Myrmecia is a genus of first established by Danish zoologist Johan Christian Fabricius in 1804. The genus is a member of the subfamily Myrmeciinae of the family Formicidae. Myrmecia is a large genus of ants, comprising at least 93 species that are found throughout Australia and its coastal islands, while a single species is only known from New Caledonia. One species has been introduced out of its natural distribution and was found in New Zealand in 1940, but the ant was last seen in 1981. These ants are commonly known as bull ants, bulldog ants or jack jumper ants, and are also associated with many other common names. They are characterized by their extreme aggressiveness, ferocity, and painful stings. Some species are known for the jumping behavior they exhibit when agitated.
Species of this genus are also characterized by their elongated mandibles and large Arthropod eye that provide excellent vision. They vary in colour and size, ranging from . While workers and queens are hard to distinguish from each other due to their similar appearance, males are identifiable by their perceptibly smaller mandibles. Almost all Myrmecia species are monomorphic, with little variation among workers of a given species. Some queens are ergatoid and have no wings, while others have either stubby or completely developed wings. Nests are mostly found in soil, but they can be found in rotten wood and under rocks. One species does not nest in the ground at all; its colonies can only be found in trees.
A queen will mate with one or more males, and during colony foundation she will hunt for food until the brood have fully developed. The life cycle of the ant from egg to adult takes several months. Myrmecia workers exhibit greater longevity in comparison to other ants, and workers are also able to reproduce with male ants. Myrmecia is one of the most primitive group of ants on earth, exhibiting differentiated behaviors from other ants. Workers are solitary hunters and do not lead other workers to food. Adults are omnivores that feed on sweet substances, but the larvae are carnivores that feed on captured prey. Very few predators eat these ants due to their sting, but their larvae are often consumed by and , and a number of parasites infect both adults and brood. Some species are also effective pollinators.
Myrmecia stings are very potent, and the venom from these ants is among the most toxic in the insect world. In Tasmania, 3% of the human population are allergic to the venom of M. pilosula and can suffer life-threatening anaphylaxis if stung. People prone to severe allergic reactions can be treated with allergen immunotherapy (desensitisation).
Ants of this genus are popularly known as bulldog ants, bull ants, or jack jumper ants due to their ferocity and the way they hang off their victims using their mandibles, and also due to the jumping behaviour displayed by some species. Other common names include "inch ants", "sergeant ants", and "soldier ants".
In 1911, Emery classified the Subgenus Myrmecia, Pristomyrmecia, and Promyrmecia, based on the shape of their mandibles. Wheeler established the subgenus Halmamyrmecia, and the ants placed in it were characterized by their jumping behavior. The taxon Wheeler described was not referred to in his later publications, and the genera Halmamyrmecia and Pristomyrmecia were synonymised by John Clark. At the same time, Clark reclassified the subgenus Promyrmecia as a full genus. He revised the whole subfamily Myrmeciinae in 1951, recognizing 118 species and subspecies in Myrmecia and Promyrmecia; five species groups were assigned to Myrmecia and eight species groups to Promyrmecia. This revision was rejected by entomologist William Brown due to the lack of morphological evidence that would make the two genera distinct from each other. Due to this, Brown classified Promyrmecia as a synonym of Myrmecia in 1953. Clark's revision was the last major taxonomic study on the genus before 1991, and only a single species was described in the intervening years. In 2015, four new Myrmecia ants were described by Robert Taylor, all exclusive to Australia. Currently, 94 species are described in the genus, but as many as 130 species may exist.
Under the present classification, Myrmecia is the only extant genus in the tribe Myrmeciini, subfamily Myrmeciinae. It is a member of the family Formicidae in the order Hymenoptera. The type species for the genus is Myrmecia gulosa, discovered by Joseph Banks in 1770 during his expedition with James Cook on HMS Endeavour. M. gulosa is among the earliest Australian insects to be described, and the specimen Banks collected is housed in the Joseph Banks Collection in the Natural History Museum, London. M. gulosa was described by Fabricius in 1775 under the name Formica gulosa and later designated as the type species of Myrmecia in 1840.
+A summary of the nine species groups described in the genus Myrmecia | ||||
M. aberrans species group | Wide-jawed bull ants | These medium-to-large ants are distinctive members of the genus. The mandibles and legs are short. They are found in the south-eastern region of Australia, and their colonies are small. Specimens have rarely been collected. | M. aberrans, Myrmecia formosa, M. forggatti, Myrmecia maura, and Myrmecia nobilis | |
M. cephalotes species group | These ants are characterised by their bright colours and black heads. Species of this group are medium in size and rare. Colonies dwell inland and they can be found in the eastern and western regions of Australia. | Myrmecia callima, M. cephalotes and Myrmecia hilli | ||
M. gulosa species group | Giant bull ants | Members of this group are large and slender, and have long legs. They are commonly found throughout most of Australia, although they are rarely or never found in the north-western coastal areas and Tasmania; one species has also been introduced to New Zealand. The mandibles vary in shape, and the number of teeth range from three to six. | Myrmecia analis, Myrmecia arnoldi, M. athertonensis, M. auriventris, M. borealis, M. brevinoda, M. browningi, Myrmecia comata, M. desertorum, M. dimidiata, Myrmecia erecta, M. esuriens, M. eungellensis, M. fabricii, M. ferruginea, M. flavicoma, Myrmecia forceps, M. forficata, Myrmecia fulgida, M. fuscipes, M. gratiosa, Myrmecia gulosa, Myrmecia hirsuta, Myrmecia inquilina, Myrmecia midas, M. minuscula, M. mjobergi, M. nigriceps, M. nigriscapa, Myrmecia pavida, M. picticeps, Myrmecia pulchra, M. pyriformis, M. regularis, M. rowlandi, M. rubripes, M. rufinodis, M. simillima, M. subfasciata, Myrmecia tarsata, M. tridentata, and Myrmecia vindex | |
M. mandibularis species group | Toothless bull ants | The ants of this group are medium-sized, and can be distinguished from other Myrmecia ants by their oddly shaped mandibles. While their bodies are black, their appendages may vary in colour. They are known to live in the eastern regions of Australia and Tasmania. Colonies have also been found in the south coastal areas and Western Australia. | M. fulviculis, M. fulvipes, M. gilberti, M. luteiforceps, M. mandibularis, M. piliventris and Myrmecia potteri | |
M. nigrocincta species group | These ants are medium in size with slender bodies and long legs, confined to the east of Australia. Members of this group look similar to those of the M. gulosa species group. | M. flammicollis, M. nigrocincta, and M. petiolata | ||
M. picta species group | These ants are small and can be found throughout southern Australia. This species group has only two members, making it the smallest of all the species groups. | Myrmecia fucosa and Myrmecia picta | ||
M. pilosula species group | Jack jumper ants | The majority of these ants are small in size, and colouration varies between species. They are distributed throughout Australia and Tasmania, and one member of this group is endemic to New Caledonia. The species group is known to be heterogeneous. | M. apicalis, Myrmecia banksi, Myrmecia chasei, M. chrysogaster, M. croslandi, Myrmecia cydista, Myrmecia dispar, Myrmecia elegans, Myrmecia harderi, M. haskinsorum, Myrmecia imaii, M. impaternata, Myrmecia ludlowi, M. michaelseni, M. occidentalis, M. pilosula, M. queenslandica, Myrmecia rugosa, and Myrmecia varians | |
M. tepperi species group | Buck-toothed bull ants | These ants are small or medium-sized, and have similar characteristics to the M. pilosula group. They are found in the south-western and south-eastern regions of Australia. | Myrmecia acuta, Myrmecia clarki, Myrmecia swalei, Myrmecia tepperi, and M. testaceipes | |
M. urens species group | Baby bull ants | Members of this group are all small, and colouration varies widely between species. Most specimens collected to date are from the coastal regions of Australia. | M. dichospila, Myrmecia exigua, Myrmecia infima, Myrmecia nigra, Myrmecia loweryi, M. rubicunda, and Myrmecia urens |
Their colouration is variable; black combined with red and yellow is a common pattern, and many species have golden-coloured pubescence (hair). Many other species are brightly coloured which Aposematism to avoid them. The Formicinae ant Camponotus bendigensis is similar in appearance to M. fulvipes, and data suggest C. bengdigensis is a batesian mimicry of M. fulvipes. The number of malpighian tubules differs between castes; in M. dispar, males have 16 tubules, queens range from 23 to 26, and workers have 21 to 29.
Worker ants are usually the same size as each other, although this is not true for some species; worker ants of M. brevinoda, for example, vary in length from . The mandibles of the workers are long with a number of teeth, and the clypeus is short. The antennae consist of 12 segments and the eyes are large and convex. Based on a study on the antennal sensory of M. pyriformis, the antennal Sensillum are known to have eight types. Large ocelli are always present.
Queens are usually larger than the workers, but are similar in colour and body shape. The head, node, and postpetiole are broader in the queen, and the mandibles are shorter and also broad. Myrmecia queens are unique in that particular species either have fully winged queens, queens with poorly developed wings, or queens without any wings. For example, M. aberrans and M. esuriens queens are ergatoid, meaning that they are wingless. Completely excavated nests showed no evidence of any winged queen residing within them. Some species have queens which are subapterous, meaning they are either wingless or only have rudiments of wings; the queens can be well developed with or without these wing buds. M. nigrocincta and M. tarsata are "Brachyptery", where queens have small and rudimentary wings which render the queen flightless. Dealated queens with developed wings and thoraces are considered rare. In some species, such as M. brevinoda and M. pilosula, three forms of queens exist, with the dealated queens being the most recognisable.
Males are easy to identify due to their perceptibly broad and smaller mandibles. Their antennae consist of 13 segments, and are almost the same length as the ants' bodies. Ergatandromorph (an ant that exhibits both male and worker characteristics) males are known; in 1985, a male M. gulosa was collected before it hatched from its cocoon, and it had a long but excessively curved left mandible while the other mandible was small. On the right side of its body, it was structurally male, but the left side appeared female. The head was also longer on the female side, its colour was darker, and the legs and prothorax were smaller on the male side. Male genitalia are retracted into a genital cavity that is located in the posterior end of the gaster. The sperm is structurally the same to other animal sperm, forming an oval head with a long tail.
Among the largest larvae examined were those of M. simillima, reaching lengths of . The pupae are enclosed in dark cocoons.
Ants of this genus prefer to inhabit grasslands, forests, heath, urban areas and woodland. Nests are found in Callitris forest, dry marri forest, Eucalyptus woodland and forests, mallee scrub, in paddocks, Riparian zone woodland, and wet and dry sclerophyll. They also live in dry sandplains, and coastal plain. When a queen establishes a new colony, the nest is at first quite simple structurally. The nest gradually expands as the colony grows larger. Nests can be found in debris, decaying tree stumps, rotten logs, rocks, sand, and soil, and under stones. While most species nest underground, M. mjobergi is an arboreal nesting species found on Epiphyte ferns of the genus Platycerium. Two types of nests have been described for this genus: a simple nest with a noticeable shaft inside, and a complex structure surrounded by a mound. Some species construct dome-shaped mounds containing a single entrance, but some nests have numerous holes that are constantly used and can extend several metres underground. Sometimes, these mounds can be 0.5 m (20 in) high. Workers decorate these nests with a variety of items, including charcoal, leaves, plant fragments, pebbles, and twigs. Some ants use the warmth by decorating their nests with dry materials that heat quickly, providing the nest with solar energy traps.
Most species are Diurnality, and forage on the ground or onto low vegetation in search of food, but a few are nocturnality and only forage at night. Most Myrmecia ants are active during the warmer months, and are dormancy during winter. However, M. pyriformis is a nocturnal species that is active throughout the whole year. M. pyriformis also has a unique foraging schedule; 65% of individuals who went out to forage left the nest in 40–60 minutes, while 60% of workers would return to the nest in the same duration of time at dusk. Foraging workers rely on landmarks for navigation back home. If displaced a short distance, they will scan their surroundings, and then rapidly move in the direction of the nest. M. vindex ants carry dead nest-mates out of their nests and place them on refuse piles, a behaviour known as necrophoresis.
Myrmecia is one of the very few genera where the workers lay , or infertile eggs laid as food for viable offspring. Workers laying trophic eggs have only been reported in two species; these species are M. forceps and M. gulosa. Depending on the species, colonies specialise in trophallaxis; queens and larvae eat eggs that are laid by worker individuals, but the workers do not feed on eggs. Neither adults nor larvae consume food during winter, but cannibalism among larvae is known to occur throughout the year. The larvae only cannibalise each other; this is most likely to happen when no dead insects are available.
The host association between Myrmecia and Eucharitidae wasps began several million years ago; M. forficata larvae are the host to Austeucharis myrmeciae, being the first recorded eucharitid Parasitoid wasp of an ant, and Austeucharis fasciiventris is a parasitoid to M. gulosa pupae. M. pilosula is affected by a Gregarinasina parasite that changes an ant's colour from their typical black appearance to brown. This was discovered when brown workers were dissected and found to have gregarinasina spores, while black workers showed no spores. Another unidentified gregarine parasite is known to infect the larvae of M. pilosula and other Myrmecia species. This gregarine parasite also softens the ant's cuticle. Other parasites include Beauveria bassiana, Paecilomyces lilacinus, Chalcura affinis, Tricoryna wasps, and various Mermithidae nematodes.
M. hirsuta and M. inquilina are the only known species in this genus that are inquilines and live in other Myrmecia colonies. An M. inquilina queen has been found in an Myrmecia vindex colony.
During the process of founding a colony, as many as four queens cooperate with each other to find a suitable nesting ground, but after the first generation of workers is born, they fight each other until one queen is left alive. However, occasional colonies are known to have as many as six queens coexisting peacefully in the presence of workers. A queen searches for a suitable nest site to establish her colony, and excavates a small chamber in the soil or under logs and rocks, where she takes care of her young. A queen also hunts for prey instead of staying in her nest, a behaviour known as claustral colony founding. Although queens do provide sufficient amounts of food to feed their larvae, the first workers are "nanitics" (or minims), smaller than the smallest workers encountered in older developed colonies. Several species do not have any worker caste, and solitary queens will raid a colony, kill the residing queen, and take over the colony. The first generation of workers may take a while to fully develop into adults; for example, M. forficata eggs take around 100 days to fully develop, while other species may take up to eight months.
Queens lay around eight eggs, but less than half of these eggs develop. Some species, such as M. simillima and M. gulosa, lay their eggs singly on the colony floor, while M. pilosula ants may lay eggs in a clump. These clumps have two to 30 eggs each with no larvae present. Certain Myrmecia species do not lay their eggs singly and form clumps of eggs, instead.
Both independent and dependent colony foundation can occur after mating. Isolation by distance (IBD) patterns have been recorded with M. pilosula queens, where nests that tend to be closer together were more genetically related to each other in comparison to other nests further away. Independent colony foundation is closely associated with queens which engage in nuptial flight in areas far from their home colony, showing that dependent colony foundation mostly occurs if they mate near their nest. In some cases, queens could seek adoption into alien colonies if there are no suitable areas to find a nest or independent colony foundation cannot be carried out. Other queens could try to return to their home nest after nuptial flight, but they may end up in another nest near the nest they originally came from. In multiple-queen societies, the egg-laying queens are generally unrelated to one another, but one study showed that it is possible for multiple queens in the same colony to be genetically related to each other. Depending on the species, the number of individuals present in a colony can range from 50 to over 2,200 individuals. A colony with less than 100 workers is not considered a mature colony. M. dispar colonies have around 15 to 329 ants, M. nigrocincta have over 1,000, M. pyriformis have from 200 to over 1,400 and M. gulosa have nearly 1,600. A colony can last for a number of years. Foraging behaviour among smaller workers which never usually leave the nest can be a sign of a colony's impending demise.
Workers are known to produce their own eggs, but these eggs are unfertilised and hatch into male ants. There is a chance of workers attacking a particular individual who has successfully produced male offspring due to a change in a workers cuticular hydrocarbon; cuticular hydrocarbons are believed to play a vital role in the regulation of reproduction. However, this is not always the case. Myrmecia is one of several ant genera which possess gamergate workers, where a female worker is able to reproduce with mature males when the colony is lacking a queen. Myrmecia workers are highly fertile and can successfully mate with males. A colony of M. pyriformis without a queen was collected in 1998 and kept in captivity, during which time the gamergates produced viable workers for three years. Ovarian dissections showed that three workers of this colony mated with males and produced female workers. Queens have bigger ovaries than the workers, with 44 ovarioles while workers have 8 to 14. Spermatheca is present in M. gulosa workers, based on eight dissected individuals showing a spermatheca structurally similar to those found in queens. These spermathecas did not have any sperm. Why the queen was not replaced is still unknown.
Fatalities associated with Myrmecia stings are well known, and have been attested to by multiple sources. In 1931 two adults and an infant girl from New South Wales died from ant stings, possibly from M. pilosula or M. pyriformis. Another fatality was reported in 1963 in Tasmania. Between 1980 and 2000, there were six recorded deaths, five in Tasmania and one in New South Wales. Four of these deaths were due to M. pilosula, while the remaining two died from a M. pyriformis sting. Half of the victims had known ant-sting allergies, but only one of the victims was carrying adrenaline before being stung. Most victims died within 20 minutes of being stung, but one of the victims died in just five minutes from a M. pyriformis sting. No death has been officially recorded since 2003, but M. pilosula may have been responsible for the death of a man from Bunbury in 2011. Prior to the establishment of a desensitisation program, Myrmecia stings caused one fatality every four years.
Emergency treatment is only needed if a person is showing signs of a severe allergic reaction. Prior to calling for help, stung persons should be laid down, and their legs elevated. An EpiPen or an Anapen is given to people at risk of anaphylaxis, to use in case they are stung. If someone experiences anaphylactic shock, Epinephrine and intravenous infusions are required, and those who suffer cardiac arrest require resuscitation. Desensitisation (also called allergy immunotherapy) is offered to those who are susceptible to M. pilosula stings, and the program has shown effectiveness in preventing anaphylaxis. However, the standardisation of M. pilosula venom is not validated, and the program is poorly funded. The Royal Hobart Hospital and the Royal Adelaide Hospital are the only known hospitals to run desensitisation programs. During immunotherapy, patients are given an injection of venom under the skin. The first dose is small, but dosage gradually increases. This sort of immunotherapy is designed to change how the immune system reacts to increased doses of venom entering the body.
Before venom immunotherapy, whole-body extract immunotherapy was widely used due to its apparent effectiveness, and it was the only immunotherapy used for ant stings.
Notable Australian poet Diane Fahey wrote a poem about Myrmecia, which is based on Schopenhauer's description, and a music piece written by German composer Karola Obermüller was named after the ant.
Distribution and habitat
Behaviour and ecology
Foraging
Pollination
Diet
Predators, parasites and associations
Life cycle
Reproduction
Vision
Sting
Interaction with humans
Venom
Sting treatment
Prevention
Human uses
Cultural representations
See also
Notes
Cited texts
External links
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