Parasitic ant

A parasitic ant is a type of ant that exploits the social structure of another ant species to aid in its own survival and reproduction. The most common types of parasitic ants usually infiltrate a colony of a closely related species by using identical pheromones of the colony's workers to blend in and avoid conflict. The parasite then lays eggs alongside existing ones for the host colony's workers to raise and nurture as their own.
Parasites usually induce harmful effects on the target colony and can inhibit the colony's growth and development. Depending on the severity and quantity of parasites, the colony can experience extreme damage, sometimes to the point of colony collapse. Parasites evolve their anatomy to reflect that of their targeted species, which causes some to remain undetected inside of a colony for the majority of their lifespans. The parasites may also experience social parasitic syndrome, a series of changes to their anatomy during their evolution towards exploitative behaviors.
Species and characteristics
[edit]Of the 12,000 individual species of ants, 230 are considered parasites.[1] The genus Formica contains many parasitic species,[2] with over half of its 172 species confirmed or suspected to be parasites.[3] Other genera–such as Acromyrmex (leafcutter) ants and Myrmica red ants–also show parasitic behaviors.[4]
Eusociality
[edit]Typical ant colonies usually have at least three castes: queens, workers and drones (males).[5] Some species also have a fourth caste that consist of soldiers who defend the colony.[6] However, some socially parasitic ant species have developed a separate pseudo–caste of mutated worker ants that exhibit queen–like phenotypes.[7] The mutated ants contain a supergene that allows them to develop queen–like traits such as wings, larger eyes and ovaries.[7]
Features
[edit]Parasitic ant species can evolve over time to reflect the preferred host species. Rather than become visually homogeneous, such as matching their host's colors, they instead evolve similar anatomical shapes and features in order to blend in with the hosts through touch.[8] The parasitic queen's features are also significantly altered, with the vast majority of species' queens shrunken to at least 35% of a normal queen's size.[8]
The pheromones of parasitic ants are also altered to reflect the host species', usually through a process known as "rubbing".[9] The parasite uses its mandible to attach itself to a host ant and cover its own body with the host's pheromones. This is done so as to obtain the host's cuticular hydrocarbon profile, which is a collection of chemicals specific to each individual colony, and is used for identification among colony members.[10] Covering themselves with the host's profile can make the parasite indistinguishable from the host workers. This is done to minimize any potential conflict whilst the host colony is being entered.[9][1] The species Formicoxenus provancheri (shampoo ant) utilizes a slightly different method of imitating pheromones, in which they use their glossae (a tongue like structure) to obtain the host's pheromones, then use it to cover themselves.[11]
Methods
[edit]There are several methods that parasitic ants use to exploit other colonies.[11] A common method observed involves a parasitic queen ant infiltrating a colony and killing the existing queen, so as to become accepted by the hosts as a permanent member of the colony.[12] Another method observed in Teleutomyrmex schneideri involves the parasite clinging onto the body of a host ant and consuming nutrients from the colony off of the host's back.[13] Most parasites (excluding inquiline ants) eventually kill the host queen by either attacking the queen or killing off the workers who care for her.[1]
Slaver ants
[edit]
Slaver ants are a type of parasite that enter a host colony and transport their offspring back to their own colony in order to raise them as their own.[14] Around 50 individual ant species exhibit slave-making behavior.[15] After the host's offspring hatch inside the parasite colony, they are unable to differentiate between the parasitic species and their own, and therefore behave as workers for the slaver colony.[16]
Slaver colonies usually send out scouts to search for taxonomically similar species nearby in order to infiltrate the colony with minimal conflict from host workers, before returning back to the colony to prepare for an attack known as a slave raid.[17] They then launch a raid where they transport thousands of offspring in various stages of growth back to their own colony.[18] The slave-making ant's bodies are specialized to parasitize either a single species or a group of related species, and are often closely related taxonomically to their hosts.[19] The slave-makers may be either permanent social parasites or facultative slave-makers.[20][21] The permanent social parasites are unable to independently survive without a host to exploit, whereas facultative parasites are not reliant on a host but still engage in parasitic behaviors. Slave-making behavior is unusual among ants but has evolved several times independently.[22]
Temporary social parasites
[edit]Some parasitic ants are considered temporary social parasites, whose method of exploitation involves infiltrating a host colony and the parasitic queen killing the host queen, and then using the host workers to develop a new colony.[23][12] After the host queen is killed, the parasite queen then begins laying her own eggs among existing ones. The host workers then behave with the parasite as though it is their new queen.[1][24] They are considered temporary as they do not depend on their hosts after their new colony is established.[1] Temporary social parasitism has evolved 60 times in multiple ant species, marked by the parasites losing the ability to form their own colonies.[25]
Invading parasites typically disguise themselves with a similar chemical profile as the hosts through rubbing, which reduces conflict while they enter the colony.[26][27] Queen ants of some socially parasitic species such as Polyergus breviceps are unable to care for their own offspring and are entirely reliant on a host colony to raise their brood.[12]
Temporary social parasites eventually replace the population of the original host colony entirely. This process usually takes around 1 year for all of the individual ants born prior to the parasite attack to die off.[28] As the host queen is usually killed during the initial attack by the parasitic queen, the host colony is no longer able to lay new eggs and maintain their population. The existing workers are usually not attacked by the parasites, and are instead used to nurture and care for the parasitic brood until the host population eventually dies off.[28]
Guest ants
[edit]Some species of the genus Megalomyrmex, also known as parasitic guest ants, are social parasites that live among and consume the food and brood of fungus-growing ants.[29] These ants target fungus-growing ants as they do not have a soldier caste and their mandibles are significantly weaker than that of the parasites', making it difficult for the hosts to keep them out.[30] Guest ants do not mimic the hydrocarbon profile of their hosts. Instead, they possess an alkaloid-derived venom[31] that is used to pacify the host colony so they can consume the host's food and offspring with less resistance.[32] The venom can be detected by other ants outside of the host colony, so host ants often allow the parasites to reside inside of the nest in order to deter nearby hostile ant colonies despite their negative effects.[29]
Species
[edit]- Megalomyrmex adamsae[33]
- Megalomyrmex mondabora[33]
- Megalomyrmex symmetochus[33]
- Megalomyrmex wettereri[33]
Inquiline ants
[edit]
Inquiline ants are a type of social parasite that exhibit permanent social parasitism, and are considered the most severe form of parasitism in ants.[34] There are around 100 ant species that have evolved to exhibit inquiline behavior.[35] These ants are typically not hostile to the host colony and co-exist with the present host queen.[36] These ants live in their host colony for the vast majority of their lifespan.[36] Inquiline queens leave their home colony in search of a host one, leaving the queens workerless and forcing them to rely on the host colony's workers to provide for their grooming and feeding needs.[37] The ants practice polygyny with the male hosts,[35] with the males mating simultaneously with the parasite and the existing queen.[35] While they are not physically hostile to the hosts, the eggs they lay among the host colony's divert resources away from the host's, harming the fitness of the host's children.[38] The inquiline parasite's brood are almost always capable of reproduction.[34]
Almost all inquiline ants follow Emery's rule, which is a trend of social parasites that suggests social parasites should be closely related to each other taxonomically.[36][39] They usually follow either the strict form of Emery's rule (ants that are from the same genus) or the relaxed form (from the same or similar family).[39] The following of Emery's rule is one of the major distinctions of inquiline ants from temporary social parasites.[36]
Rodeo ants
[edit]Some species of Solenopsis (fire ants) may attach themselves to the backs of other ants in a parasitic fashion, where they can infiltrate a host colony easier.[40] These rodeo ants clamp onto the thorax of a host ant (usually the queen), and cover themselves with the pheromones of the ant in order to remain undetected by the rest of the colony.[41] All rodeo ants are queens and therefore lack workers to care for them, so the parasite lays her eggs off the back of the host ant alongside the host's brood, in an effort to convince the host colony to raise the parasite's offspring.[42] These ants have not been identified outside of the US state of Texas, and are the least common type of parasitic ant.[43]
Social parasitic syndrome
[edit]Social parasitic syndrome is a series of changes that can occur in parasitic ant species during their evolution towards exploitative behaviors. It is usually marked by the loss of a worker caste, shrinkage of body structures and overall size, broadening of the petioles, and a reduction in mandible size.[44] Queens ants affected by the syndrome contain a mutated supergene on chromosome 13, which causes increased egg production, altered metabolism and significantly shrunken bodies.[11] The supergene is a collection of 200 individual genes that contribute to the syndrome's effects.[11] The supergene is inherited in a single generation and is rarely not passed down to future generations.[45] Affected queens have a similar visual appearance in size and structure to workers of their own or closely related species, with the exception of wings.[45] After inheriting the supergene and being affected by the syndrome, the ants are then considered obligate parasites as they are no longer able to independently survive without exploiting a host colony.[45]
Effects on host colony
[edit]Depending on the method of parasitism, effects on the host colony can vary drastically. If a host colony is repeatedly subjected to slave raids, their ability to replenish their workers is significantly reduced and may eventually lead to colony collapse.[46] Colonies subjected to temporary social parasites may gradually experience a decline in their population as offspring from the new parasite queen replace the existing host workforce.[2][47] Parasitic queens do not productively contribute any resources or support to the host colony, and sometimes remove the wings of virgin potential queens.[48] The resources and care that are usually diverted towards a host colony's own brood are used for the parasite's offspring, which can lead to significantly less healthy and fit host worker ants.[38][49]
Parasites can cause an evolutionary arms race against their hosts, with colonies rapidly evolving to develop methods to attack and defend against each other.[50][30] Some parasites have developed a chemical toxin that is released from their dufour gland. This toxin can induce infighting among the host colony, in an attempt for the parasites to more easily infiltrate and exploit the host colony.[50] Chemicals used by permanent inquiline hosts can also influence the genes and behaviors of the host.[51]
Colony defense
[edit]When an infected host colony is raided by a separate colony, some parasites residing inside have been observed defending the hosts from the raiding colony.[51] Parasites release an alkaloid venom in order to help deter the attackers from beginning a raid.[51] They also use violent physical means to scare off or kill the raiders who ignore chemical deterrents.[52] Scouts from raiding colonies can detect whether or not a colony has social parasites within, and the presence of them can be enough to discourage the raid from beginning in the first place.[52]
Parasite–host pairs
[edit]- Rossomyrmex–Proformica[22]
- Polyergus–Formica[22]
- Formica–Formica[22]
- Leptothorax–Chalepoxenus[22]
- Leptothorax–Epimyrma[22]
- Leptothorax–Harpagoxenus[22]
- Leptothorax–Leptothorax[22]
- Leptothorax–Protomognathus[22]
- Myrmoxenus–Temnothorax[53]
- Strongylognathus–Tetramorium[22]
See also
[edit]References
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