worker, the ant for which the term ''gamergate'' was originally coined<ref group="note">In the original ''Naturwissenschaften'' paper where the term "gamergate" was first used, ''O. berthoudi'' is referred to by the synonym ''Pachycondyla berthoudi''.</ref>]]


A '''gamergate''' ({{IPAc-en|ˈ|ɡ|æ|m|ər|ˌ|ɡ|eɪ|t}} {{Respell|GAMM|ər|gayt}}) is a mated ] 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 functional sperm reservoir (']'). In some species, gamergates reproduce in addition to winged queens (usually upon the death of the original foundress), while in other species the queen caste has been completely replaced by gamergates. In gamergate species, all workers in a colony have similar reproductive potentials, but as a result of physical interactions, a dominance hierarchy is formed and only one or a few top-ranking workers can mate (usually with foreign males) and produce eggs. Subsequently, however, aggression is no longer needed as gamergates secrete chemical signals that inform the other workers of their reproductive status in the colony.
⚫ ]
A '''gamergate''' is a reproductively viable female ] that is able to reproduce with mature males when the ] is lacking a ]. Most commonly occurring within the primitive species of the ], gamergate females differentiate from their fellow workers by a combination of elevated fecundity and aggression-related mutilation of competitors' secondary sexual characteristics. Subsequent to their first mating event, however, aggression is no longer needed as females secrete chemical signals that lead the workers to accept their role as reproducers for the colony.


Depending on the species, there can be one gamergate per colony (]) or several gamergates (]). Most gamergate species have colonies with a few hundred or fewer workers.
Gamergates exist in colonies with winged and ergatoid queens as well as singly in monogynous colonies and alongside other reproductively viable gamergates in polygynous colonies. Most gamergate species are solitary generalist foragers living in arid environments.


==Etymology== ==Etymology==
The term "gamergate" derives from the Greek words {{lang|grc|γάμος}} (''gámos'') and {{lang|grc|ἐργάτης}} (''ergátēs'') and means "married worker." It was coined in 1983 by geneticist ]<ref name=peeters84/> and was first used in scientific literature by entomologists Christian Peeters and Robin Crewe in a 1984 paper published in '']''.<ref name=veron/> The definition typically found in entomological dictionaries is "mated, egg-laying worker,"<ref name=barrows/><ref name=gordh/> and is drawn from the glossary of ] and ]'s 1990 book, '']''.<ref name=barrows/> ''Gamergate'' derives from the Greek words {{lang|grc|γάμος}} ({{Lang|grc-latn|gámos}}) and {{lang|grc|ἐργάτης}} ({{Lang|grc-latn|ergátēs}}) and means 'married worker'. It was coined in 1983 by geneticist ]<ref name=peeters84/> and was first used in scientific literature by entomologists ] and ] in a 1984 paper published in '']''.<ref name=veron/> The definition typically found in entomological dictionaries is 'mated, egg-laying worker',<ref name=barrows/><ref name=gordh/> and is drawn from the glossary of ] and ]'s 1990 book, '']''.<ref name=barrows/>


==Overview== ==Description==
There exist between one hundred and two hundred different species of queenless ants (roughly 1% of all ants), most of which fall within the ]. Whereas female workers in colonies with a queen are ] incapable of mating, in gamergate societies a single reproductive worker and sometimes a cadre of dominant female workers have active ovaries.<ref name=noel/> Gamergate lifespan is short compared to queens in queenright colonies, but gamergates can be replaced by other dominant workers in the colony without risking colony survival. Reproductive investment in gamergate females is thus optimized because non-differentiated gamergates (i.e. reproductively inactive workers) function as laborers.<ref name=choe/> There are 100–200 different species in which gamergates reproduce (roughly 1% of all ants), most of which fall within the ]. Whereas workers (which are all females) in most ant species are ] incapable of storing sperm, in gamergate species one or several workers mate and have active ].<ref name=noel/> Gamergate lifespan is short compared to queens in ] colonies, but gamergates can be replaced by other dominant workers in the colony without risking colony survival. Reproductive investment in gamergate females is thus optimized because non-differentiated gamergates (i.e. reproductively inactive workers) function as laborers.<ref name=choe/>


===Caste structure=== ===Caste structure===
] ]
Within gamergate colonies all females are born reproductively viable and thus represent potential gamergates. Prior to differentiation as a gamergate a dominant female workers must physically inhibit its sisters. For example in the case of ''Diacamma australe'', the first females to reach maturity will clip off the thoracic gemmae of their sisters. This mutilation greatly reduces the attractiveness of the female as a mate. Thereafter persistent domination of worker females by gamergates via physical aggression all but ensure that they will not produce male offspring.<ref name=bourke/> In ''Diacamma nilgiri'' gamergates use dominance interactions to monopolize reproduction without mutilation of sister workers.<ref name=karnik/> The same is true for ''Streblognathus peetersi'' which engage in non-injurious aggression "games" to determine dominance.<ref name=veron/> For all gamergate species, the act of mating eliminates the need to physically dominate female workers. Instead newly produced pheromones or signaling chemicals ensure that workers remain nonreproductive. Although it is unknown to what degree these chemicals act as pheromones or as signals, support for the signaling hypothesis can be found in the loss of gamergate reproductive inhibition of workers as the gamergate grows older and her fecundity diminishes.<ref name=bourke/> Within gamergate colonies, all workers are born reproductively viable and are thus potential gamergates. Prior to differentiation as a gamergate, a dominant worker must physically inhibit its sisters. For example, in the case of '']'', the first female to become reproductively active will clip off the ] gemmae of her sisters, thus greatly reducing their sexual attractiveness. In other genera, persistent domination of worker females by gamergates via physical aggression all but ensure that they will not produce male offspring.<ref name=bourke/> In '']'', gamergates use dominance interactions to monopolize reproduction without mutilation of sister workers.<ref name=cournault/> The same is true for '']'', which engage in non-injurious aggression to determine dominance.<ref name=veron/> For most gamergate species, the start of ovarian activity eliminates the need to physically dominate nestmate workers. Instead newly produced ]s or signaling chemicals ensure that workers remain nonreproductive. Although it is unknown to what degree these chemicals act as pheromones or as signals, support for the signaling hypothesis can be found in the loss of reproductive inhibition of workers as the gamergate grows older and her ] diminishes.<ref name=bourke/>


When a reproductive gamergate dies, it is replaced by a former dominant worker who becomes a new gamergate. New gamergates often emerge from the female cohort closest to maturity at the time of death of the previous gamergate, but in some cases this event triggers adult female workers to differentiate directly to gamergates. For example, when a queen dies in a '']'' colony, workers of the colony will begin to fight for dominance to become the next queen and reproduce the next eggs.<ref name=gorman/> Because reproductively inactive workers are able to become reproductive again after the death of the gamergate, some gamergate species can be considered ] rather than truly ]s.<ref name=crespi/> Mechanisms of gamergate replacement vary among monogynous and polygynous species. When a gamergate dies, it is usually replaced by a formerly submissive worker who proceeds to mate and begins ovarian activity. A new gamergate often originates from a younger ]. For example, when the original founding queen dies in a '']'' colony, younger workers begin to fight for dominance and some become the next reproductives.<ref name=gorman/> Because reproductively inactive workers are able to activate their ovaries after the death of the gamergate, some gamergate species can be considered ] rather than truly ]s.<ref name=crespi/>


In colonies with queens, gamergates, and workers, gamergates occupy an intermediate caste.<ref name=noel/> Research on ] species displaying gamergate social structures has found that there is a fecundity-based hierarchy within the gamergate caste as well. In a study on '']'' sp., it was found that higher-ranked gamergates had more fully developed ]s than low-ranked gamergates. The near absence of sterile gamergates suggests to researchers that particularly low-ranked gamergates may be expelled from their colonies.<ref name=ito/> A three-tiered gamergate hierarchy also occurs within ''Streblognathus peetersi'' with only alpha-gamergates reproducing while beta- and gamma-gamergates await a chance to reproduce when an alpha-gamergate has lost her fecundity or died.<ref name=veron/> Challenges to top-ranked gamergates from workers of the lower-hierarchies are risky for the challenger because gamergates of species like '']'' may mark the challenger by stinging her with special chemicals only produced by the dominant gamergate. These chemicals signal other workers to immobilize the challenger by biting her appendages and holding her for up to a few days until the threat has passed.<ref name=monnin/> In colonies with both queens and gamergates, the latter function as secondary reproductives.<ref name=noel/> Research on ] species has shown that there is a fecundity-based hierarchy among gamergates. In '']'', it was found that higher-ranked gamergates had more fully developed ]s than low-ranked gamergates.<ref name=ito/> In ''Streblognathus peetersi,'' only the alpha worker mates and becomes the gamergate; younger workers await a chance to reproduce when the current gamergate exhibits decreased fecundity or dies.<ref name=veron/> Challenges to gamergates from subordinate workers are risky because the gamergate in species like '']'' may mark the challenger by rubbing special chemicals produced only by the gamergate. These chemicals signal to other workers to immobilize the challenger by biting her appendages and immobilizing her for a few days until her ] levels return to normal.<ref name=monnin/> Subordinate workers play an important policing role in the ] future gamergates and are thus able to increase their indirect fitness.<ref name=noel/>

The role that the dominated worker plays in selection of reproductive females is seen in the expulsion of infertile gamergates in ''Amblyopone'' colonies,<ref name=ito/> and it is even more apparent in ''Dinoponera quadriceps'' where workers may choose to ignore the chemical marker stung by the dominant gamergate. If the dominant gamergate is less reproductively fit than the challenger, workers may instead bite the appendages of the gamergate and hold her immobilized allowing a more fecund challenger to establish herself as the new alpha-gamergate. By playing a part in the selection of gamergates, dominated workers are able to increase their indirect reproduction.<ref name=noel/>


===Social structure variation and ecology=== ===Social structure variation and ecology===
There is variation within the social structure of ant colonies with a gamergate caste. Some species such as ''Harpegnathos saltator'',<ref name=peeters95/> '']'' spp., ''Gnamptogynes menadensis'', and ''Rhytidoponera confusa'' have a winged alate queen caste, a gamergate caste, and a non-reproductive worker caste.<ref name=peeters01/> Some species that normally have a queen caste have been shown to persist for long periods of time by relying on the gamergate caste. For example, a colony of '']'' in 1998 was collected and lasted three years reproducing without a queen ant.<ref name=dietemann/> There is much variation in the social structure of ant colonies with gamergates. Some species such as '']'',<ref name=peeters95/> '']'', '']'', and '']'' have a winged ] queen caste as well as gamergates.<ref name=peeters01/> Queenless species with only gamergates and workers may have a ] structure with a single gamergate or they may have a ] structure with multiple gamergates. Examples of monogynous queenless species include '']'', '']'', ''Diacamma australe'', '']'', '']'', '']'', and '']''.<ref name=peeters91/> Examples of polygynous queenless species include '']'', '']'',<ref name="Peetere1985">{{cite journal |last1=Peetere |first1=C. P. |last2=Crewe |first2=R. M. |year=1985 |title=Queenlessness and reproductive differentiation in ''Ophthalmopone hottentota'' |journal=South African Journal of Zoology |volume=20 |issue=4 |page=268 |doi=10.1080/02541858.1985.11447948|doi-access=free }}</ref> and all known queenless species of '']''.<ref name=peeters91/> In the queenless ''Ophthalmopone berthoudi'', foreign males visit underground nests to mate with young workers.<ref name=antweb-pachycondyla/>


⚫ Ecologically, gamergate species from different ] and ] often tend to share certain characteristics. Many gamergate species are solitary generalist foragers living in ] environments.<ref name="choe" /> Similar to species with ] queens, the evolution of gamergate reproduction is hypothesized to be associated with a shift to ]. Myrmecologists Christian Peeters and ] have also suggested that "the evolution of gamergate reproduction appears strongly associated with the adaptive benefits of secondary polygyny (e.g. increased colony lifespan and resource inheritance), and it is the preferred option in species having workers able to store sperm."<ref name="peeters01" />
Queenless species with only gamergates and workers may have a ] structure with a single gamergate female reproducing for the entire colony like a queen or they may have a ] structure with multiple reproductive gamergates laying ] eggs. Examples of monogynous queenless species include ''Pachycondyla krugeri'', ''P. sublaevis'', ''Diacamma australe'', ''D. rugosum'', ''Platythyrea lamellosa'', and ''Streblognathus aethiopicus'' among others.<ref name=peeters91/> For monogynous gamergates, social regulation is based on morphometry (sexual attractiveness) and fecundity (ovarian oogenesis). Examples of polygynous queenless species include ''Ophthalmopone berthoudi'', ''O. hottentota'', ''Dinoponera quadriceps'', and all known queenless species of ''Rhytidoponera'' spp.<ref name=peeters91/> In the queenless '']'' (junior synonym of '']''), foreign males visit underground nests to mate with the polygynous gamergate workers.<ref name=antweb-pachycondyla/>

Yet other systems exist for example in ''Pachycondyla'' spp. where gamergates and ]s share a colony with workers.<ref name=tebeau/>

⚫ Ecologically, gamergate species from different tribes and genera often tend to share certain characteristics. Most gamergate species are solitary generalist foragers living in arid environments.<ref name=choe/> Like ergatoid systems, the evolution of gamergate social structure is hypothesized to be a response to frequent colonial fission events such as periodic flooding or changes in microclimate such as might occur in harsh arid climates. Myrmecologists Christian Peeters and Fuminori Ito have also suggested that "the evolution of gamergate reproduction appears strongly associated with the adaptive benefits of secondary polygyny (e.g. increased colony lifespan and resource inheritance), and it is the preferred option in species havinf workers able to reproduce sexually."<ref name=peeters01/>


==Classification dispute== ==Classification dispute==
The utility of "gamergate" as a morphological designation is not without critics. Within the field of ] it is a matter of dispute whether ] should be defined primarily by reproductive role or by physical morphology. Notably, Alfred Buschinger has argued that the term "worker" should be applied only to those ants who make up the non-reproductive caste and "queen" should be applied only to reproductively viable female ants regardless of their physical appearance. Hölldobler and Wilson suggest that the two positions can be semantically resolved and that the most fruitful approach would be to keep classification "somewhat loose, incorporating either anatomy or roles in a manner that maximizes convenience, precision, and clarity of expression."<ref name=holldobler/> The utility of ''gamergate'' as a morphological designation is not without critics. Within the field of ] it is a matter of dispute whether ] should be defined primarily by reproductive role or by physical morphology. Notably, ] has argued that the term ''worker'' should be applied only to those ants who make up the non-reproductive caste and ''queen'' should be applied only to reproductively viable female ants regardless of their physical appearance. Hölldobler and Wilson suggest that the two positions can be semantically resolved and that the most fruitful approach would be to keep classification "somewhat loose, incorporating either anatomy or roles in a manner that maximizes convenience, precision, and clarity of expression."<ref name=holldobler/>


==Genera with gamergates== ==Genera with gamergates==
The existence of gamergates is ], or ancestral, in ants.<ref name=dietemann/> They are known to be present in the following genera:
This list may be incomplete and may require expansion:


*Poneromorph subfamilies *]
**Amblyoponinae **]
***'']''<ref name=ito/><ref group="note">Throughout Ito's 1993 paper for the ''Journal of Natural History'', he refers not to ''Stigmatomma'' but to ''Amblyopone''. At the time ''Stigmatomma'' was considered to be a synonym of ''Amblyopone''.</ref>
⚫ ***'']''<ref name=ito/>
**Ectatomminae **]
⚫ ***'']''<ref name=gobin/>
***'']''<ref name=peeters87/> ***'']''<ref name=peeters87/>
**Ponerinae **]
***'']''<ref name=antweb-diacamma/> ***'']''<ref name="Schmidt&Shattuck_2014"/>
***'']''<ref name=haskins/> ***'']''<ref name="Schmidt&Shattuck_2014"/><ref name=antweb-diacamma/>
***'']''<ref name=peeters00/> ***'']''<ref name="Schmidt&Shattuck_2014"/><ref name=haskins/>
***'']''<ref name=schmidt/> ***'']''<ref name="peeters2016"/>
***'']''<ref name=peeters85/>/'']''<ref name=antweb-pachycondyla/> ***'']''<ref name="Schmidt&Shattuck_2014"/>
***'']''<ref name=schilder/> ***'']''<ref name="Schmidt&Shattuck_2014"/><ref name=peeters00/>
***'']''<ref name=antweb-streblognathus/> ***'']''<ref name="Schmidt&Shattuck_2014"/>
***'']''<ref name="Schmidt&Shattuck_2014"/><ref name=peeters85/>
⚫ *Myrmicinae
::*'']''<ref name=dietemann/> ***'']''<ref name="Schmidt&Shattuck_2014"/><ref name=schilder/>
***'']''<ref name="Schmidt&Shattuck_2014"/>
::*'']'' - Note: Although the asexual ''Pristomyrmex'' females may technically meet the "gamergate" definition, Hölldobler and Wilson argue that it stretches the definition beyond its useful limits when applied to species practicing ].<ref name=holldobler/>
***'']''<ref name="Schmidt&Shattuck_2014"/><ref name=antweb-streblognathus/>
***'']''<ref name="Schmidt&Shattuck_2014"/>
*]
::*'']''<ref name=dietemann/>
⚫ *]
::*'']''<ref name=Holldobler2002/>


==See also== ==See also==
*] * ]
* ]
* ]
* ]

==Notes==
{{Reflist|group=note}}


==References== ==References==
{{Reflist|2|refs= {{Reflist|30em|refs=
<ref name=antweb-diacamma>{{cite web |url=http://www.antweb.org/description.do?name=Diacamma&rank=genus&project=allantwebants |title= Genus: ''Diacamma'' |date= |website=antweb.org |publisher=] |accessdate=12 August 2014}}</ref> <ref name=antweb-diacamma>{{cite web |url=http://www.antweb.org/description.do?name=Diacamma&rank=genus&project=allantwebants |title= Genus: ''Diacamma'' |website=antweb.org |publisher=] |access-date=12 August 2014}}</ref>
<ref name=antweb-pachycondyla>{{cite web |url=http://www.antweb.org/description.do?genus=pachycondyla&species=berthoudi&rank=species&project=allantwebants |title= Species: ''Pachycondyla berthoudi'' |date= |website=antweb.org |publisher=] |accessdate=12 August 2014}}</ref> <ref name=antweb-pachycondyla>{{cite web |url=http://www.antweb.org/description.do?genus=pachycondyla&species=berthoudi&rank=species&project=allantwebants |title= Species: ''Pachycondyla berthoudi'' |website=antweb.org |publisher=] |access-date=12 August 2014}}</ref>
<ref name=antweb-streblognathus>{{cite web |url=http://www.antweb.org/description.do?subfamily=ponerinae&genus=streblognathus&rank=genus&project=allantwebants|title= Genus: ''Streblognathus'' |date= |website=antweb.org |publisher=] |accessdate=12 August 2014}}</ref> <ref name=antweb-streblognathus>{{cite web |url=http://www.antweb.org/description.do?subfamily=ponerinae&genus=streblognathus&rank=genus&project=allantwebants|title= Genus: ''Streblognathus'' |website=antweb.org |publisher=] |access-date=12 August 2014}}</ref>
<ref name=barrows>{{cite book |last= Barrows|first= Edward M.|date= 2011|title= Animal Behavior Desk Reference: A Dictionary of Animal Behavior, Ecology, and Evolution|edition= Third|chapter= Caste - Gamergate|location= |publisher= CRC Press|page= 75|isbn= 9781439836514}}</ref> <ref name=barrows>{{cite book |last= Barrows|first= Edward M.|date= 2011|title= Animal Behavior Desk Reference: A Dictionary of Animal Behavior, Ecology, and Evolution|edition= Third|chapter= Caste – Gamergate|publisher= CRC Press|page= 75|isbn= 9781439836514}}</ref>
<ref name=bourke>{{cite book |last= Bourke|first= Andrew F. G.|date= 1995|title= Social Evolution in Ants: Monographs in behavior and ecology|chapter= 7 - Kin Conflict: Reproduction (Part 2 - Queen Policing, Queen Control, and Queen Signaling)|location= |publisher= ]|pages= 239–240|isbn= 9780691044262}}</ref> <ref name=bourke>{{cite book |last= Bourke|first= Andrew F. G.|date= 1995|title= Social Evolution in Ants: Monographs in behavior and ecology|chapter= 7 – Kin Conflict: Reproduction (Part 2 – Queen Policing, Queen Control, and Queen Signaling)|publisher= ]|pages= 239–240|isbn= 9780691044262}}</ref>
<ref name=choe>{{cite book |last1= Choe|first1= Jae C.|last2= Crespi|first2= Bernard J.|date= 1997|title= The Evolution of Social Behaviour in Insects and Arachnids|chapter= Morphologically 'Primitive' Ants|location= |publisher= ]|page= 385|isbn= 9780521589772}}</ref> <ref name=choe>{{cite book |last1= Choe|first1= Jae C.|last2= Crespi|first2= Bernard J.|author-link2=Bernard Crespi|date= 1997|title= The Evolution of Social Behaviour in Insects and Arachnids|url= https://archive.org/details/evolutionsocialb00choe|url-access= limited|chapter= Morphologically 'Primitive' Ants|publisher= ]|page= |isbn= 9780521589772}}</ref>
<ref name=crespi>{{cite book |last= Crespi|first= Bernard J.|editor-last= Martins|editor-first= Emília P.|date= 1996|title= Phylogenies and the Comparative Method in Animal Behavior|chapter= 9 - Comparative Analysis of the Origins and Losses of Eusociality: Causal Mosaics and Historical Uniqueness (Part 6 - Formicidae)|location= |publisher= ]|page= 272|isbn= 9780195092103}}</ref> <ref name=crespi>{{cite book |last= Crespi|first= Bernard J.|editor-last= Martins|editor-first= Emília P.|date= 1996|title= Phylogenies and the Comparative Method in Animal Behavior|chapter= 9 – Comparative Analysis of the Origins and Losses of Eusociality: Causal Mosaics and Historical Uniqueness (Part 6 – Formicidae)|publisher= ]|page= 272|isbn= 9780195092103}}</ref>
<ref name=dietemann>{{cite journal|last1= Dietemann|first1= V.|last2= Peeters|first2= C|last3= Hölldobler|first3= B.|authorlink3= Bert Hölldobler|date= 2004|title= Gamergates in the Australian ant subfamily Myrmeciinae |journal= ]|publisher= ]|issn= 0028-1042|volume= 91|issue= 9|pages= 432–435}}</ref> <ref name=dietemann>{{cite journal|last1= Dietemann|first1= V.|last2= Peeters|first2= C|last3= Hölldobler|first3= B.|author-link3= Bert Hölldobler|date= 2004|title= Gamergates in the Australian ant subfamily Myrmeciinae |journal= ]|issn= 0028-1042|volume= 91|issue= 9|pages= 432–435 |doi=10.1007/s00114-004-0549-1 |pmid=15278223|bibcode = 2004NW.....91..432D |s2cid= 26161972}}</ref>
<ref name=gobin>{{cite journal |last1= Gobin |first1= B. |last2= Peeters |first2= C. |last3= Billen |first3= J. |title= Production of trophic eggs by virgin workers in the ponerine ant Gnamptogenys menadensis |date= September 1998 |journal= Physiological Entomology |volume= 23 |issue= 4 |pages= 329–336 |doi= 10.1046/j.1365-3032.1998.234102.x |s2cid= 83569093 |url= http://ecologie.snv.jussieu.fr/socialite/PhysEnto%20Gobin.pdf |access-date= 26 August 2016 |archive-url= https://web.archive.org/web/20160827050750/http://ecologie.snv.jussieu.fr/socialite/PhysEnto%20Gobin.pdf |archive-date= 27 August 2016 |url-status= dead }}</ref>
<ref name=gordh>{{cite book |last= Gordh|first= Gordon|authorlink= Gordon Gordh|date= 2011|title= A Dictionary of Entomology|chapter= Gamergate|location= |publisher= ]|page= 608|isbn= 9781845935429}}</ref> <ref name=gordh>{{cite book |last= Gordh|first= Gordon|author-link= Gordon Gordh|date= 2011|title= A Dictionary of Entomology|chapter= Gamergate|publisher= ]|page= 608|isbn= 9781845935429}}</ref>
<ref name=gorman>{{cite news|last1= Gorman|first1= James|title= The Ant Queen Is Dead. Let the Battles Begin|url= http://www.nytimes.com/2014/05/27/science/the-ant-queen-is-dead-let-the-battles-begin.html|accessdate= 12 August 2014|publisher= ]|issn= 0362-4331|date= 27 May 2014}}</ref> <ref name=gorman>{{cite news|last1= Gorman|first1= James|title= The Ant Queen Is Dead. Let the Battles Begin|url= https://www.nytimes.com/2014/05/27/science/the-ant-queen-is-dead-let-the-battles-begin.html|access-date= 12 August 2014|work= ]|issn= 0362-4331|date= 27 May 2014}}</ref>
<ref name=haskins>{{cite journal|last1= Haskins|first1= Caryl Parker|authorlink1= Caryl Parker Haskins|last2= Zahl|first2= P. A.|date= 1971|title= The reproductive pattern of Dinoponera grandis Roger (Hymenoptera, Ponerinae) with notes on the ethology of the species|journal= ]|publisher= ]|issn= 0033-2615|volume= 78|issue= |pages= 1–11|doi= }}</ref>
<ref name=holldobler>{{cite book |last1= Hölldobler|first1= Bert|authorlink1= Bert Hölldobler|last2= Wilson|first2= E. O.|authorlink2= E. O. Wilson|date= 1990|title= ]|chapter= Caste and Division of Labor|location= |publisher= ]|pages= 301 & 305|isbn= 9780674040755}}</ref> <ref name=haskins>{{cite journal|last1= Haskins|first1= Caryl Parker|author-link1= Caryl Parker Haskins|last2= Zahl|first2= P. A.|date= 1971|title= The reproductive pattern of Dinoponera grandis Roger (Hymenoptera, Ponerinae) with notes on the ethology of the species|journal= ]|issn= 0033-2615|volume= 78|issue= 1–2|pages= 1–11|doi= 10.1155/1971/914147|doi-access= free}}</ref>
<ref name=holldobler>{{cite book|last1= Hölldobler|first1= Bert|author-link1= Bert Hölldobler|last2= Wilson|first2= E. O.|author-link2= E. O. Wilson|date= 1990|title= The Ants|chapter= Caste and Division of Labor|publisher= ]|pages= |isbn= 9780674040755|title-link= The Ants}}</ref>
<ref name=ito>{{cite journal |last= Ito|first= Fuminori|date= 1993|title= Social organization in a primitive ponerine ant: queenless reproduction, dominance hierarchy and functional polygyny in Amblyopone sp. (reclinata group) (Hymenoptera: Formicidae: Ponerinae)|journal= ]|publisher= ]|issn= 0022-2933|volume= 27|issue= 6|pages= 1315–1324|doi= 10.1080/00222939300770751}}</ref>
<ref name=karnik>{{cite journal |last= Karnik|first= Nutan|last2= Channaveerappa|first2= H.|last3= Ranganath|first3= H. A.|last4= Gadagkar|first4= Raghavendra|authorlink4= Raghavendra Gadagkar|date= 2010|title= Karyotype instability in the ponerine ant genus ''Diacamma''|journal= ]|publisher= ]|issn= 0022-1333|volume= 89|issue= 2|pages= 173–82|doi= 10.1007/s12041-010-0023-0}}</ref> <ref name=Holldobler2002>{{cite journal|last1=Hölldobler|first1=Bert|last2=Liebig|first2=Jürgen|last3=Alpert|first3=Gary|title=Gamergates in the myrmicine genus ''Metapone'' (Hymenoptera:Formicidae)|journal=Naturwissenschaften|date=2002|volume=89|issue=7|pages=305–307|doi=10.1007/s00114-002-0329-8|pmid=12216860|bibcode = 2002NW.....89..305H |s2cid=6945541}}</ref>
<ref name=monnin>{{cite journal |last= Monnin|first= Thibaud|last2= Ratnieks|first2= Francis L. W.|last3= Jones|first3= Graeme R.|last4= Beard|last4= Richard|date= 5 September 2002|title= Pretender punishment induced by chemical signalling in a queenless ant|journal= ]|publisher= ]|issn= 0028-0836|volume= 419|issue= 6902|pages= 61–65|doi= 10.1038/nature00932}}</ref> <ref name=ito>{{cite journal |last= Ito|first= Fuminori|date= 1993|title= Social organization in a primitive ponerine ant: queenless reproduction, dominance hierarchy and functional polygyny in Amblyopone sp. (reclinata group) (Hymenoptera: Formicidae: Ponerinae)|journal= ]|issn= 0022-2933|volume= 27|issue= 6|pages= 1315–1324|doi= 10.1080/00222939300770751|bibcode= 1993JNatH..27.1315I}}</ref>
<ref name=cournault>{{cite journal |last1= Cournault|first1= Laurent|last2= Peeters|first2= Christian|date= 2012|title= Aggression regulates monogyny in non-mutilating ''Diacamma'' ants|journal= ]|volume= 59|issue= 4|pages= 533–539|doi= 10.1007/s00040-012-0251-9|s2cid= 16179257}}</ref>
⚫ <ref name=noel>{{cite press release |last= Noël|first= Carine|date= 6 September 2002|title= How queenless ants regulate their conflicts|url= http://www.cnrs.fr/cw/en/pres/compress/fourmis2.htm|location= Paris|publisher= ]|accessdate=2014-09-10}}</ref>
<ref name=peeters84>{{cite journal |last= Peeters|first= Christian|last2= Crewe|first2= Robin|date= 1984|title= Insemination Controls the Reproductive Division of Labour in a Ponerine Ant|journal= ]|publisher= ]|issn= 0028-1042|volume= 71|issue= 1|pages= 50–51|doi= 10.1007/BF00365989}}</ref> <ref name=monnin>{{cite journal |last1= Monnin|first1= Thibaud|last2= Ratnieks|first2= Francis L. W.|last3= Jones|first3= Graeme R.|last4= Beard|first4= Richard|date= 5 September 2002|title= Pretender punishment induced by chemical signalling in a queenless ant|journal= ]|issn= 0028-0836|volume= 419|issue= 6902|pages= 61–65|doi= 10.1038/nature00932|bibcode = 2002Natur.419...61M |pmid=12214231|s2cid= 4372841|url= http://eprints.whiterose.ac.uk/99/1/ratnieksflw2.pdf}}</ref>
⚫ <ref name=noel>{{cite press release|last= Noël|first= Carine|date= 6 September 2002|title= How queenless ants regulate their conflicts|url= http://www.cnrs.fr/cw/en/pres/compress/fourmis2.htm|location= Paris|publisher= ]|access-date= 2014-09-10|url-status= dead|archive-url= https://web.archive.org/web/20030302185140/http://www.cnrs.fr/cw/en/pres/compress/fourmis2.htm|archive-date= 2 March 2003|df= dmy-all}}</ref>
⚫ <ref name=peeters85>{{cite journal |last= Peeters|first= Christian|last2= Crewe|first2= Robin M.|date= 1985|title= Worker reproduction in the ponerine ant Ophthalmopone berthoudi: an alternative form of eusocial organization|journal= ]|publisher= ]|issn= 0340-5443|volume= 18|issue= 1|pages= 29–37|doi= 10.1007/BF00299235}}</ref>
<ref name=peeters87>{{cite journal |last= Peeters|first= Christian P.|date= 1987|title= The Reproductive Division of Labour in the Queenless Ponerine Ant Rhytidoponera sp. 12|journal= ]|publisher= ]|issn= 0020-1812|volume= 34|issue= 2|pages= 75–86|doi= 10.1007/BF02223826}}</ref> <ref name=peeters84>{{cite journal |last1= Peeters|first1= Christian|last2= Crewe|first2= Robin|date= 1984|title= Insemination Controls the Reproductive Division of Labour in a Ponerine Ant|journal= ]|issn= 0028-1042|volume= 71|issue= 1|pages= 50–51|doi= 10.1007/BF00365989 |bibcode=1984NW.....71...50P|s2cid= 43871631}}</ref>
⚫ <ref name=peeters85>{{cite journal |last1= Peeters|first1= Christian|last2= Crewe|first2= Robin M.|date= 1985|title= Worker reproduction in the ponerine ant Ophthalmopone berthoudi: an alternative form of eusocial organization|journal= ]|issn= 0340-5443|volume= 18|issue= 1|pages= 29–37|doi= 10.1007/BF00299235|bibcode= 1985BEcoS..18...29P|s2cid= 20041163|url= https://zenodo.org/record/25727}}</ref>
<ref name=peeters91>{{cite book|last= Peeters|first = Christian P.|editor1-last= Veeresh|editor1-first= G. K.|editor2-last= Mallik|editor2-first= B.|title= Social Insects and the Environment: Proceedings of the 11th International Congress of IUSSI, 1990 (International Union for the Study of Social Insects)|date= 1991|publisher= Brill Academic Pub|isbn= 978-9004093164|page= 234|url= http://books.google.com/books?id=ZsMUAAAAIAAJ&pg=PA234|accessdate= 12 August 2014}}</ref>
<ref name=peeters95>{{cite journal |last= Peeters|first= Christian|last2= Hölldobler|first2= Bert|authorlink2= Bert Hölldobler|date= November 1995|title= Reproductive cooperation between queens and their mated workers: The complex life history of an ant with a valuable nest|journal= ]|publisher= ]|issn= 0027-8424|volume= 92|issue= 24|pages= 10977–10979|doi= 10.1073/pnas.92.24.10977}}</ref> <ref name=peeters87>{{cite journal |last= Peeters|first= Christian P.|date= 1987|title= The Reproductive Division of Labour in the Queenless Ponerine Ant Rhytidoponera sp. 12|journal= ]|issn= 0020-1812|volume= 34|issue= 2|pages= 75–86|doi= 10.1007/BF02223826|s2cid= 26116621}}</ref>
<ref name=peeters00>{{cite journal |last= Peeters|first= Christian|last2= Hölldobler|first2= Bert|authorlink2= Bert Hölldobler|date= 2000|title= Sexual reproduction by both queens and workers in the ponerine ant ''Harpegnathos saltator''|journal= ]|publisher= ]|issn= 0020-1812|volume= 47|issue= 4|pages= 325–332|doi= 10.1007/PL00001724}}</ref> <ref name=peeters91>{{cite book|last= Peeters|first = Christian P.|editor1-last= Veeresh|editor1-first= G. K.|editor2-last= Mallik|editor2-first= B.|title= Social Insects and the Environment: Proceedings of the 11th International Congress of IUSSI, 1990 (International Union for the Study of Social Insects)|date= 1991|publisher= Brill Academic Pub|isbn= 978-9004093164|page= 234|url= https://books.google.com/books?id=ZsMUAAAAIAAJ&pg=PA234|access-date= 12 August 2014}}</ref>
<ref name=peeters01>{{cite journal |last= Peeters|first= Christian|last2= Ito|first2= Fuminori|date= 2001|title= Colony Dispersal and the Evolution of Queen Morphology in Social Hymenoptera|journal= ]|publisher= ]|issn= 0066-4170|volume= 46|issue= |pages= 601–30|doi= 10.1146/annurev.ento.46.1.601}}</ref> <ref name=peeters95>{{cite journal |last1= Peeters|first1= Christian|last2= Hölldobler|first2= Bert|author-link2= Bert Hölldobler|date= November 1995|title= Reproductive cooperation between queens and their mated workers: The complex life history of an ant with a valuable nest|journal= ]|issn= 0027-8424|volume= 92|issue= 24|pages= 10977–10979|doi= 10.1073/pnas.92.24.10977 |pmid=11607589 |pmc=40553|bibcode = 1995PNAS...9210977P |doi-access= free}}</ref>
<ref name=schilder>{{cite journal |last1= Schilder|first1= Klaus|last2= Heinze|first2= Jürgen|last3= Hölldobler|first3= Bert|authorlink3= Bert Hölldobler|date= January 1999|title= Colony structure and reproduction in the thelytokous parthenogenetic ant ''Platythyrea punctata'' (F. Smith) (Hymenoptera, Formicidae) |journal= ]|publisher= ]|issn= 0020-1812|volume= 46|issue= 2|pages= 150–158|doi= 10.1007/s000400050126}}</ref> <ref name=peeters00>{{cite journal |last1= Peeters|first1= Christian|last2= Hölldobler|first2= Bert|author-link2= Bert Hölldobler|date= 2000|title= Sexual reproduction by both queens and workers in the ponerine ant ''Harpegnathos saltator''|journal= ]|issn= 0020-1812|volume= 47|issue= 4|pages= 325–332|doi= 10.1007/PL00001724|s2cid= 8728206}}</ref>
<ref name=schmidt>{{cite book|last1= Schmidt|first1= Chris A.|title= Molecular phylogenetics and taxonomic revision of ponerine ants (Hymenoptera: Formicidae: Ponerinae)|date= 2011|publisher= ProQuest, UMI Dissertation Publishing|isbn= 978-1244009004|page= 142|url= http://books.google.com/books?id=Sby7w0Hec6EC&pg=PA142|accessdate= 12 August 2014}}</ref> <ref name=peeters01>{{cite journal |last1= Peeters|first1= Christian|last2= Ito|first2= Fuminori|date= 2001|title= Colony Dispersal and the Evolution of Queen Morphology in Social Hymenoptera|journal= ]|issn= 0066-4170|volume= 46|pages= 601–30|doi= 10.1146/annurev.ento.46.1.601|pmid= 11112181|citeseerx= 10.1.1.562.7031}}</ref>
<ref name=peeters2016>{{cite journal |last1= Peeters |first1= C. |last2= Fisher |first2= B.L. |title= Gamergates (mated egg-laying workers) and queens both reproduce in Euponera sikorae ants from Madagascar. |journal= African Entomology |volume= 24 |issue= 1 |pages= 180–187 |doi= 10.4001/003.024.0180 |date= 14 March 2016 |s2cid= 51851380 |url= http://www.fisherlab.org/uploads/3/9/4/4/39442661/128_peeters_fisher_2016.pdf |access-date= 26 August 2016}}</ref>
⚫ <ref name=tebeau>{{Cite thesis |last= Tebeau|first= Andrew|title= Reproductive strategies and colony relatedness in the invasive ponerine and ''Pachycondyla chinensis'' (Emery)|type= |pages= 18–19|url= |year= 2009|publisher= ]. Republished by ''All Theses'' (Paper 626)|accessdate= }}</ref>
<ref name=schilder>{{cite journal |last1= Schilder|first1= Klaus|last2= Heinze|first2= Jürgen|last3= Hölldobler|first3= Bert|author-link3= Bert Hölldobler|date= January 1999|title= Colony structure and reproduction in the thelytokous parthenogenetic ant ''Platythyrea punctata'' (F. Smith) (Hymenoptera, Formicidae) |journal= ]|issn= 0020-1812|volume= 46|issue= 2|pages= 150–158|doi= 10.1007/s000400050126|s2cid= 13729041}}</ref>
⚫ <ref name=veron>{{cite journal |last= Véron|first= Géraldine|date= February 2005|title= La reine des fourmis couronnée au combat|url= http://www2.cnrs.fr/presse/journal/1965.htm|archiveurl= http://web.archive.org/web/20050210024631/http://www2.cnrs.fr/presse/journal/1965.htm|archivedate=10 February 2005|journal= Le Journal du CNRS|publisher= ]|issn= 0994-7647|volume= |issue= 181|pages= 50–51|doi= |accessdate=9 September 2014}}</ref>
<ref name="Schmidt&Shattuck_2014">{{cite journal |last1=Schmidt |first1=C. A |last2=Shattuck |first2=S. O. |s2cid=4540972 |date=2014 |title=The Higher Classification of the Ant Subfamily Ponerinae (Hymenoptera: Formicidae), with a Review of Ponerine Ecology and Behavior |journal=] |volume=3817 |issue=1 |pages=1–242 |doi=10.11646/zootaxa.3817.1.1 |pmid=24943802}}</ref>
⚫ <!--REF UNUSED AS OF AUGUST 2015<ref name=tebeau>{{Cite thesis |last= Tebeau|first= Andrew|title= Reproductive strategies and colony relatedness in the invasive ponerine and ''Pachycondyla chinensis'' (Emery)|pages= 18–19|year= 2009|publisher= ]. Republished by ''All Theses'' (Paper 626)}}</ref>-->
⚫ <ref name=veron>{{cite journal |last= Véron|first= Géraldine|date= February 2005|title= La reine des fourmis couronnée au combat|url= http://www2.cnrs.fr/presse/journal/1965.htm|archive-url= https://web.archive.org/web/20050210024631/http://www2.cnrs.fr/presse/journal/1965.htm|archive-date=10 February 2005|journal= Le Journal du CNRS|issn= 0994-7647|issue= 181|pages= 50–51|access-date=9 September 2014}}</ref>
}} }}

{{Eusociality}}


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Latest revision as of 22:55, 11 November 2024

Reproductively viable female worker ant

Lateral view of an Ophthalmopone berthoudi worker, the ant for which the term gamergate was originally coined

A gamergate (/ˈɡæmərˌɡeɪt/ GAMM-ər-gayt) 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 functional sperm reservoir ('spermatheca'). In some species, gamergates reproduce in addition to winged queens (usually upon the death of the original foundress), while in other species the queen caste has been completely replaced by gamergates. In gamergate species, all workers in a colony have similar reproductive potentials, but as a result of physical interactions, a dominance hierarchy is formed and only one or a few top-ranking workers can mate (usually with foreign males) and produce eggs. Subsequently, however, aggression is no longer needed as gamergates secrete chemical signals that inform the other workers of their reproductive status in the colony.

Depending on the species, there can be one gamergate per colony (monogyny) or several gamergates (polygyny). Most gamergate species have colonies with a few hundred or fewer workers.

Etymology

Gamergate derives from the Greek words γάμος (gámos) and ἐργάτης (ergátēs) and means 'married worker'. It was coined in 1983 by geneticist William Louis Brown and was first used in scientific literature by entomologists Christian Peeters and Robin Crewe in a 1984 paper published in Naturwissenschaften. The definition typically found in entomological dictionaries is 'mated, egg-laying worker', and is drawn from the glossary of Bert Hölldobler and E. O. Wilson's 1990 book, The Ants.

Description

There are 100–200 different species in which gamergates reproduce (roughly 1% of all ants), most of which fall within the poneromorph subfamilies. Whereas workers (which are all females) in most ant species are morphologically incapable of storing sperm, in gamergate species one or several workers mate and have active ovaries. Gamergate lifespan is short compared to queens in queenright colonies, but gamergates can be replaced by other dominant workers in the colony without risking colony survival. Reproductive investment in gamergate females is thus optimized because non-differentiated gamergates (i.e. reproductively inactive workers) function as laborers.

Caste structure

Dorsal view of a nonreproductive female Diacamma australe worker, lacking anterior thoracic gemmae (buds).

Within gamergate colonies, all workers are born reproductively viable and are thus potential gamergates. Prior to differentiation as a gamergate, a dominant worker must physically inhibit its sisters. For example, in the case of Diacamma australe, the first female to become reproductively active will clip off the thoracic gemmae of her sisters, thus greatly reducing their sexual attractiveness. In other genera, persistent domination of worker females by gamergates via physical aggression all but ensure that they will not produce male offspring. In Diacamma nilgiri, gamergates use dominance interactions to monopolize reproduction without mutilation of sister workers. The same is true for Streblognathus peetersi, which engage in non-injurious aggression to determine dominance. For most gamergate species, the start of ovarian activity eliminates the need to physically dominate nestmate workers. Instead newly produced pheromones or signaling chemicals ensure that workers remain nonreproductive. Although it is unknown to what degree these chemicals act as pheromones or as signals, support for the signaling hypothesis can be found in the loss of reproductive inhibition of workers as the gamergate grows older and her fecundity diminishes.

Mechanisms of gamergate replacement vary among monogynous and polygynous species. When a gamergate dies, it is usually replaced by a formerly submissive worker who proceeds to mate and begins ovarian activity. A new gamergate often originates from a younger cohort. For example, when the original founding queen dies in a Harpegnathos saltator colony, younger workers begin to fight for dominance and some become the next reproductives. Because reproductively inactive workers are able to activate their ovaries after the death of the gamergate, some gamergate species can be considered cooperative breeders rather than truly eusocial insects.

In colonies with both queens and gamergates, the latter function as secondary reproductives. Research on Amblyoponinae species has shown that there is a fecundity-based hierarchy among gamergates. In Stigmatomma reclinatum, it was found that higher-ranked gamergates had more fully developed oocytes than low-ranked gamergates. In Streblognathus peetersi, only the alpha worker mates and becomes the gamergate; younger workers await a chance to reproduce when the current gamergate exhibits decreased fecundity or dies. Challenges to gamergates from subordinate workers are risky because the gamergate in species like Dinoponera quadriceps may mark the challenger by rubbing special chemicals produced only by the gamergate. These chemicals signal to other workers to immobilize the challenger by biting her appendages and immobilizing her for a few days until her hormonal levels return to normal. Subordinate workers play an important policing role in the selection of future gamergates and are thus able to increase their indirect fitness.

Social structure variation and ecology

There is much variation in the social structure of ant colonies with gamergates. Some species such as Harpegnathos saltator, Pseudoneoponera tridentata, Gnamptogenys menadensis, and Rhytidoponera confusa have a winged alate queen caste as well as gamergates. Queenless species with only gamergates and workers may have a monogynous structure with a single gamergate or they may have a polygynous structure with multiple gamergates. Examples of monogynous queenless species include Pachycondyla krugeri, P. sublaevis, Diacamma australe, D. rugosum, Dinoponera quadriceps, Platythyrea lamellosa, and Streblognathus aethiopicus. Examples of polygynous queenless species include Ophthalmopone berthoudi, O. hottentota, and all known queenless species of Rhytidoponera. In the queenless Ophthalmopone berthoudi, foreign males visit underground nests to mate with young workers.

Ecologically, gamergate species from different tribes and genera often tend to share certain characteristics. Many gamergate species are solitary generalist foragers living in arid environments. Similar to species with ergatoid queens, the evolution of gamergate reproduction is hypothesized to be associated with a shift to colonial fission. Myrmecologists Christian Peeters and Fuminori Ito have also suggested that "the evolution of gamergate reproduction appears strongly associated with the adaptive benefits of secondary polygyny (e.g. increased colony lifespan and resource inheritance), and it is the preferred option in species having workers able to store sperm."

Classification dispute

The utility of gamergate as a morphological designation is not without critics. Within the field of myrmecology it is a matter of dispute whether caste should be defined primarily by reproductive role or by physical morphology. Notably, Alfred Buschinger has argued that the term worker should be applied only to those ants who make up the non-reproductive caste and queen should be applied only to reproductively viable female ants regardless of their physical appearance. Hölldobler and Wilson suggest that the two positions can be semantically resolved and that the most fruitful approach would be to keep classification "somewhat loose, incorporating either anatomy or roles in a manner that maximizes convenience, precision, and clarity of expression."

Genera with gamergates

The existence of gamergates is symplesiomorphic, or ancestral, in ants. They are known to be present in the following genera:

See also

Notes

  1. In the original Naturwissenschaften paper where the term "gamergate" was first used, O. berthoudi is referred to by the synonym Pachycondyla berthoudi.
  2. Throughout Ito's 1993 paper for the Journal of Natural History, he refers not to Stigmatomma but to Amblyopone. At the time Stigmatomma was considered to be a synonym of Amblyopone.

References

  1. Peeters, Christian; Crewe, Robin (1984). "Insemination Controls the Reproductive Division of Labour in a Ponerine Ant". Naturwissenschaften. 71 (1): 50–51. Bibcode:1984NW.....71...50P. doi:10.1007/BF00365989. ISSN 0028-1042. S2CID 43871631.
  2. ^ Véron, Géraldine (February 2005). "La reine des fourmis couronnée au combat". Le Journal du CNRS (181): 50–51. ISSN 0994-7647. Archived from the original on 10 February 2005. Retrieved 9 September 2014.
  3. ^ Barrows, Edward M. (2011). "Caste – Gamergate". Animal Behavior Desk Reference: A Dictionary of Animal Behavior, Ecology, and Evolution (Third ed.). CRC Press. p. 75. ISBN 9781439836514.
  4. Gordh, Gordon (2011). "Gamergate". A Dictionary of Entomology. CABI. p. 608. ISBN 9781845935429.
  5. ^ Noël, Carine (6 September 2002). "How queenless ants regulate their conflicts" (Press release). Paris: CNRS. Archived from the original on 2 March 2003. Retrieved 10 September 2014.
  6. ^ Choe, Jae C.; Crespi, Bernard J. (1997). "Morphologically 'Primitive' Ants". The Evolution of Social Behaviour in Insects and Arachnids. Cambridge University Press. p. 385. ISBN 9780521589772.
  7. ^ Bourke, Andrew F. G. (1995). "7 – Kin Conflict: Reproduction (Part 2 – Queen Policing, Queen Control, and Queen Signaling)". Social Evolution in Ants: Monographs in behavior and ecology. Princeton University Press. pp. 239–240. ISBN 9780691044262.
  8. Cournault, Laurent; Peeters, Christian (2012). "Aggression regulates monogyny in non-mutilating Diacamma ants". Insectes Sociaux. 59 (4): 533–539. doi:10.1007/s00040-012-0251-9. S2CID 16179257.
  9. Gorman, James (27 May 2014). "The Ant Queen Is Dead. Let the Battles Begin". The New York Times. ISSN 0362-4331. Retrieved 12 August 2014.
  10. Crespi, Bernard J. (1996). "9 – Comparative Analysis of the Origins and Losses of Eusociality: Causal Mosaics and Historical Uniqueness (Part 6 – Formicidae)". In Martins, Emília P. (ed.). Phylogenies and the Comparative Method in Animal Behavior. Oxford University Press. p. 272. ISBN 9780195092103.
  11. ^ Ito, Fuminori (1993). "Social organization in a primitive ponerine ant: queenless reproduction, dominance hierarchy and functional polygyny in Amblyopone sp. (reclinata group) (Hymenoptera: Formicidae: Ponerinae)". Journal of Natural History. 27 (6): 1315–1324. Bibcode:1993JNatH..27.1315I. doi:10.1080/00222939300770751. ISSN 0022-2933.
  12. Monnin, Thibaud; Ratnieks, Francis L. W.; Jones, Graeme R.; Beard, Richard (5 September 2002). "Pretender punishment induced by chemical signalling in a queenless ant" (PDF). Nature. 419 (6902): 61–65. Bibcode:2002Natur.419...61M. doi:10.1038/nature00932. ISSN 0028-0836. PMID 12214231. S2CID 4372841.
  13. Peeters, Christian; Hölldobler, Bert (November 1995). "Reproductive cooperation between queens and their mated workers: The complex life history of an ant with a valuable nest". Proceedings of the National Academy of Sciences of the United States of America. 92 (24): 10977–10979. Bibcode:1995PNAS...9210977P. doi:10.1073/pnas.92.24.10977. ISSN 0027-8424. PMC 40553. PMID 11607589.
  14. ^ Peeters, Christian; Ito, Fuminori (2001). "Colony Dispersal and the Evolution of Queen Morphology in Social Hymenoptera". Annual Review of Entomology. 46: 601–30. CiteSeerX 10.1.1.562.7031. doi:10.1146/annurev.ento.46.1.601. ISSN 0066-4170. PMID 11112181.
  15. ^ Peeters, Christian P. (1991). Veeresh, G. K.; Mallik, B. (eds.). Social Insects and the Environment: Proceedings of the 11th International Congress of IUSSI, 1990 (International Union for the Study of Social Insects). Brill Academic Pub. p. 234. ISBN 978-9004093164. Retrieved 12 August 2014.
  16. Peetere, C. P.; Crewe, R. M. (1985). "Queenlessness and reproductive differentiation in Ophthalmopone hottentota". South African Journal of Zoology. 20 (4): 268. doi:10.1080/02541858.1985.11447948.
  17. "Species: Pachycondyla berthoudi". antweb.org. AntWeb. Retrieved 12 August 2014.
  18. Hölldobler, Bert; Wilson, E. O. (1990). "Caste and Division of Labor". The Ants. Harvard University Press. pp. 301 & 305. ISBN 9780674040755.
  19. ^ Dietemann, V.; Peeters, C; Hölldobler, B. (2004). "Gamergates in the Australian ant subfamily Myrmeciinae". Naturwissenschaften. 91 (9): 432–435. Bibcode:2004NW.....91..432D. doi:10.1007/s00114-004-0549-1. ISSN 0028-1042. PMID 15278223. S2CID 26161972.
  20. Gobin, B.; Peeters, C.; Billen, J. (September 1998). "Production of trophic eggs by virgin workers in the ponerine ant Gnamptogenys menadensis" (PDF). Physiological Entomology. 23 (4): 329–336. doi:10.1046/j.1365-3032.1998.234102.x. S2CID 83569093. Archived from the original (PDF) on 27 August 2016. Retrieved 26 August 2016.
  21. Peeters, Christian P. (1987). "The Reproductive Division of Labour in the Queenless Ponerine Ant Rhytidoponera sp. 12". Insectes Sociaux. 34 (2): 75–86. doi:10.1007/BF02223826. ISSN 0020-1812. S2CID 26116621.
  22. ^ Schmidt, C. A; Shattuck, S. O. (2014). "The Higher Classification of the Ant Subfamily Ponerinae (Hymenoptera: Formicidae), with a Review of Ponerine Ecology and Behavior". Zootaxa. 3817 (1): 1–242. doi:10.11646/zootaxa.3817.1.1. PMID 24943802. S2CID 4540972.
  23. "Genus: Diacamma". antweb.org. AntWeb. Retrieved 12 August 2014.
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