Red imported fire ant

The red imported fire ant (Solenopsis invicta), also known as the fire ant or RIFA, is a species of ant native to South America. A member of the genus Solenopsis in the subfamily Myrmicinae, it was described by Swiss entomologist Felix Santschi as a variant in 1916. Its current name invicta was given to the ant in 1972 as a separate species. However, the variant and species were the same ant, but the name was preserved due to its wide use. The red imported fire ant is native to South America but it has been accidentally introduced in Australia, New Zealand, several Asian and Caribbean countries and the United States. The red imported fire ant is polymorphic as workers appear in different shapes and sizes. The ant's colours are red and somewhat yellowish with a brown or black gaster. However, males are completely black. Red imported fire ants are dominant in altered areas and live in a wide variety of habitats. They can be found in: rain forests, in disturbed areas, deserts, grasslands, alongside roads and buildings, and in electrical equipment. Colonies form large mounds constructed from dirt with no visible entrances because foraging tunnels are built and workers emerge far away from the nest.

These ants exhibit a wide variety of behaviours, such as building rafts when they sense that water levels are rising. They also show necrophoric behaviour, where nestmates discard scraps or dead ants on refuse piles outside the nest. Foraging takes place on warm or hot days, although they may remain outside at night. Workers communicate by a series of semiochemicals and pheromones which are used for recruitment, foraging, and defence. They are omnivores and eat dead mammals, arthropods, insects, seeds, and sweet substances such as honeydew from hemipteran insects with whom they have developed relationships. Predators include arachnids, birds and many insects including other ants, dragonflies, earwigs, and beetles. The ant is a host to parasites and to a number of pathogens, nematodes, and viruses which have been viewed as potential biological control agents. Nuptial flight occurs during the warm seasons, and the alates may mate for as long as 30 minutes. Colony founding can be done via a single queen or a group of queens, who will later contest for dominance once the first workers emerge. Workers can live for several months while queens can live for years; colony numbers can vary from 100,000 to 250,000 individuals. Two forms of society in the red imported fire ant exist: polygynous colonies (nests with multiple queens) and monogynous colonies (nests with one queen).

Venom plays an important role in the ant's life as it is used to capture prey or for defence. 95% of the venom's components are water-insoluble piperidine alkaloids, and it is particularly potent on sensitive humans. More than 14 million people are stung by them in the United States annually, where many are expected to develop allergies to the venom. Most victims experience intense burning and swelling, followed by the formation of sterile pustules which may remain for several days. However 0.6% to 6% of people may suffer from anaphylaxis which can be fatal if left untreated. Common symptoms include dizziness, chest pain, nausea, severe sweating, low blood pressure, loss of breath, and slurred speech. More than 80 deaths have been recorded from red imported fire ant attacks. Treatment depends on the symptoms; those who only experience pain and pustule formation require no medical attention, but those who suffer from anaphylaxis are given epinephrines. Whole body extract immunotherapy is used to treat victims and is regarded as highly effective.

The ant is viewed as a notorious pest, causing billions of dollars in damages annually and impacting wildlife. The ants thrive in urban areas, so their presence may deter outdoor activities. Nests can be built under structures such as pavements and foundations, which may cause structural problems, or cause them to collapse. Not only can they damage or destroy structures, red imported fire ants can damage equipment and infrastructure and impact business, land, and property values. As workers are attracted to electricity, they can swarm electrical equipment and destroy it. In agriculture, they can damage crops, damage machinery, and threaten pastures. They are known to invade a wide variety of crops, and mounds built on farmland may prevent harvesting. They also pose a threat to animals and livestock, capable of inflicting serious injury or killing them, especially weak or sick animals. Despite this, they may be beneficial because they consume common pest insects on crops. Common methods of controlling these ants include baiting and fumigation; other methods may be ineffective or dangerous. Due to its notoriety and importance, the ant has become one of the most studied insects on the planet, even rivalling the western honey bee (Apis mellifera).^[2][3]

Etymology and common names

The specific epithet of the red imported fire ant, invicta, derives from Latin, and means "invincible" or "unconquered".^[4][5][6] The epithet originates from the phrase Roma invicta ("unconquered Rome"), used as an inspirational quote until the fall of the Western Roman Empire in 476 AD. This symbolic statement was printed on minted coins.^[7] The generic name, Solenopsis, translates as "appearance" or "face" from Ancient Greek. It is a compound of two Ancient Greek words–solen, meaning "pipe" or "channel", and opsis, meaning "appearance" or "sight".^[8][9] The ant is commonly known as the "red imported fire ant" (abbreviated as RIFA), because of the burning sensation caused by its sting.^[10][11] Alternative names include: the "fire ant", "red ant" or "tramp ant".^[12][13] In Brazil, locals call the ant toicinhera, which derives from the Portuguese word toicinho (pork fat).^[14] The ants are called "fourmi de feu" and "rote importierte Feuerameise" in French and German.^[15]

Taxonomy

The red imported fire ant was first described by Swiss entomologist Felix Santschi in a 1916 journal article published by Physis.^[16] Originally named Solenopsis saevissima wagneri from a syntype worker collected from Santiago del Estero, Argentina, Santschi believed the ant was a variant of S. saevissima; the specific epithet, wagneri, derives from the surname of E.R. Wagner, who collected the first specimens.^[17] The type material is currently housed in Naturhistorisches Museum Basel, Switzerland, but additional type workers are possibly housed in the Muséum national d'histoire naturelle, Paris.^[18] In 1930, American myrmecologist William Creighton reviewed the genus Solenopsis and reclassified the taxon as Solenopsis saevissima electra wagneri at infrasubspecific rank, noting that he could not collect any workers that referred to Santschi's original description.^[19] In 1952, the S. saevissima species complex was examined and, together with nine other species-group names, S. saevissima electra wagneri was synonymised with S. saevissima saevissima.^[20] This reclassification was accepted by Australian entomologist George Ettershank in his revision of the genus and in Walter Kempf's 1972 catalogue of Neotropical ants.^[21][22]

In 1972, American entomologist William Buren described what he thought was a new species, naming it Solenopsis invicta.^[23] Buren collected a holotype worker from Cuiabá in Mato Grosso, Brazil and provided the first official description of the ant in a journal article published by the Georgia Entomological Society. He accidentally misspelled invicta as invica above the description pages of the species, although it was clear that invicta was the intended spelling because of the constant use of the name in the article.^[24] The type material is currently housed in the National Museum of Natural History, Washington, D.C.^[23]

In a 1991 review of the species complex, American entomologist James Trager synonymised S. saevissima electra wagneri and S. wagneri together.^[24] Trager incorrectly cites Solenopsis saevissima electra wagneri as the original name, erroneously believing that the name S. wagneri was unavailable and used Buren's name S. invicta. Trager previously believed that S. invicta was conspecific with S. saevissima until comparing the material with S. wagneri. Trager notes that even though S. wagneri has priority over S. invicta, the name was never used above infrasubspecific rank. The use of the name since Santschi has not been associated with collected specimens, and as a result is nomen nudum.^[24] In 1995, English myrmecologist Barry Bolton corrected Trager's error, recognising S. wagneri as the valid name and synonymised S. invicta.^[25] He states that Trager wrongfully classified S. wagneri as an unavailable name and cites S. saevissima electra wagneri as the original taxon. He concludes that S. wagneri is, in fact, the original name and has priority over S. invicta.^[25][26]

In 1999, Steve Shattuck and colleagues proposed conserving the name S. invicta.^[18] Since the first description of S. invicta, over 1,800 scientific papers using the name were published discussing a wide range of topics about its ecological behaviour, genetics, chemical communication, economic impacts, methods of control, population, and physiology. They state that the use of S. wagneri is a "threat" to nomenclatural stability towards scientists and non-scientists; taxonomists may have been able to adapt to such name change, but name confusion may arise if such case occurred. Due to this, Shattuck and his colleagues proposed the continued use of S. invicta and not S. wagneri, as this name has been rarely used; between 1995 and 1998, over 100 papers were published using S. invicta and only three using S. wagneri. They requested that the International Commission on Zoological Nomenclature (ICZN) use plenary powers to suppress S. wagneri for the purpose of the Principle of Priority and not for the Principle of Homonymy. Furthermore, they requested that the name S. invicta be added to the Official List of Specific Names in Zoology and that S. wagneri be added to the Official Index of Rejected Invalid Specific Names in Zoology.^[18] Upon review, the proposal was voted on by the entomological community and was supported by all but one voter. They note that there is no justification in suppressing S. wagneri; instead, it would be better to give precedence to S. invicta over S. wagneri whenever an author treated them as conspecific. The ICZN would conserve S. invicta and suppress S. wagneri in a 2001 review.^[27] Under the present classification, the red imported fire ant is a member of the genus Solenopsis in the tribe Solenopsidini, subfamily Myrmicinae. It is a member of the family Formicidae, belonging to the order Hymenoptera, an order of insects containing ants, bees, and wasps.^[28]

Phylogeny

The red imported fire ant is a member of the S. saevissima species-group. Members can be distinguished by their two-jointed clubs at the end of the funiculus in workers and queens, and the second and third segments of the funiculus are twice as long and broad in larger workers. Polymorphism occurs in all species and the mandibles bear four teeth.^[23] The following cladogram shows the position of the red imported fire ant among other members of the S. saevissima species-group:^{[lower-alpha 1][29]}

Solenopsis	Solenopsis geminata Solenopsis daguerrei Solenopsis quinquecupsis Solenopsis macdonaghi Solenopsis megergates Solenopsis invicta Solenopsis richteri Solenopsis interrupta Solenopsis altipunctata Solenopsis weyrauchi Solenopsis saevissima Solenopsis pythia Solenopsis electra Solenopsis pusillignis

Phenotypic and genetic data suggests that the red imported fire ant and the black imported fire ant (Solenopsis richteri) differentiate from each other, but they do share a close genetic relationship.^{[30][31][32][33]} Hybridisation between the two ants occurs in areas where they make contact, with the hybrid zone located in Mississippi. Such hybridisation has resulted from secondary contact between these two ants several decades ago, when they first encountered each other in southern Alabama.^[30][34] Based on mitochondrial DNA, examined haplotypes do not form a monophyletic clade. Some of the examined haplotypes form a closer relationship to S. megergates, S. quinquecuspis and S. richteri than they do with other S. invicta haplotypes. The occurrence of a possible paraphyletic grouping suggests that the red imported fire ant and S. quinquecuspis are possible cryptic species groups composed of several species that cannot be distinguished morphologically.^[33][35]

Genetics

A social chromosome is present in the red imported fire ant. This chromosome can differentiate the social organisation of a colony carrying one of two variants of a "supergene" (B and Bb) which contains more than 600 genes. For example, colonies exclusively carrying the B variant of this chromosome accept single BB queens, but colonies with multiple B variants will accept multiple Bb queens.^[36] Differences in another single gene can also determine whether the colony will have single or multiple queens.^[37] Studies show that mitochondrial DNA variation occurs substantially in polygyne societies (nests with multiple queens),^[38] but no variation is detected in monogyne societies (nests with a single queen).^[39] Triploidy occurs in red imported fire ants at high rates (as high as 12% in non-reproductive females), which is linked to the high frequency of diploid males.^[40] The red imported fire ant is the first species shown to possess a green-beard gene, by which natural selection can favour altruistic behaviour. Workers containing this gene are able to distinguish between queens containing it, and those that do not, apparently by using odour cues. The workers kill queens that do not contain the gene.^[41][42] In 2011, scientists announced they had fully sequenced the red imported fire ant genome from a male.^[43]

Description

Red imported fire ant workers range in size from small to medium, making them polymorphic. Workers measure between 2.4 to 6 millimetres (0.094 to 0.236 in).^[44] The head measures 0.66 to 1.41 millimetres (0.026 to 0.056 in) and is 0.65 to 1.43 millimetres (0.026 to 0.056 in) wide. In the larger workers (as in the major workers), their heads measure 1.35 to 1.40 millimetres (0.053 to 0.055 in) and 1.39 to 1.42 millimetres (0.055 to 0.056 in) wide. The antenna scapes measure 0.96 to 1.02 millimetres (0.038 to 0.040 in) and the thoracic length is 1.70 to 1.73 millimetres (0.067 to 0.068 in).^[23] The head becomes wider behind the eyes with rounded occipital lobes present, and unlike the similar-looking S. richteri, the lobes peak further than the midline, but the occipital excision is not as crease-like. The scapes in major workers do not extend beyond occipital peak by one or two scape diameters; this feature is more noticeable in S. richteri. In medium-sized workers, the scapes reach the occipital peaks and exceed the rear border in the smallest workers. In small and medium workers, the head tends to have more elliptical sides. The head of small workers is wider out front than it is behind.^[23] In the major workers, the pronotum does not have any angular shoulders, nor does it have any sunken posteromedian area. The promesonotum is convex and the propodeum base is rounded and also convex. The base and declivity are of equal length. The suture of the promesonotum is either strong or weak in larger workers. The petiole has a thick and blunt scale; if observed from behind, it is not as rounded above in contrast to S. richteri, and sometimes it may be subtruncate. The postpetiole is large and broad, and in the larger workers, it is broader than its length. The postpetiole tends to be less broad in front and broader behind. On the rear side of the dorsal surface, a transverse impression is present. In S. richteri, this feature is also present but much weaker.^[23]

The sculpture is very similar to S. richteri.^[23] The punctures are where pilosity arises from, and these are often elongated on the dorsal and ventral portions of the head. On the thorax, striae are present, but they are less engraved with fewer punctures than in S. richteri. On the petiole, the punctates are located on the sides. The postpetiole, when viewed above, has a strong shagreen with distinct transverse punctostriae. The sides are covered in deep punctures, where they appear smaller but deeper. In S. richteri, the punctures are larger and more shallow. This gives a more opaque appearance to the surface. In some cases, punctostriae may be present around the rear portion.^[23] The pilosity appears similar to that of S. richteri. These hairs are erect and vary in length, appearing long on each side of the pronotum and mesonotum; on the head, the long hairs are seen in longitudinal rows. There are numerous appressed pubescent hairs on the petiolar scale; this is the opposite in S. richteri, as these hairs are sparse.^[23] Workers appear red and somewhat yellowish with a brown or completely black gaster.^[16] Gastric spots are sometimes seen in larger workers, where they are not as brightly coloured as those in S. richteri. The gastric spot usually covers a small portion of the first gastric tergite. The thorax is concolorous, ranging from light reddish-brown to dark-brown. The legs and coxae are usually lightly shaded. The head has a consistent colour pattern in large workers, with the occiput and vertex appearing brown. Other parts of the head including the front, genae and the central region of the clypeus are either yellowish or yellowish brown. The anterior border of the genae and mandibles are dark-brown; they also both appear to share the same coloured shade with the occiput. The scapes and funiculi range from being the same colour as the head or shares the same shade with the occiput. Light coloured areas of the head in small to medium-sized workers is restricted to only the frontal region, with a dark mark resembling an arrow or rocket being present. On occasion, nests may have a series of different colours. For example, workers may be much darker, and the gastric spot may be completely absent or appear dark-brown.^[23]

Queens have a head length of 1.27 to 1.29 millimetres (0.050 to 0.051 in) and a width of 1.32 to 1.33 millimetres (0.052 to 0.052 in).^[23] The scapes measure 0.95 to 0.98 millimetres (0.037 to 0.039 in) and the thorax is 2.60 to 2.63 millimetres (0.102 to 0.104 in). The head is almost indistinguishable from S. richteri, but the occipital excision is less crease-like and the scapes are considerably shorter. Its petiolar scale is convex and resembles that of S. richteri. The postpetiole has straight sides that never concave, unlike in S. richteri where they concave. The thorax is almost identical, but the clear space between the metapleural striate area and propodeal spiracles is either a narrow crease or not present. The side portions of the petiole are punctate. The sides of the postpetiole are opaque with punctures present, but no irregular roughening is seen. The anterior of the dorsum is shagreen, and the middle and rear regions bear transverse puncto-striae. All these regions have erect hairs. The anterior portions of both the petiole and postpetiole have appressed pubescence that is also seen on the propodeum. The colour of the queen is similar to that of a worker: the gaster is dark brown and the legs, scapes and thorax are light brown with dark streaks on the mesoscutum. The head is yellowish or yellowish-brown around the central regions, the occiput and mandibles are a similar colour to the thorax, and the wing veins range from colourless to pale brown.^[23] Males appear similar to S. richteri, but the upper border of the petiolar scales are more concave. In both species, the postpetiole's and petiole's spiracles strongly project. The whole body of the male is concolorous black, but the antennae are whitish. Like the queen, the wing veins are colourless or pale brown.^[23]

The red imported ant can be misidentified as the similar looking S. richteri.^[23] The two species can be distinguished from each other through morphological examinations of the head, thorax and postpetiole. In S. richteri, the sides of the head are broadly elliptical-shaped and the cordate shape seen in the red imported fire ant is absent. The region of the occipital lobes that are situated nearby the midline and occipital excision appear more crease-like in S. richteri than it does in the red imported fire ant. The scapes of S. richteri are longer than they are in the red imported fire ant, and the pronotum has strong angulate shoulders. Such character is almost absent in the red imported fire ant. A shallow but sunken area is only known in the larger workers of S. richteri, which is located in the posterior region of the dorsum of the pronotum. This feature is completely absent in larger red imported fire ant workers. The red imported fire ant has a promesonotum that is strongly convex, whereas this feature is weakly convex in S. richteri. Upon examination the base of the propodeum is elongated and straight in S. richteri while convex and shorter in the red imported fire ant. It also has a wide postpetiole with either straight or diverging sides. The postpetiole in S. richteri is narrower with converging sides. In S. richteri, the transverse impression on the posterodorsal portion of the postpetiole is strong but weak or absent in the red imported fire ant.^[23] As well as that, S. richteri workers are 15% larger than red imported fire ant workers, are blackish-brown and they have a yellow stripe on the dorsal side of the gaster.^[45][46]

Brood

Eggs are tiny and oval-shaped, remaining the same size for approximately a week. After one week, the egg will assume the shape of an embryo and form as a larva when the egg shell is removed.^[47] Larvae measure 3 millimetres (0.12 in).^[48] They show a similar appearance to S. geminata larvae, but they can be distinguished by the integument with spinules on top of the dorsal portion of the posterior somites. The body hairs measure 0.063 to 0.113 millimetres (0.0025 to 0.0044 in) with a denticulate tip. The antennae both have two or three sensilla. The labrum is smaller with two hairs on the anterior surface that are 0.013 millimetres (0.00051 in). The maxilla has a sclerotised band between the cardo and stipes. The labium also has a small sclerotised band.^[48] The tubes of the labial glands are known to produce or secrete a proteinaceous substance that has a rich level of digestive enzymes, which includes proteases and amylases that function as an extraintestinal digestion of solid food. The midgut also contains amylases, roteases and upases. The narrow cells in its reservoir have little to no function in secretion.^[49] The pupae resemble adults of any caste, except that their legs and antennae are held tightly against the body. They appear white, but over time, the pupae turns darker when they are almost ready to mature.^[50]

Four larval instars have been described based on their morphological characters.^[47][51] With the exception of size differences, the larvae of the minor and major workers are almost impossible to distinguish. Major worker 4th instar larvae tend to be larger than minor worker pre-pupae with a wider width. Only the reproductive larvae are larger than major workers and in general, are very robust. Even male 3rd instar larvae tend to be longer and larger than 3rd instar workers. No significant morphological difference can distinguish male and queen larvae, but the internal gonopodal imaginal discs differ. Most larval changes are not significant, but the moulting of 3rd instar larvae and studies of their eclosion support four larval instar stages.^[47]

Polymorphism

The red imported fire ant is polymorphic with two different castes of workers: minor workers and major workers (soldiers). Like many ants that exhibit polymorphism, young, smaller ants do not forage and tend to the brood instead, while the larger workers go out and forage.^{[52][53][54][55]} In incipient colonies, polymorphism does not exist, but instead they are occupied by monomorphic workers called "minims" or "nanitics". The average head-width in tested colonies increases during the first six months of development.^[56][57] In five-year old colonies, the head width of minor workers decreases but for major workers the head-width remains the same. The total weight of a major worker is twice that of a minor worker when they first arrive, and by six months, major workers are four times heavier than minor workers. Once major workers develop, they can make up a large portion of the workforce, with as many as 35% being major workers in a single colony.^[56] This does not affect colony performance, as polymorphic colonies and nests with small workers produce broods at roughly the same rate, and polymorphism is not an advantage or disadvantage when food sources are not limited. However, polymorphic colonies are more energetically efficient, and under conditions where food is limited, polymorphism may provide a small advantage in brood production but this depends on the levels of food stress.^[58]

One study shows that as worker ants grow to larger sizes, the shape of the head changes. This is due to the head length growing at the same time as the total body length, and the head width may grow by 20%. The length of the antennae only grows slowly; the antennae may only grow 60% longer by the time the body doubles its length, thus the relative antennal length decreases by 20% as the length of the body doubles.^[57] All individual legs of the body are isometric with body length meaning that even when the length of the body doubles, the legs will also double. However, not all of the legs are the same length; the prothoracic portion accounts for 29% of leg length, the mesothoracic 31%, and the metathoracic 41%. The first two pairs of legs are of equal length to one another, whereas the final pair is longer.^[57] Overall, the morphological appearance of a worker changes dramatically when it grows larger. The head exhibits the greatest shape change and the height of the alinotum grows quicker than its length, where there is a height/length ratio of 0.27 in minor workers and 0.32 in major workers.^[57] Due to this, larger workers tend to have a humped-shape and robust alinotum in contrast to smaller workers. No petiole segment exhibits any change in shape as the size of the body changes. The width of the gaster grows more rapidly than its length, where the width may be 96% of its length but increases to 106%.^[57]

Physiology

Like other insects, the red imported fire ant breathes through a system of gas-filled tubes called tracheae connected to the external environment through spiracles. The terminal tracheal branches (tracheoles) make direct contact with internal organs and tissue. The transport of oxygen to cells (and carbon dioxide out of cells) occurs through diffusion of gases between the tracheoles and the surrounding tissue and is assisted by a discontinuous gas exchange.^[59] As with other insects, the direct communication between the tracheal system and tissues eliminates the need for a circulating fluid network to transport O₂.^[60] Thus, red imported fire ants and other arthropods can have a modest circulatory system though they have highly expensive metabolic demands.^[61]

The excretory system consists of three regions. The basal region has three cells found within the posterior portion of the midgut. The anterior and superior cavity are formed by the bases of four malpighian tubules.^[62] The superior cavity opens into the lumen of the small intestine. The rectum is a large but thin-walled sac that occupies the posterior fifth of the larvae. The release of waste is controlled by the rectal valves that lead to the anus.^[62] Sometimes, the larvae secrete a liquid that consists of uric acid, water and salts.^[63] These contents are often carried outside by workers and ejected, but colonies under water stress may consume the contents.^[62] In the reproductive system, queens release a pheromone that prevents dealation and oogenesis in virgin females; those tested in colonies without a queen begin oocyte development after dealation and take up the egg-laying role.^[64] Flight muscle degeneration is initiated by mating and juvenile hormones, and prevented by corpus allatectomy.^[65][66] Histolysis begins with the dissolution of the myofibril and the slow breakdown of the myofilaments. Such dissolution continues until it reaches the only free Z-line materials, which would also disappear; only the nuclei and lamellar bodies remain.^[62] In one study, the amino acids increase in the hemolymph after insemination.^[67]

The glandular system contains four glands: the mandibular, maxillary, labial and post-pharyngeal glands.^[62] The post-pharyngeal is well developed in the queen while the other glands are larger in workers. The post-pharyngeal gland functions as a vacuum to absorb fatty acids and triglycerides, as well as a gastric caecum.^[68] The functions of the other glands remain poorly understood. In one study discussing the enzymes of the digestion system of adult ants, lipase activity was found in the mandibular and labial glands, as well as invertase activity. The Dufour's gland is found in the ant and acts as a source of trail pheromones, although scientists believed the poison gland was the source of the queen pheromone.^[62][69][70] The neurohormone pheromone biosynthesis activating neuropeptide (PBAN) is found in the ant that activates the biosynthesis of pheromones from the Dufour's gland.^[71] The spermatheca gland is found in queens, which functions in sperm maintenance. Males appear to lack these glands, but those associated with its head are morphologically similar to those found in workers, but these glands may act differently.^[68]

The ant faces many respiratory challenges due to its highly variable environment, which can cause increased desiccation, hypoxia, and hypercapnia. Hot, humid climates cause an increase in heart rate and respiration which increases energy and water loss.^[60][72] Hypoxia and hypercapnia can result from red imported fire ant colonies living in poorly ventilated thermoregulatory mounds and underground nests. Discontinuous gas exchange (DGE) may allow ants to survive the hypercapnic and hypoxic conditions frequently found in their burrows;^[59] it is ideal for adapting to these conditions because it allows the ants to increase the period of O₂ intake and CO₂ expulsion independently through spiracle manipulation. The invasion success of the red imported fire ant may possibly be related to its physiological tolerance to abiotic stress, being more heat tolerant and more adaptable to desiccation stress than S. richteri. This means that the ant is less vulnerable to heat and desiccation stress. Although S. richteri has higher water body content than the red imported fire ant, S. richteri was more vulnerable to desiccation stress. The lower sensitivity to desiccation is due to a lower water loss rate.^[73] Colonies living in unshaded and warmer sites tend to have a higher heat tolerance than those living in shaded and cooler sites.^[74]

Metabolic rate, which indirectly affects respiration, is also influenced by environmental temperature. Peak metabolism occurs at about 32 °C.^[75] Metabolism, and therefore respiration rate, increases consistently as temperature increases. DGE stops above 25 °C, although the reason for this is currently unknown.^[76]

Respiration rate also appears to be influenced significantly by caste. Males show a considerably higher rate of respiration than females and workers, due, in part, to their capability for flight and higher muscle mass. In general, males have more muscle and less fat, resulting in a higher metabolic O₂ demand.^[76] While the metabolic rate is highest at 32 °C, colonies often thrive at slightly cooler temperatures (around 25 °C). The high rate of metabolic activity associated with warmer temperatures is a limiting factor on colony growth because the need for food consumption is also increased. As a result, larger colonies tend to be found in cooler conditions because the metabolic demands required to sustain a colony are decreased.^[75]

Distribution and habitat

Red imported fire ants are native to the tropical areas of Central and South America, where they have an expansive geographical range that expands from southeastern Peru to central Argentina, and to the south of Brazil.^{[77][78][79][80]} In contrast to its geographical range in North America, its range in South America is significantly different. It has an extremely long north-south range but a very narrow east-west distribution. The northernmost record of the red imported fire ant is Porto Velho in Brazil and its southernmost record is Resistencia in Argentina; this is a distance of about 3,000 kilometres (1,900 mi). In comparison, the width of its narrow range is about 350 kilometres (220 mi), and this is most likely narrower into southern Argentina and Paraguay and into the northern areas of the Amazon River basin.^[81] Most known records of the red imported fire ant are around the Pantanal region of Brazil. However, the interior of this area has not been examined thoroughly, but it is certain that the species occurs in favourable locations around it. The Pantanal region is hypothesised to be the original homeland of the red imported fire ant; hydrochore dispersal via floating ant rafts could easily account for the far south populations around the Paraguay and Guaporé Rivers.^[81] The western extension of its range is not known exactly, but its abundance there may be limited. It may be extensive in easternmost Bolivia, owing to the presence of the Pantanal region. It is also unknown why the ant has penetrated farther into the east; in areas where related Solenopsis ants are found, the red imported is excluded from these areas, making these not entirely insalubrious. Red imported fire ants have not been found in the cerrado area to the east of the Pantanal, where scientists have suggested that its absence is due to the lack of moisture during the dry seasons. They have also not been found in the Paraná River while other Solenopsis ants have been, but this may be due to competition with dominant ants such as Pheidole.^[81] In areas it is native, red imported fire ants prefer to inhabit rainforests, even if these sites are at risk of inundation.^[80]

These ants are native to Argentina, and sources suggest that the red imported fire ant most likely came from here when they first invaded the United States; in particular, populations of these ants have been found in the provinces of Chaco, Corrientes, Formosa, Santiago del Estero, Santa Fe, and Tucumán.^[24][81][82] The northeastern regions of Argentina are the most credible guess where the invading ants originate.^[79] In Brazil, they are found in northern Mato Grosso and in Rondônia and in São Paulo state. The red imported fire ant and S. saevissima are parapatric in Brazil, with contact zones known in Mato Grosso do Sul, Paraná state and São Paulo.^[78][83] In Paraguay they are found throughout the country, and have been recorded in Boquerón, Caaguazú, Canindeyú, Central, Guairá, Ñeembucú, Paraguarí, and Presidente Hayes departments; Trager claims that the ant is distributed in all regions of the country.^[84][85][86] They are also found in a large portion of northeastern Bolivia and, to a lesser extent, in northwestern Uruguay.^[81][87]

The red imported fire ant is able to dominate altered areas and live in a variety of habitats. It can survive the extreme weather of the South American rain forest, and in disturbed areas nests are seen frequently alongside roads and buildings.^[88][89] The ant has been observed frequently around the floodplains of the Paraguay River.^[90] In areas where water is present, they are commonly found around: irrigation channels, lakes, ponds, reservoirs, rivers, streams, riverbanks and, mangrove swamps.^[84][88] Nests are found in: agricultural areas, coastlands, wetlands, coastal dune remnants, deserts, forests, grasslands, natural forests, oak woodland, mesic forest, leaf-litter, beach margins, shrublands, alongside rail and roads and in urban areas.^[15] In particular, they are found in cultivated land, managed forests and plantations, disturbed areas, intensive livestock production systems, and greenhouses.^[88][91] Red imported fire ants have been found to invade buildings, including medical facilities.^[92] In urban areas, colonies dwell in open areas, especially if the area is sunny.^[92] This includes: urban gardens, picnic areas, lawns, playgrounds, schoolyards, parks, and golf courses.^[84][92] In some areas, there are on average 200 mounds per acre.^[93] During winter, colonies move under pavements or into buildings, and newly mated queens move into pastures.^[88][92] Red imported fire ants are mostly found at altitudes of between 5 to 145 m (16 to 476 ft) above sea level.^[84]

Mounds range from small to large, measuring 10 to 60 cm (3.9 to 23.6 in) in height and 46 cm (18 in) in diameter with no visible entrances.^[88][94] Workers are only able to access their nests by a series of subterranean tunnels that protrude from the central region. Constructed from dirt, mounds are oriented so that the long portions of the mound face toward the sun during the early morning and before sunset.^[88][94] Mounds tend to be oval-shaped with the long axis of the nest orientating itself in a north-south direction.^[95] These ants also spend large amounts of energy in nest construction and transporting brood, which is related with thermoregulation. The brood is transported to areas where temperatures are high; not only does mound temperature increase due to the sun, the temperature is greater than the ground heat if a mound is shaded. With this said, workers track temperature patterns and do not rely on behavioural habits.^[96] Inside nests, mounds contain a series of narrow horizontal tunnels, with subterranean shafts and nodes reaching grass roots 10 to 20 centimetres (3.9 to 7.9 in) below the surface; these shafts and nodes connect the mound tunnels to the subterranean chambers. These chambers are approximately 5 cm² (0.77 inch²) and reach depths of 10 to 80 centimetres (3.9 to 31.5 in). The mean number of ants in a single subterranean chamber is around 200 individuals.^[97][98][99] Aside from mounds, nests may be found under logs, sidewalks, walls in buildings and electric and water utility boxes.^[50] Red imported fire ants and S. geminata tend to be difficult to distinguish; both species form large colonies and defend their territories, build galleried mounds, forage through underground tunnels, and have workers approximately the same size. S. geminata can be distinguished by its smaller colony population, preference towards wooded areas, and differentiated diet.^[100] In areas where the red imported fire ant and S. xyloni meet, S. xyloni nests can be distinguished by their irregular shape with obscure entrances and scattered soil around the nest.^[50]

Introductions

Red imported fire ants are among the worst invasive species in the world.^[101][102] However, some scientists would consider the red imported fire ant to be a "disturbance specialist"; this is because human disturbance to the environment may be a major factor behind the ants' impact (fire ants tend to favour disturbed areas). This is shown through one experiment, demonstrating that mowing and plowing in studied areas almost diminished the diversity and abundance of native ant species, whereas red imported fire ants found on undisturbed forest plots had only diminished a couple of species.^[103][104]

In the United States, the red imported fire ant first arrived in the seaport of Mobile, Alabama by cargo ship between 1933 and 1945.^{[lower-alpha 2][lower-alpha 3][81][107][108][109]} Arriving with an estimated 9 to 20 unrelated queens,^[110][111] The red imported fire ant was only rare at the time, as entomologists were unable to collect any specimens. The earliest observations of these ants were by E.O. Wilson in 1942, and the population expansion most likely occurred after 1937; the population of these ants exploded by the 1950s.^{[112][113][114]} Since its introduction to the United States, the red imported fire ant has spread throughout the southern states and the north-east of Mexico, negatively affecting wildlife and causing economic damage.^{[77][115][116][117]} The expansion of red imported fire ants may be limited since they are almost wiped out during Tennessee winters, and thus they may be reaching their northernmost range.^{[118][119][120]} However, global warming may allow the red imported fire ant to expand its geographical range.^[121] As of 2004, the ant is found in 13 states and occupies over 128 million hectares of land, and as many as 400 mounds can be found on a single acre of land.^[122][123] The United States Department of Agriculture estimates that they expand 193 kilometres (120 miles) per year.^[92]

Red imported fire ants were first discovered in Queensland, Australia in 2001.^[124][125] The ants were believed to be present in shipping containers arriving at the Port of Brisbane, most likely from North America.^[126] Anecdotal evidence suggests fire ants may have been present in Australia for six to eight years prior to formal identification. The potential damage from the red imported fire ant prompted the Australian government to respond rapidly. The initial emergency response was followed by the formation of the Fire Ant Control Centre in September 2001. A joint state and federal funding of A$175 million was granted for a six-year eradication programme involving the employment of more than 600 staff.^{[127][128][129]} Following years of eradication, eradication rates of greater than 99% from previously infested properties were reported. The program received extended Commonwealth funding of around A$10 million for at least another two years, until June 2009, to treat the residual infestations found most recently.^[130] In December 2014, a nest was identified at Port Botany, Sydney, in New South Wales. The port has been quarantined, and a removal operation took place.^[131] In September 2015, populations originating from the United States were found at a Brisbane airport.^[132]

Red imported fire ants have spread beyond North America. The Invasive Species Specialist Group (ISSG) reports the ant inhabiting from three of the Cayman Islands. However, the sources the ISSG cited give no report about them on the island, but recent collections indicate that they are present.^[77] In 2001, red imported fire ants were discovered in New Zealand, but they were successfully eradicated several years later.^[77][133] Red imported fire ants have been reported in India,^[134] Malaysia,^[135] the Philippines^[136] and Singapore.^[77] However, these reports were found to be incorrect as the ants collected there were incorrectly identified as the red imported fire ant. In Singapore, the ants were most likely misidentified as well. In India, surveyed ants in Sattur Taluk, India listed the red imported fire ant there in high populations; meanwhile, there were no reports of the ant outside the surveyed area. The reports in the Philippines most likely misidentified collected material as the red imported fire ant, as no populations have been found there.^[77] It was discovered in Hong Kong and mainland China in 2004, where they have spread into several provinces as well as Macau and Taiwan in 2005.^{[137][138][139][140]} The ants in China most likely appeared in 1995, where it has spread quickly; these ants most likely arrived from the United States. No geographic or climatic barriers prevent these ants from spreading further, thus it may spread throughout the tropical and subtropical regions of Asia.^[122][141] In Europe, a single nest was found in the Netherlands in 2002.^[80]

Around 1980, red imported fire ants began spreading throughout the West Indies where they were first reported in Puerto Rico and the U.S. Virgin Islands.^[142][143] Between 1991 and 2001, the ant was recorded from Trinidad and Tobago, several areas in the Bahamas, the British Virgin Islands, Antigua, and the Turks and Caicos Islands.^{[77][144][145]} Since then, red imported fire ants have been recorded on more islands and regions, with new populations discovered in: Anguilla, Saint Martin, Barbuda, Montserrat, Saint Kitts, Nevis, Aruba, and Jamaica.^[77] The ants recorded from Aruba and Jamaica have only been found on golf courses; these courses import sod from Florida, so such importation may be an important way for the ant to spread throughout the West Indies.^[77]

Populations found outside North America originate from the United States. In 2011, the DNA of specimens from Australia, China and Taiwan was analysed with results showing that they are related to those in the United States.^[146] Despite its spread, S. geminata has a greater geographical range than the red imported fire ant, but it can be easily displaced by it. Because of this, almost all of its exotic range in North America has been lost and the ant has almost disappeared there. On roadsides in Florida, 83% of these sites had S. geminata present when the red imported fire ant was absent, but only 7% when it is present.^[77] This means that the ant can probably invade many tropical and subtropical regions where S. geminata populations are present.^[77]

Behaviour and ecology

Red imported fire ants are extremely resilient and have adapted to contend with both flooding and drought conditions. If the ants sense increased water levels in their nests, they link together and form a ball or raft that floats, with the workers on the outside and the queen inside.^{[147][148][149]} The brood is transported to the highest surface.^[150] They are also used as the founding structure of the raft, except for the eggs and smaller larvae. Before submerging, the ants will tip themselves into the water and sever connections with the dry land. In some cases, workers may deliberately remove all males from the raft, resulting in drowning. The longevity of a raft can be as long as 12 days. Ants that are trapped underwater escape by lifting themselves to the surface using bubbles which are collected from submerged substrate.^[150] Owing to their greater vulnerability to predators, red imported fire ants are significantly more aggressive when rafting. Workers tend to deliver higher doses of venom, which reduces the threat of other animals attacking. Due to this, and the fact that a higher workforce of ants is available, rafts are potentially dangerous to those that encounter them.^[151]

Necrophoric behaviour occurs in the red imported fire ant. Workers discard uneaten food and other such wastes away from the nest. The active component was not identified, but the fatty acids accumulating as a result of decomposition were implicated and bits of paper coated with synthetic oleic acid typically elicited a necrophoric response. The process behind this behaviour in imported red fire ants was confirmed by Blum (1970): unsaturated fats, such as oleic acid, elicit corpse-removal behaviour.^[152] Workers also show differentiated responses towards dead workers and pupae. Dead workers are usually taken away from the nest, whereas the pupae may take a day for a necrophoric response to occur. Pupae infected by Metarhizium anisopliae are usually discarded by workers at a higher rate; 47.5% of unaffected corpses are discarded within a day, but for affected corpses this figure is 73.8%.^[153]

Foraging and communication

Colonies of the red imported fire ant have tunneling surfaces that protrude out of the surfaces where workers forage.^[154][155] These areas of protrusion tend to be within their own territory, but greater ant colonisation can affect this.^[156] Tunnels are designed to allow effective body, limb and antennae interactions with walls, and a worker can also move exceptionally fast inside them (more than nine bodylengths per second).^[157] The holes exit out of any point within the colony's territory, and foraging workers may need to travel half a metre to reach the surface. Assuming the average forager travels five metres, over 90% of foraging time is inside the tunnels during the day and rarely at night. Workers forage in soil temperatures reaching 27 °C (80 °F) and surface temperatures of 12–51 °C (53–123 °F).^[155] Workers exposed to temperatures of 42 °C (107 °F) are at risk of dying from the heat.^[155] The rate of workers foraging drops rapidly by autumn, and they rarely emerge during winter. This may be due to the effects of soil temperature, and a decreased preference for food sources. These preferences only decrease when there is low brood production. In the northern regions of the United States, areas are too cold for the ant to forage, but in other areas such as Florida and Texas, foraging may occur all year round. When it is raining, workers do not forage outside, as exit holes are temporarily blocked, pheromone trails are washed away, and foragers may be physically struck by the rain. The soil's moisture may also effect the foraging behaviour of workers.^[155]

When workers are foraging, it is characterised by three steps: searching, recruitment and transportation.^[158] Workers tend to search for honey more often than other food sources, and the weight of food has no impact on searching time. Workers may recruit other nestmates if the food they have found is too heavy, taking as much as 30 minutes for the maximum number of recruited workers to arrive. Lighter food sources take less time and are usually transported rapidly.^[158] Foraging workers become scouts and search solely for food outside the surface, and may subsequently die two weeks later from old age.^[159]

Workers communicate by a series of semiochemicals and pheromones. These communication methods are used in a variety of activities, such as nestmate recruitment, foraging, attraction, and defence; for example, a worker may secrete trail pheromones if a food source it discovered is too large to carry.^[160] These pheromones are synthesized by the Dufour's gland and may trail from the discovered food source back to the nest.^[161][162] The components in these trail pheromones are also species-specific to this ant only, in contrast to other ants with common tail pheromones.^[163] The poison sack in this species has been identified as being the novel storage site of the queen pheromone; this pheromone is known to elicit orientation in worker individuals, resulting in the deposition of brood.^[70][164] It is also an attractant, where workers aggregate toward areas where the pheromone has been released.^[160] A brood pheromone is possibly present, as workers are able to segregate brood by their age and caste, which is followed by licking, grooming and antennation.^[160] If a colony is under attack, workers will release alarm pheromones.^[165] However, these pheromones are poorly developed in workers. Workers can detect pyrazines which are produced by the alates; these pyrazines may be involved in nuptial flight, as well as an alarm response.^[166]

Red imported fire ants can distinguish nestmates and non-nestmates through chemical communication and specific colony odours.^[160][167] Workers prefer to dig into nest materials from their own colony and not from soil in unnested areas or from other red imported fire ant colonies. One study suggests that as a colony's diet is similar, the only difference between nested and unnested soil was the nesting of the ants themselves. Therefore, it is possible that workers transfer colony odour within the soil.^[167] Colony odour can be affected by the environment, as workers in lab-reared colonies are less aggressive than those in the wild.^[168] Queen-derived cues are able to regulate nestmate recognition in workers and amine levels. However, these cues do not play a major role in colony-level recognition, but they can serve as a form of caste-recognition within nests.^[169][170] Workers living in monogyne societies tend to be extremely aggressive and attack intruders from neighbouring nests. In queenless colonies, the addition of alien queens or workers does not increase aggression among the population.^[171]

Diet

Red imported fire ants are omnivores, and foragers are considered to be scavengers rather than predators.^[172] The ants' diet consists of dead mammals, arthropods,^[173] insects, earthworms, vertebrates, and solid food matter such as seeds. However, this species prefers liquid over solid food. The liquid food the ants collect is sweet substances from plants or honeydew-producing hemipterans.^{[94][172][174][175]} Arthropod prey may include dipteran adults, larvae and pupae, and termites. The consumption of sugar amino acid is known to affect recruitment of workers to plant nectars. Mimic plants with sugar rarely have workers to feed on them, whereas those with sugar and amino have considerable numbers.^[176][177] The habitats they live in may determine the food they collect the most; for example, forage success rates for solid foods are highest in lake shore sites, while high levels of liquid sources were collected from pasture sites.^[178] Specific diets can also alter the growth of a colony, with laboratory colonies showing high growth if fed honey-water. Colonies that feed on insects and sugar-water can grow exceptionally large in a short period of time whereas those that do not feed on sugar-water grow substantially slower. Colonies that do not feed on insects cease brood production entirely.^[175] Altogether, the volume of food digested by nestmates is regulated within colonies.^[179] Larvae are able to display independent appetites for sources such as solid proteins, amino acid solutions, and sucrose solutions, and they also prefer these sources over dilute solutions. Such behaviour is due to their capability to communicate hunger to workers. The rate of consumption depends on the type, concentration, and state of the food they feed on. Workers tend to recruit more nestmates to food sources filled with high levels of sucrose than to protein.^[179]

Food distribution plays an important role in a colony. This behaviour varies in colonies, with small workers receiving more food than larger workers if a small colony is seriously deprived of food. In larger colonies, however, the larger workers receive more food. Workers can donate sugar water efficiently to other nestmates, with some acting as donors. These "donors" distribute their food sources to recipients, who may also act as donors. Workers may also share a greater portion of their food with other nestmates.^[180] In colonies that are not going through starvation, food is still distributed among the workers and larvae. One study shows that honey and soybean oil were fed to the larvae after 12 to 24 hours of being retained by the workers. The ratio distribution of these food sources was 40% towards the larvae and 60% towards the worker for honey, and for soybean oil this figure was around 30% and 70% respectively.^[181] Red imported fire ants also stockpile specific food sources such as insect pieces rather than consuming them immediately. These pieces are usually transported below the mound surface and in the driest and warmest locations.^[182]

This species engages in trophallaxis with the larvae.^[183] Regardless of the attributes and conditions of each larva, they are fed roughly the same amount of liquid food. The rate of trophallaxis may increase with larval food deprivation, but such increase depends on the size of each larva. Larvae that are fed regularly tend to be given small amounts. In order to reach satiation, all larvae regardless of their size generally require the equivalent of eight hours of feeding.^[184]

Predators

A number of insects, arachnids, and birds prey on these ants, especially when queens are trying to establish a new colony.^[185] Many species of dragonfly including Anax junius, Pachydiplax longipennis, Somatochlora provocans and Tramea carolina, capture the queens while they are in flight. 16 species of spiders including the wolf spider Lycosa timuga and the southern black widow spider (Latrodectus mactans) actively kill red imported fire ants. L. mactans captures all castes of the species (the workers, queens and males) within its web. These ants constitute 75% of prey captured by the spider. Juvenile L. mactans spiders have also been seen capturing the ants.^[185][186] Other invertebrates that prey on red imported fire ants are earwigs (Labidura riparia) and tiger beetles (Cicindella punctulata).^[185] Birds that eat these ants include the chimney swift (Chaetura pelagica), the eastern kingbird (Tyrannus tyrannus), and the eastern bobwhite (Colinus virginianus virginianus). The eastern bobwhite attacks these ants by digging out the mounds looking for young queens.^[185]

Many species of ants have been observed attacking queens and killing them. Predatory ants include: Ectatomma edentatum, Ephebomyrmex, Lasius neoniger, Pheidole, Pogonomyrmex badius, and Conomyrma insana, which is among the most significant.^{[185][187][188]} C. insana ants are known to be effective predators against founding queens in studied areas of Northern Florida. The pressure of attacks initiated by C. insana increase over time causing queens to exhibit different reactions including escaping, concealment, or defence. Most queens that are attacked by these ants are ultimately killed.^[187] Queens that are in groups have higher chances of survival than solitary queens if they are attacked by S. geminata.^[189] Ants can attack queens on the ground and invade nests by stinging and dismembering them. Other ants such as P. porcula try to take the head and gaster, and C. clara invade in groups. Also, certain ants try to drag queens out of their nests by pulling on the antennae or legs.^[188] Small, monomorphic ants rely on recruitment to kill queens and do not attack them until reinforcements arrive. Aside from killing the queen, some ants may steal the eggs for consumption or emit a repellent that is effective against red imported fire ants.^[188] Certain ant species may raid colonies and destroy them.^[190]

Parasites, pathogens and viruses

Flies in the genus Pseudacteon (phorid flies) are known to be parasitic to ants. These flies are parasitoids of the red imported fire ant in its native range in South America, and can be attracted through the ants' venom alkaloids.^[191] One species, Pseudacteon obtusus, attacks the ant by landing on the posterioral portion of the head and laying an egg. The location of the egg makes it impossible for the ant to successfully remove it.^[192] The larvae migrate to the head, then develop by feeding on the hemolymph, muscle tissue, and nervous tissue. After about two weeks, they cause the ant's head to fall off by releasing an enzyme that dissolves the membrane attaching the head to its body. The fly pupates in the detached head capsule, emerging two weeks later.^[192][193] P. tricuspis is another phorid fly that is a parasitoid to this species. Although parasitism pressures by these flies do not affect the ants' population density and activity, it has a small effect on a colony population.^[194] The strepsipteran insect Caenocholax fenyesi is known to infect male ants of this species and attack the eggs,^[195][196] and the mite Pyemotes tritici has been considered a potential biological agent against red imported fire ants, capable of parasitising every caste within the colony.^[197] Bacteria, such as Wolbachia, has been found in the red imported fire ant; three different variants of the bacteria are known to infect the red imported fire ant. However, its effect on the ant is unknown.^[198][199] Solenopsis daguerrei is a reproductive parasite to red imported fire ant colonies.^[200]

A large variety of pathogens and nematodes also infect red imported fire ants. Pathogens include Myrmecomyces annellisae, Mattesia, Steinernema,^[201] a mermithid nematode,^[202] Vairimorpha invictae, which can be transmitted via live larvae and pupae and dead adults^[203][204] and Tetradonema solenopsis, which can be fatal to a large portion of a colony.^[205] Individuals infected by Metarhizium anisopliae tend to perform trophallaxis more frequently and have an enhanced preference to quinine, an alkaloid substance.^[206] Phorid flies with Kneallhazia solenopsae can serve as vectors in transmitting the disease to the ants.^[207] Weakening the colony, infections from this disease are localised within the body fat, with spores only occurring in adult individuals.^[208][209] The mortality of an infected colony tends to be greater in contrast to those that are healthy.^[202] These ants are a host to Conidiobolus,^[210] Myrmicinosporidium durum, and Beauveria bassiana, each of which are parasitic fungi. Infected individuals have spores all over their bodies and appear darker than usual.^{[211][212][213]}

A virus, S. invicta 1 (SINV-1), has been found in about 20% of fire ant fields, where it appears to cause the slow death of infected colonies. It has proven to be self-sustaining and transmissible. Once introduced, it can eliminate a colony within three months. Researchers believe the virus has potential as a viable biopesticide to control fire ants.^{[214][215][216]} Two more viruses have also been discovered: S. invicta 2 (SINV-2) and S. invicta 3 (SINV-3). Polygynous colonies tend to face greater infections in contrast to monogynous colonies. Multiple virus infections can also occur.^[217][218]

Life cycle and reproduction

Nuptial flight in red imported fire ants begins during the warmer seasons of the year (spring and summer), usually two days after it has rained. The time alates emerge and mate is between 12:00 pm to 3:00 pm.^[219][220] Nuptial flights recorded in North Florida have, on average, 690 female and male alates participating in a single flight.^[221] Males are the first to leave the nest and both sexes readily undertake flight with little to no pre-flight activity. However, workers swarm the mound excitedly stimulated by mandibular glands within the head of the alates.^[222][223] As mounds do not have holes, workers form holes during nuptial flight as a way for the alates to emerge. This behaviour in workers, elicited by the pheromones, includes rapid running and back-and-forth movements, and increased aggression. Workers will also cluster themselves around the alates as they climb up on vegetation, and, in some cases, attempt to pull them back down before they take flight. Chemical cues from males and females during nuptial flight attracts workers, but chemical cues released by workers do not attract other nestmates. It also induces alarm-recruitment behaviour in workers which results in a higher rate of alate retrieval.^[222][224]

Based on aerial trapping, males fly much higher than females: captured males are usually 100 to 300 metres (328 to 984 ft) above the surface, whereas the females are only 60 to 120 metres (197 to 394 ft) above the surface. A nuptial flight takes place for roughly half an hour and females generally fly for less than 1.6 kilometres (0.994 mi) before landing. Studies show that 95% of queens successfully mate and only mate once; some males may be infertile due to the testicular lobes failing to develop.^{[219][225][226][227][228][229]} In polygyne colonies, males do not play a significant role and most are therefore sterile; one of the reasons for this is to avoid mating with other ant species. This also makes male mortality selective, which may affect the breeding system, mating success and, gene flow.^[230][231] Ideal conditions for a nuptial flight to begin is when humidity levels are above 80% and when the soil temperature is above 185 °C (64 °F). Nuptial flights only occur when the ambient temperature is 24–32 °C (75–89 °F).^[219]

Colony founding can be done by an individual or in groups, known as pleometrosis.^[232] This joint effort of the co-foundresses contributes to the growth and survival of the incipient colony; nests founded by multiple queens begin the growth period with three times as many workers when compared to colonies founded by a single queen. Despite this, such associations are not always stable.^{[233][234][235][236]} The emergence of the first workers instigates queen-queen and queen-worker fighting. In pleometrotic conditions, only one queen will emerge victorious, whereas the queens who lost are subsequently killed by the workers.^[237] The two factors that could affect the survival of individual queens are their relative fighting capabilities and their relative contribution to worker production. Experimentation indicates that size, an indicator of fighting capacity, positively correlates with survival rates. However, manipulation of the queen's relative contribution to worker production had no correlation with survival rate.^[238]

A single queen will lay approximately 10 to 15 eggs 24 hours after mating.^[94] In established nests, a queen will apply venom onto each egg that perhaps contains a signal calling for workers to move it.^[239] These eggs remain unchanged in size for one week until they hatch into larvae. By this time, the queen will have laid approximately 75 to 125 more eggs. The larvae that hatch from their eggs are usually covered in their shell membranes for several days. The larvae can free their mouthparts from their shells using body movements, but still need assistance from workers with hatching. The larval stage is divided into four instars, as observed through the moulting stages. At the end of each moult, a piece of unknown material is seen connected to the exuviae if they are isolated from the workers. The larval stage lasts between six and 12 days before their bodies expand significantly and become pupae; the pupal stage lasts between nine and 16 days.^[47][94]

As soon as the first individuals reach the pupal stage, the queen will cease egg production until the first workers mature. This process takes approximately two weeks to one month. The young larvae are fed oils which are regurgitated from her crop, as well as trophic eggs or secretions. She will also feed the young her wing muscles providing the young with needed nutrients. The first generation of workers will always be small because of the limited nutrients needed for development. These workers are known as minims nanitics who burrow out of the queen's chamber and commence foraging for food needed for the colony. Mound construction also occurs at this time. Within a month after the first generation is born, larger workers (major workers) will start to develop, and within six months, the mound will be noticeable, if viewed, and will house several thousand residents. A mature queen is capable of laying 1,500 eggs per day; all workers are sterile and cannot reproduce.^{[94][240][241][242]}

The life expectancy of a worker ant depends on its size, although the overall average is around 62 days.^[243] Minor workers are expected to live for about 30 to 60 days, whereas the larger workers will live much longer. Larger workers, which have a life expectancy of 60 to 180 days, will live 50–140% longer than their smaller counterparts, but this depends on the temperature.^[94][244] However, workers kept in laboratory conditions have been known to live for 10 to 70 weeks (70 days to 490 days); the maximum recorded longevity of a worker is 97 weeks (or 679 days).^[245] The queens live much longer than the workers, with a life span ranging from two years to nearly seven years.^[94][245]

There are several factors that contribute to colony growth. Temperature plays a major role in colony growth and development; colony growth ceases below 24 °C and developmental time decreases from 55 days at temperatures of 24 °C to 23 days at 35 °C. Growth in established colonies only occurs at temperatures between 24 °C to 36 °C. Nanitic brood also develops far quicker than minor worker brood (approximately 35% faster), which is beneficial for founding colonies.^[246] Colonies that have access to an unlimited amount of insect prey are known to grow substantially, but this growth is further accelerated if they are able to access plant resources colonised by hemipteran insects.^[247] In incipient monogyne colonies where diploid males are produced, colony mortality rates are significantly high and colony growth is slow. In some cases, monogyne colonies experience 100% mortality rates in the early stages of development.^[248]

Based on several studies, the population of a colony can grow exceptionally fast. One observation shows that colonies that housed 15–20 workers in May grew to over 7,000 by September. These colonies started to produce reproductive ants when they were a year old, and by the time they were two years old they had over 25,000 workers. The population doubled to 50,000 when these colonies were three years old.^[249] At maturity, a colony can house up to 100,000 to 250,000 individuals, but other reports suggest that colonies can hold more than 400,000.^{[lower-alpha 4][250][251][252][253]} Polygyne colonies have the potential to grow much larger than monogyne colonies.^[251]

In colonies, queens are the only ants able to alter sex ratios which can be predicted. For example, queens originating from male-producing colonies tend to produce predominantly males; while a queen that came from a female-favoured sex ratio colony will tend to produce females.^[254][255] Queens also exert control over the production of sexuals through pheromones that influence the behaviours of workers toward both male and female larvae.^[256]

Monogyny and polygyny

There are two forms of society in the red imported fire ant: polygynous colonies and monogynous colonies.^{[257][258][257]} Polygynous colonies differ substantially from monogynous colonies in social insects. The former experience reductions in queen fecundity, dispersal, longevity, and nestmate relatedness.^{[259][260][261][262]} Polygynous queens are also less physogastric than monogynous queens and workers are smaller.^[263][264] Understanding the mechanisms behind queen recruitment is integral to understanding how these differences in fitness are formed. It is unusual that the number of older queens in a colony does not influence new queen recruitment. Levels of queen pheromone, which appears to be related to queen number, play important roles in the regulation of reproduction. It would follow that workers would reject new queens when exposed to large quantities of this queen pheromone. Moreover, evidence supports the claim that queens in both populations enter nests at random, without any regard for the number of older queens present.^[265] There is no correlation between the number of older queens and the number of newly recruited queens. Three hypotheses have been posited to explain the acceptance of multiple queens into established colonies: mutualism, kin selection, and parasitism.^[266] The mutualism hypothesis states that cooperation leads to an increase in the personal fitness of older queens. However, this hypothesis is not consistent with the fact that increasing queen number decreases both queen production and queen longevity.^[267] Kin selection also seems unlikely given that queens have been observed to cooperate under circumstances where they are statistically unrelated.^[268] Therefore, queens experience no gain in personal fitness by allowing new queens into the colony. Parasitism of preexisting nests appears to be the best explanation of polygyny. One theory is that so many queens attempt to enter the colony that the workers get confused and inadvertently allow several queens to join it.^[268]

Monogyne workers kill foreign queens and aggressively defend their territory. However, not all behaviours are universal, primarily because worker behaviours depend on the ecological context in which they develop, and the manipulation of worker genotypes can elicit change in behaviours. Therefore, behaviours of native populations can differ from those of introduced populations.^[269] In a study to assess the aggressive behaviour of monogyne and polygyne red fire ant workers by studying interaction in neutral arenas, and to develop a reliable ethogram for readily distinguishing between monogyne and polygyne colonies of red imported fire ants in the field,^[270] monogyne and polygyne workers discriminated between nestmates and foreigners as indicated by different behaviours ranging from tolerance to aggression. Monogyne ants always attacked foreign ants independently if they were from monogyne or polygyne colonies, whereas polygyne ants recognised, but did not attack, foreign polygyne ants, mainly by exhibiting postures similar to behaviours assumed after attacks by Pseudacteon phorids. Hostile versus warning behaviours were strongly dependent on the social structure of workers. Therefore, the behaviour toward foreign workers was a method of characterising monogyne and polygyne colonies.^[271] Most colonies in the southeastern and south-central US tend to be monogynous.^[236]

The monogynous red imported fire ant colony territorial area and the mound size are positively correlated, which, in turn, is regulated by the colony size (number and biomass of workers), distance from neighbouring colonies, prey density, and by the colony's collective competitive ability. In contrast, nestmate discrimination among polygynous colonies is more relaxed as workers tolerate conspecific ants alien to the colony, accept other heterozygote queens, and do not aggressively protect their territory from polygyne conspecifics.^[272] These colonies might increase their reproductive output as a result of having many queens and the possibility of exploiting greater territories by means of cooperative recruitment and interconnected mounds.^[273]

Relationship with other organisms

When polygyne forms invade areas where colonies have not yet been established, the diversity of native arthropods and vertebrates declines greatly.^[274] This is evident as populations of isopods, mites and tumblebug scarabs decline significantly. They can also significantly alter the populations of many fly and beetle families, including: Calliphoridae, Histeridae, Muscidae, Sarcophagidae, Silphidae, and Staphylinidae. Despite this, one review found that certain insects may be unaffected by red imported fire ants; for example, the density of isopods decreases in red imported fire ant infested areas, but crickets of the genus Gryllus are unaffected. There are some cases where the diversity of certain insect and arthropod species increase in areas where red imported fire ants are present.^{[275][276][277]} Red imported fire ants are important predators on cave invertebrates, some of which are endangered species. This includes harvestmen, pseudoscorpions, spiders, ground beetles, and pselaphid beetles. The biggest concern is not the ant itself, but the bait used to treat them because this can prove fatal. Stock Island tree snails (Orthalicus reses) are extinct in the wild; predation by red imported fire ants is believed to be the major factor in the snail's extinction.^[275] Overall, red imported fire ants prefer specific arthropods to others, although they attack and kill any invertebrate that cannot defend itself or escape.^[278] Arthropod biodiversity increases once red imported fire ant populations are either reduced or eradicated.^[115][279]

Interactions between red imported fire ants and mammals have been rarely documented. However, deaths of live-trapped animals by red imported fire ants have been observed.^{[275][280][281]} Mortality rates in eastern cottontail (Sylvilagus floridanus) young range from 33 to 75% because of red imported fire ants.^[282] It is believed that red imported fire ants have a strong impact on many herpetofauna species; scientists have noted population declines in the Florida kingsnake (Lampropeltis getula floridana), and eggs and adults of the eastern fence lizard (Sceloporus undulatus) and six-lined racerunner (Aspidoscelis sexlineata) are a source of food.^{[283][284][285]} Because of this, eastern fence lizards have adapted to have longer legs and new behaviours to escape the red imported fire ant.^[286][287] Adult three-toed box turtles (Terrapene carolina triunguis), Houston toad (Anaxyrus houstonensis) juveniles, and American alligator (Alligator mississippiensis) hatchlings are also attacked and killed by these ants.^{[275][288][289][290]} Despite this mostly-negative association, one study shows that red imported fire ants may be capable of impacting vector-borne disease transmissions by regulating tick populations and altering vector and host dynamics, thereby reducing transmission rates not only to animals, but to humans as well.^[291]

Mortality rates have been well observed in birds; there have been instances where no young have survived to adulthood in areas with high fire ant density. Many birds including cliff nesting swallows, ducks, egrets, quail, and terns have been affected by red imported fire ants.^[292] Ground nesting birds, particularly the least tern (Sterna antillarum), are vulnerable to fire ant attacks.^[293][290] The impact of red imported fire ants on colonial breeding birds is especially severe; waterbirds can experience a mortality rate of 100%, although this factor was lower for early-nesting birds. Brood survival decreases in American cliff swallows (Petrochelidon pyrrhonota) if they are exposed to foraging workers.^[294] Songbird nest survival decreases in areas with red imported fire ants present, but survival rates in white-eyed vireo (Vireo griseus) and black-capped vireo (Vireo atricapilla) nests increase from 10% to 31% and 7% to 13% whenever fire ants are not present or when they are unable to attack them.^[295] Red imported fire ants may indirectly contribute to low brood survival in the Attwater's prairie chicken.^[296] It was first thought that the ants were linked to the decline of overwintering birds such as the loggerhead shrike (Lanius ludovicianus), but a later study showed that ant eradication efforts using the pesticide Mirex, which was known to have toxic side effects, was largely to blame.^[297][298]

Red imported fire ants are strong competitors with many ant species. They have managed to displace many native ants which has led to a number of ecological consequences.^[299] However, studies show that these ants are not always superior competitors that suppress native ants. Habitat disturbance prior to their arrival, and recruitment limitations, are more plausible reasons why native ants are suppressed.^[300] Between Tapinoma melanocephalum and Pheidole fervida, the red imported fire ant is stronger than both species but shows different levels of aggression. For example, they are less hostile towards T. melanocephalum in contrast to P. fervida. Mortality rates in T. melanocephalum and P. fervida when fighting with red imported fire ants are high, being 31.8% and 49.9% respectively. The mortality rate for red imported fire ant workers, however, is only 0.2% to 12%.^[301] The imported crazy ant (Nylanderia fulva) exhibits greater dominance than the red imported fire ant and has been known to displace them in habitats where they encounter each other in.^[302] Larger colonies of pavement ants (Tetramorium caespitum) can destroy red imported fire ant colonies, leading entomologists to conclude that this conflict between the two species may help impede the spread of the red imported fire ant.^[303][304] Individuals infected by SINV-1 can be killed faster than healthy individuals by Monomorium chinense. This means that ants infected with SINV-1 are weaker than their healthy counterparts and more than likely will be eliminated by M. chinense. However, major workers, whether they are infected or not, are rarely killed.^[305]

In areas it is native to, the red important fire ant is still a dominant species and coexists with 28 ant species in gallery forest gaps and ten species in xerophytic forest grassland, winning most aggressive interactions with other ants. However, some ants can be co-dominant in areas where they coexist such as the Argentine ant, where they compete symmetrically.^[306][307] Workers regularly engage in food competition with other ants, and can suppress the exploitation of food resources from honeydew-producing hemipterans (specifically from Phenacoccus solenopsis) from native ants; however, red important fire ants are unable to eliminate T. melanocephalum completely although they consume a higher proportion of food. Instead, the two ants may peacefully coexist and share the honeydew.^[308][309] When encountering neighboring fire ants, workers may take on death-feigning behaviours to avoid them with success. However, such behaviour is only seen in young workers, as older workers either flee or fight back when threatened.^[310]

As mentioned, red imported fire ants and Argentine ants compete with each other. Mortality rates vary in different scenarios (i.e. mortality rates in colony confrontation is lower than those confronting each other in the field). Major workers can also withstand more injuries to their bodies, thus increasing the mortality rate of Argentine ants. For an Argentine ant colony to successfully wipe out a monogyne colony of 160,000 workers, the colony would need 396,800 workers. A colony that has reduced in size due to successful bait treatment are prone to predation by Argentine ants. The ants may play a vital role in removing weakened fire ant colonies, and they may also be important in slowing the spread of these fire ants, especially in heavily infested Argentine ant areas.^[311] Despite this, Argentine ant populations in the southeastern United States have declined following the introduction of the red imported fire ant.^[312]

Red imported fire ants have formed a relationship with an invasive mealybug, Phenacoccus solenopsis. Colony growth is known to increase if ants have access to resources produced by P. solenopsis, and the population density of these mealybugs is significantly higher in areas where red imported fire ants are present (the spread of P. solenopsis in areas it is not native is attributed to the presence of the ants). As well as that, the life expectancy and reproductive rate of P. solenopsis both increase. These mealybugs may even be transported by workers back to their nest.^[313] Predation of P. solenopsis decreases due to tending workers interfering with the predation and parasitism by natural enemies. Mealybugs are preyed on by the lady beetle (Menochilus sexmaculatus) and are hosts of two parasites (Aenasius bambawalei and Acerophagus coccois) if there are no ants present, but this is different if ants are present. Lady beetles are less frequently seen on plants with red imported fire ants, and the rate of "mummy" nymphs is significantly lower.^[313] This is clearly evident as scientists observed that cotton aphid (Aphis gossypii) populations, and the predation of sentinel bollworm eggs, increased in areas with red imported fire ant presence.^{[314][315][316]} Red imported fire ants have developed a mutual relationship with another mealybug (Dysmicoccus morrisoni). The ants promote the colony growth of D. morrisoni through protection, covering the colonies up with debris and collecting the honeydew they secrete.^[317]

Toxicology

Venom

Venom plays an important role in a red imported fire ant colony, being used to capture prey items, for defence, and antimicrobial action.^[318][319] On average, however, a worker produces very little venom (less than 1 µg/kg).^[320] Newborn workers contain little to no venom within their reservoirs, but workers that are only one-day old can produce 1.17 µg/kg. However, workers that are 15 days old only produce 0.3 µg/kg. Workers deliver 0.66 nl of venom when they sting, which amounts to 3.1% of their supply. Older workers deliver less venom when they sting, but middle-aged workers and nest-defenders deliver much higher quantities.^[318] Like all fire ant species, venom is secreted by the venom gland and is stored in the poison sac. When in use, it is ejected through the main duct. Capacity is between 20 and 40 nl, but this depends on the worker's size.^[321] The Schmidt sting pain index, a pain scale which ranks the pain intensity of an insect's sting from 0 to 4, ranks the pain at "1". American entomologist Justin O. Schmidt described it as being "sharp, sudden, mildly alarming. Like walking across a shag carpet & reaching for the light switch."^[322]

95% of the venom components are water-insoluble piperidine alkaloids. Piperidines include trans-2-methyl-6-n-undecylpiperidines, trans-2-Methyl-6-n-tridecylpiperidine, trans-2-Methyl-6-(cis-4-tridecenyl) piperidines, trans-2-methyl-6-n-pentadecylpiperidine, trans-2-methyl-6-(cis-6-pentadecenyl)piperidine and 2,6-dialkylpiperidines (the ants' venom is dominated by the trans stereoisomers of this specific ingredient).^{[23][323][324]} Trans-2-methyl-6-n-undecylpiperidine (solenopsin) has been shown to have cytotoxic, hemolytic, necrotic, insecticidal, antibacterial, antifungal, and anti-HIV properties.^[325] As well as that, the alkaloid has shown antiangiogenic activity.^[326] These components are responsible for the formation of hives, and also for the development of sterile pustules on areas where the ant has stung.^[327] Experiments indicate that the median lethal dose (LD₅₀) on tested female rats is 0.36 mg/kg.^[328]

Approximately 46 proteins have been identified in the red imported fire ant's venom, although scientists long believed the venom only contained alkaloids. This is most likely attributed to the low protein content found in the venom, only representing 0.1% of the venom's total weight. These proteins most likely account for the anaphylactic reactions seen in humans sensitive to the venom.^[329][330] However, not all of these proteins are involved with venom function. The venom only has four protein allergens, which have been extensively investigated: this includes Sol i 2 and 4, which are not similar to other characterised proteins and have not been described in other insect venoms. Another protein, Sol i 3, is part of the antigen 5 family, and Sol i 1 is a phospholipase A1B; Sol i 1 shows a close relation with wasp venom phospholipases.^{[327][331][332][333]} Other proteins found in the venom may benefit the colony; some of these proteins can kill off bacteria, which may explain why workers spray venom around their nests by vibrating their gasters. Other proteins also bind pheromones which may assist a worker to lay chemical trails to communicate with other nestmates.^{[320][330][334]}

Incidence

In the United States, more than 40 million people live in areas infested with fire ant populations and 14 million people are stung by them annually. A quarter of all victims stung by red imported fire ants are expected to develop sensitivity to the venom, and approximately 6,000 will suffer anaphylaxis.^[335] 51% of people who relocated themselves to infested areas report getting stung within three weeks after arrival.^[336] In a survey conducted in South Carolina, 33,000 people (or 94 per 10,000 population) received medical attention due to red imported fire ants, and 660 people (1.9 per 10 000 population) were treated for anaphylaxis.^[337] In Texas, 79% of participants in a survey stated they had been stung by red imported fire ants, while 20% had not. West Texans had the lowest incidence rate; approximately 61% of west Texans had been stung, compared to 90% in central Texas, 89% in east Texas, 86% in the gulf coastal regions, 78% in the south and 72% in the north.^[88] In another conducted survey in Georgia, 87% of individuals classed their reactions as mild, 12% as moderate and 1% as severe.^[338] In Australia, 64,000 homes are within red imported fire ant infested areas, and 140,000 consultations and 3,000 anaphylactic reaction cases are predicted every year by 2030 if government efforts to eradicate the ant fail.^[339] A survey conducted in China shows that one-third of participants in infested areas were victims of red imported fire ant stings.^[340]

Studies suggest that the rate of systemic reactions to stings may be associated with seasonal variations in the venom's potency. 51% of allergic reactions occurred in summer, and 19% in spring. However, A survey reported a higher incidence during spring (39.9%) than summer (31.9%).^[341] Younger people, usually those under 20 years, experience the highest rate of sting attacks (50%), but the rate declines with older people. Among men and women, the rate varies as some studies report more women being attacked than men and vice versa.^[338][340] Deaths from red imported fire ant stings are rare, but may become common the more the ant spreads. Many cases have also been reported in the past.^[342] It is reported that more than 80 deaths have been recorded; of these, 22 cases were recorded in Florida and 19 in Texas. However, when duplicate reports are excluded, four deaths were recorded in Alabama, 10 in Florida, two in Georgia and Louisiana, and 14 in Texas.^{[343][344][345]} People can be educated and be made aware of the dangers of red imported fire ants.^[346]

Signs and symptoms

Reactions seen in humans vary; some are hypersensitive to venom while others show resilience. Hypersensitivity can be attributed to certain medical problems such as heart conditions or diabetes. Bacterial infections attributed to sting injuries also pose a problem and may require further medical attention. Most humans can withstand many stings, but others may suffer from severe reactions such as anaphylaxis.^[10]

People who are stung by red imported fire ants experience intense burning or flare-ups, followed by reddening of the skin at the sting site. This area will swell into a bump, hive or vesicle within 20 minutes. White fluid-filled sterile pustules begin to form within hours or days after being stung.^{[10][340][347][348]} Pustules on the skin remain for a couple of days, and may become infected which would require medical attention. In most cases, pustules dry up in a matter of weeks and leave brown scars that either remain for several months or become permanent.^[10] The formation of pustules occurs in almost every person stung by the ants. In one study, 96% of participants reported the formation of pustules, whereas 2% reported large local reactions.^[336] Between 17% to 56% of people stung develop venom-specific IgE. Many of them will experience pruritic lumps around areas where the ants stung, known as late-phase responses or cutaneous allergic reactions.^{[344][349][350]}

Pustule formation can only be prevented if the ants are removed before they have a chance to sting. Once venom has been injected, pustules will form and no form of treatment will prevent them from occurring. Medications such as antibiotics, diphenylhydrazines, epinephrines or topical steroids will not affect pustular reactions.^{[10][350][351]}

Anaphylaxis occurs in 0.6 to 6% of people who have been stung by the ants, and it can be fatal if left untreated.^[342][344] Typical symptoms of anaphylaxis include dizziness, headaches, fever, severe chest pain, nausea, severe sweating, low blood pressure, loss of breath, serious swelling, and slurred speech.^{[10][340][352]} One case reports a victim feeling strong vertigo 5 to 10 minutes after being stung, followed by glassy eyes, dry mouth, paleness, unconsciousness and severe cramps on the sting sites.^[353] In addition, neuropathy, seizures (even without any evidence of prior systemic reactions), cerebrovascular accidents, and nephrotic syndrome have been associated with red imported fire ant stings.^{[10][344][354]} A series of neurotoxins have been identified in red imported fire ant venom, which may explain why some victims experience hallucinations after they have been stung.^[320]

Treatment

It is suggested that a conservative approach be used when treating sting injuries; specifically, the kind of treatment used should be based on the symptoms. For minor sting injuries, with symptoms only including pustule formations and pain, over-the-counter products are available to prevent infection. Ants should be removed by washing the area with antiseptic soap, and itchiness can be relieved through the use of nonsedative antihistamines or ice packs. This should be done with care because sterile pustules can rupture and become infected. The use of intravenous fluids or parenteral corticosteroids to treat fire ant stings is not recommended unless the person who was stung shows evidence of hypersensitivity. The use of these elements can lead to a number of health problems, including fluid retention and cardiovascular compromise. However, such conditions are often restricted to patients with preexisting cardiovascular insufficiencies. It is rare for ant sting sites to become infected, so the use of antibiotic prophylaxis is not always required.^[10][355]

Victims who show signs of anaphylaxis are treated with antihistamines, epinephrines, and parenteral corticosteroids.^[344] Epinephrine is the first product for use to treat systemic allergic responses, particularly if a patient is experiencing dyspnoea or hypotension because it is capable of reversing adverse events quickly and is very safe to use. It is recommended that people who have suffered from anaphylaxis carry an epinephrine autoinjector (EpiPen), should dyspnoea or hypotension begin to occur.^[339]

Whole body extract immunotherapy (WBE) to treat victims of anaphylaxis has been in use since 1973.^[356][357] Anyone who has a suspected allergy to the venom is redirected to an allergist for assessment.^[10] The treatment uses the entire body of the ant and not just the venom, and unlike fire ant venom immunotherapy (which is occasionally used), WBE contains venom proteins.^[358] To reduce a patient's sensitivity to the venom, gradual increases of dose extracts are injected into the body.^[359] WBE immunotherapy appears to be very effective in preventing systemic reactions; in one study of participants who completed WBE immunotherapy, two out of fifteen participants suffered from allergic reactions upon being stung 18 months after immunotherapy, while zero out of nine participants exhibited no reactions 54 months after immunotherapy.^{[355][360][361]} As mentioned, fire ant venom immunotherapy is occasionally used, and studies show it can reduce the risk of systemic reactions.^[362][363] In fact, another study claims that fire ant venom immunotherapy is more effective than WBE immunotherapy.^[364] Fire ant venom immunotherapy is not recommended for children with large local reactions, although an exception may be made for those who live in heavily infested areas. There is also an increased risk of systemic allergic reactions to future stings in children who have cutaneous manifestations after getting stung. This makes many experts put some children on fire ant venom immunotherapy, while others do not.^[355][365]

The recommended maintenance dose is between 0.5 mL of a 1:100 w/v 1:10 w/v WBE.^[366] For fire ant venom immunotherapy, the most common maintenance dose is 0.5 mL of a 1:200 (wt/vol) dilution.^[367] During the build-up phase, it is recommended that dosing is given weekly or biweekly, although some scientists suggest that rush protocols can be successful.^[355][368] It is recommended that patients going through immunotherapy receive treatment for three to five years, and lifelong therapy, although there is no consensus as to how long an individual should be treated.^[355][369]

Stings to animals

The stings of the red imported fire ant in animals are painful and sometimes life-threatening.^[370] In dogs, stings from the red imported fire ant can cause pustular dermatosis, a condition where pustules appear in crops as a result of the ant sting.^[371] After getting stung, the immediate response consists of erythema and swelling. The pustules remain for approximately 24 hours, whereas in humans they can last for several days.^[372] In livestock, red imported ants mostly sting animals in regions with no hair, particularly around the ears, eyes, muzzle, the perineum and ventral portion of the abdomen. Newborn or young livestock can be blinded or killed when attacked by the ants.^[373][374] Healthy individuals are less likely to be attacked than weak or sick animals. Red papule and mild swelling occur, followed by vesicopustule with a red halo developing within 24 to 48 hours. The eyes and eyelids are commonly damaged from the stings; in sheep and goats, ophthalmic ointment containing antibiotics and corticosteroids can be used to treat the eyes of sheep and goats, but this treatment is not recommended for horses. In non-domestic animals, cases of red imported fire ants stings in animals such as ferrets, moles squirrels, white-tailed deer, cottontail rabbits, and newborn blackbucks have been reported, as well as lizards and screech owl nestlings. The aftermath of the injuries is like those in domestic animals.^[373] In tested conditions, rats can experience seizures, respiratory arrest, or even death from the ants' venom.^[325]

Red imported fire ants do not intend to kill animals because they are too large and their venom cannot fatally injure them. Animals may trigger stinging episodes if they irritate or injure the ants on their body, with all individuals participating in the attack. These large-scale attacks on an animal are caused by chemical signals.^[373] As a result, this explains why an animal may suffer from hundreds of individual stings. It is suspected that many victims of the red imported fire ants may be depressed as a result of the effects of the toxin. Some animals may swallow red imported fire ants as they lick or bite around the sites they are stinging. This can cause additional injuries inside the animal itself, especially in the upper gastrointestinal tract. In suckling white tail deer fawns, sting sites have been found in the oesophagus and abomasum; toxins from the ingested ants may cause inflammation of the gastrointestinal lining.^[373]

Relationship with humans

As pests

Owing to its notoriety and invasive behaviour, red imported fire ants are considered to be pests. In the United States, the Food and Drug Administration estimates more than $5 billion is spent annually on medical treatment, damage, and control in infested areas.^[375][376] They can also cause $2.5 billion in damages annually in just 20 years.^[377] In Texas alone, red imported fire ants caused $300 million in damages for livestock, wildlife and public health.^[378] Approximately $36 per American household, and more than $250 million has been spent to control and eradicate red imported fire ants. Private agencies spend $25 to $40 million on pesticides annually.^[94] Unsuccessful eradication in Australia may cost the economy billions in damages annually and, based on a Queensland government study, the estimated cost could reach $43 billion over 30 years.^[379]

Red imported fire ants thrive in urban areas, especially in backyards, golf courses, parks, recreational areas, school grounds, and street verges, in which their presence may deter outdoor activities. If they enter homes or properties, they can harm pets if they are either caged, penned, tied or cannot escape.^[376][380] Nests can be built under pavements or even roads, as well as under driveways, foundations, lawns, edges of sidewalks, under patio slabs, in electrical boxes or near power lines. A colony can excavate huge quantities of soil, resulting in structural problems in driveways, pavings and walls, and can also cause the formation of potholes in roads.^{[94][376][381][382][383]} Additional damage by mounds can be inflicted on trees, yard plants, and pipes; some structures may also collapse. Colonies may migrate into human homes after heavy rain to take refuge from saturated soil.^[50][94]

Not only do they thrive in urban areas, red imported fire ants can damage equipment and infrastructure and impact business, land, and property values.^[380] They are also attracted to electricity; electrically stimulated workers release venom alkaloids, alarm pheromones, and recruitment pheromones, which in return attracts more workers to the site.^[384] As a result, red imported fire ants can destroy electrical equipment.^[385] This is known as magnetism, where scientists have identified internal magnetic materials which may play a role in orientation behaviours.^[386][387] They are known to chew through electrical insulation which causes damage to electric motors, irrigation lines, pumps, signal boxes, transformers, telephone exchanges, and other equipment.^[376][388] Colonies aggregate near electrical fields and are capable of causing short circuits or interfering with switches and equipment such as air conditioners, computers, and water pumps. They are also known to infest airport landing areas and traffic lights.^[388]

In agriculture

The red imported fire ant is a significant agricultural pest in areas where it is not native. They are capable of damaging crops, and they threaten pastures and orchards.^[380] The mounds themselves can destroy agricultural equipment such as irrigation systems and damage machinery during harvest time.^[389] The ants are known to invade soybean crops causing lower yields, and could cause $156 million in losses for soybean crops in the southeastern United States.^{[94][390][391]} Reports from Georgia and North Carolina claim that 16.8 to 49.1 kg/ha of soybeans could not be harvested due to interference from fire ant mounds; combine harvesters skipped over the mounds preventing the crops from being harvested, and farmers raised the cutter bars on their combine harvesters to avoid impacting the mounds.^{[392][393][394]} The feeding behaviour of red imported fire ants can result in significant damage to many other crops, including: beans, cabbage, citrus, corn, cucumber, eggplant, okra, peanut, potato, sorghum, sunflower, and sweet potato. The ants also interfere with plants' root systems and feed on young growth.^[94][395] Sometimes, colonies build mounds around or near the base of citrus trees chew off new growth and feed on the flowers or developing fruit. Citrus trees are often girdled or killed.^[396]

Despite its pest status and notoriety, the red imported fire ant can be beneficial. The ant is an effective insect predator, so it may serve as a biological agent against other pest species, especially in sugarcane fields.^[397] Pest insects the ant kills include: boll weevils (Anthonomus grandis) in cotton crops,^{[398][399][400]} sugarcane borers (Diatraea saccharalis) in sugarcane fields,^{[401][402][403]} horn flies (Haematobia irritans) in manure,^[404] velvetbean caterpillars (Anticarsia gemmatalis) in soybeans, and whiteflies that are found in greenhouses.^[405][406] A number of studies indicate that red imported fire ants do not interfere or attack insects of economic importance in cotton fields, which has resulted in many farmers in the southeastern regions of the United States viewing red imported fire ants as beneficial.^[407] However, some scientists have suggested that the beneficial status of the red imported fire ant is hard to predict when geography, plant size, season, soil moisture and the use of insecticides are not considered. These factors may lower the efficacy of red imported fire ants as pest control agents. Another factor is that workers are indiscriminate and kill beneficial insects such as predators that eat horn flies and other pests in pastures and predators of aphids and scale insects.^{[407][407][408][409]} They also reduce the efficacy of parasitic wasps against pest species by eating the larvae and pupae.^[407][410]

Control

In comparison to other ant species like Anoplolepis gracilipes, which rapidly take over areas they have been in contained in, red imported fire ants are fairly easy to control.^[411] The first proposals to controlling the ant occurred in 1957 when the United States Congress authorised an eradication program using federal and state funding. Research on the ant and its biology has been continuous after the establishment of an eradication program, and many chemicals were used to eliminate them. However, scientists discovered that these insecticides were killing native fauna, and the Environmental Protection Agency subsequently outlawed them. Some scientists even questioned whether the ants were pests or not.^[412][413] Today, it is unlikely that the red imported fire ant will be eradicated in areas such as the United States. Populations can be managed properly if an integrated approach is used.^[414] Some scientists have considered using the ants' natural enemies against it; this includes T. solenopsae and B. bassiana. Phorid flies have also been viewed as potential biological agents, as they can reduce foraging activity in red imported fire ants and affect population levels. However, they are unable to affect colony growth rate.^[415] In addition, parasitic ants, parasitic wasps, mites, other pathogens, nematodes, and fungi have been considered to be potential biological agents.^[416] Others suggest that populations can be maintained or reduced by manipulating several ecological factors.^[417]

Several baits have been used to control populations. Mounds are destroyed in a matter of weeks if baits are used on them. Baits are considered to be effective and simple to use against red imported fire ants, in comparison to drenching, dusting, or fumigating. They are sprinkled onto the mound then the ants take and consume them.^[346][418] Certain baits such as growth regulator baits, and boric acid-sucrose water baits, benefit native fauna and low concentrations are usually required to kill a colony.^[419][420] Others baits used against red imported fire ants include Amdro, Ascend, hydramethylnon, and Maxforce.^[421]

Researchers have also been experimenting with other methods, such as fumigation, injecting the mound, mound drenching, and surface dusting. Mound drenching entails pouring large volumes of toxic liquid into the mound. While this may affect a large portion of the mound, it is possible it may not reach the queen, thus preventing the destruction of the colony. Surface dusting is similar to mound drenching, except insecticides are applied on top of the mound and soak into the soil when wet. Mound injections use pressurising and injecting insecticides into the mound, but the queen may not be affected by this. Mounds can be eliminated effectively through fumigation. Broadcast treatment is sometimes used by spraying infected areas with agricultural equipment. Finally, spot treatment with insecticides can be used by drilling and injecting the mound with residual insecticide.^[94][346]

Aside from chemical control, other methods that can be employed against these ants include mechanical and electrical devices. However, it is unknown whether or not these devices are effective. Ant-proofing can be effective against colonies nesting inside buildings by caulking and sealing cracks, which successfully suppresses the population outside the walls. Homeowners have used their own methods to remove mounds by pouring boiling water on them or igniting them with flammable liquids. Although these methods may be effective, they are not recommended because they can be harmful to humans and the environment.^[94][346]

See also

Notes

References

Footnotes

Bibliography

External links

Library resources about Red imported fire ant

Resources in your library Resources in other libraries

Websites

• Books or journals about the red imported fire ant at Biodiversity Heritage Library
• Books or journals about the red imported fire ant at Internet Archive
• National Red Imported Fire Ant Eradication Program – Queensland Department of Agriculture and Fisheries
• Imported Fire Ants – United States Department of Agriculture
• Solenopsis invicta at the AntWiki – Bringing Ants to the World
• The S. invicta Genome Project – Hymenoptera Genome Database project

Media

• Footage of phorid flies parasitising fire ants – Captured by BBC, published by the University of Texas at Austin College of Natural Sciences
• Red Fire Ants – published by National Geographic at YouTube
• Casting a Fire Ant Colony with Molten Aluminum at YouTube

News coverage

• The Incredible Floating Fire Ant – The Washington Post, 2011