Meat ant

Meat ant
Meat ant worker feeding on honey
Not evaluated (IUCN 3.1)
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Dolichoderinae
Tribe: Leptomyrmecini
Genus: Iridomyrmex
Species: I. purpureus
Binomial name
Iridomyrmex purpureus
(Smith, 1858)
Synonyms[1]
  • Liometopum aeneum Mayr, 1862
  • Iridomyrmex detectus castrae Viehmeyer, 1925
  • Formica detecta Smith, F., 1858
  • Iridomyrmex greensladei Shattuck, 1993
  • Camponotus horni Kirby, 1896
  • Formica smithii Lowne, 1865

The meat ant (Iridomyrmex purpureus), also known as the gravel ant or southern meat ant, is a species of ant endemic to Australia. A member of the genus Iridomyrmex in the subfamily Dolichoderinae, it was described by British entomologist Frederick Smith in 1858. The meat ant is associated with many common names due to its appearance, nest building behaviour and abundance, of which its specific name, purpureus refers to its coloured appearance. It is among the best known species of ant found throughout Australia, being found in almost all states and territories except for Tasmania. Its enormous distribution and ecological importance has made this ant a dominant species.

The meat ant is monomorphic, although there is evidence that some certain populations can be polymorphic. It is characterised by its dark-bluish body and red head. It is a medium to large species, measuring 6–12 mm (0.24–0.47 in). The workers and males are approximately the same size at 6–7 mm (0.24–0.28 in) and 8 mm (0.31 in), respectively. The queens are the largest and appear mostly black, measuring 12.7 mm (0.50 in). The iridescence in workers varies, varying from green/blue to plain green and purple. It also varies in different body parts and castes. Meat ants inhabit open and warm areas in large, oval-shaped mounds that are accompanied by many entrance holes. The nest area is always cleared of vegetation and covered in materials including gravel, pebbles and dead vegetation. They are also polydomous, where a colony may be established in a series of satellite nests connected by well defined paths and trails.

Queens mate with a single male, and colonies may have more than one queen until the workers arrive, where they both exhibit antagonism. It takes around one or two months for an individual egg to develop into an adult, and colonies may house over 300,000 ants. It is a diurnal species and are active throughout most of the day, especially when it is warm. They forage on trees and collect sweet substances such as honeydew and nectar, and also capture insects or collect the remains of animals. A number of predators eat these ants, including the short-beaked echidna (Tachyglossus aculeatus), numerous species of birds, blindsnakes and spiders. This species is also a competitor of the banded sugar ant (Camponotus consobrinus). Meat ants establish territorial borders with neighbouring colonies and solve disputes through ritualised fighting.

Meat ants play an important role in both the environment and for humans. A single nest is capable of dispsering over 300,000 plant seeds and they have formed symbiotic relationships with many insects and may be used as a form of pest control to kill the cane toad, an introduced pest. They can also help farmers to remove animal carcasses by consuming and reducing them to bones in a matter of weeks. Despite this, meat ants can be pests around urban areas and eradicating them is difficult.

Taxonomy

Published illustration of a worker by W.W. Froggatt, 1907

The meat ant was first identified in 1858 by British entomologist Frederick Smith in his Catalogue of hymenopterous insects in the collection of the British Museum part VI, under the binomial name Formica purpurea from a holotype worker he collected in Melbourne, Victoria.[2] The type material is currently housed in the Natural History Museum, London.[3] In 1862, Austrian entomologist Gustav Mayr erected the new genus Iridomyrmex, of which he transferred the meat ant from the genus Formica to there.[4] In Mayr's 1862 journal article, the ant was now known as Iridomyrmex purpurea, but he would rename it to Iridomyrmex purpureus in his 1863 journal article.[5] I. purpureus was later classified as a synonym of Iridomyrmex detectus, and this taxon was designated as the type species in 1903.[6][7][8] In 1993, I. detectus was synonymised with I. purpureus and grouped into the I. purpureus species complex.[9] I. detectus was described as a separate species by Smith from two syntype queens he collected from the Hunter River in New South Wales. Although the name I. purpureus was used by many authors, Karl Wilhelm von Dalla Torre reused I. detectus and this was adopted by several authors, replacing I. purpureus.[9] By the 1960s, however, I. purpureus was reused and authors gradually favoured the name instead of I. detectus. While the two names were described in the same year and publication, the name I. purpureus is the valid name for the species in accordance to the International Code of Zoological Nomenclature. As entomologist B.T. Lowne was the first reviser of I. purpureus and I. detectus, Lowne synonymised I. detectus with I. purpureus.[9][10]

Several synonyms have either been classified as a separate species from the meat ant or even a subspecies. Iridomyrmex purpureus sanguineus, Iridomyrmex purpureus viridiaeneus and Iridomyrmex detectus castrae were classified as three subspecies of the meat ant, but I. purpureus sanguineus and I. purpureus viridiaeneus were found to be two separate species whereas I. detectus castrae was synonymised.[9][11] Camponotus horni, Formica smithii and Liometopum aeneum were described as three distinct species in three different ant genera, but these classifications were short-lived as scientists discovered that they were all the same and subsequently synonymised them with the meat ant.[9][12] Camponotus horni was described by English entomologist William Forsell Kirby in 1896 from worker and queen syntypes he collected at Palm Creek in the Northern Territory. In 1930, Australian entomologist John S. Clark reviewed these specimens and synonymized it with the meat ant. However, the queen was found to be distinct from the workers and the name C. horni was redescribed as a species. The information provided by Clark makes it sufficient to retain the validity of the name, making Clark the new author of C. horni. Due to the uncertain taxonomic status of Clark's C. horni, a proposal to rename the species cannot be done. Clark's C. horni is currently a junior homonym for Kirby's C. horni.[9]

Syntype specimen of Iridomyrmex detectus castrae, currently housed in the Natural History Museum of Geneva[13]

One synonym, Iridomyrmex greensladei, was described as a separate species from the meat ant by the anatomical differences of its head and pronotum, which are concolorous with its mesosoma. In the meat ant, the head and pronotum are usually lighter than the mesosoma.[3] Steve Shattuck notes that the two ants are nonsympatric, and supporting evidence to separate them as two distinct species emerged when the enzymes esterase and amylase were examined and results show that they had different amylase allele.[14][15] These examinations, however, did not take any account of any distinction in colour between populations previously known as I. greensladei and I. purpureus. For example, colonies residing in Western Australia may have lighter heads whereas those living in the eastern states have darker heads. An unpublished study studying the mitochondrial DNA did not find any distinction between eastern and western populations of I. greensladei and I. purpureus.[3] Additional evidence suggests that the populations found in the east and west are the same species.[3] Despite this evidence, there are several problems: forms intended to be I. greensladei were found in the Yorke Peninsula by Greenslade (1987), noting that while the ants were morphologically indistinguishable from both I. greensladei and I. purpureus populations, their nests only contained a single nest hole (whereas I. purpureus nests have many).[16] Shattuck also collected workers for his holotype and paratype specimens from Israelite Bay in Western Australia, rather than collecting those from Yorke Peninsula. The forms in Yorke Peninsula have also never been subject to study, so future research may shed light as to whether or not these ants are genetically different from the meat ant.[3]

Under the present classification, the meat ant is a member of Iridomyrmex in the tribe Leptomyrmecini, subfamily Dolichoderinae. It is a member of the family Formicidae, belonging to the order Hymenoptera.[17] The genus Froggattella is the sister group of Iridomyrmex, both of which are in a clade that is 23 million years old.[18] Fossils related to the meat ant and other species are from the Eocene and Oligocene.[19]

Forms

Differentiated colouration of heads of I. viridiaeneus (left) and I. sanguineus (right). These ants were classified as two subspecies of I. purpureus until they were raised to full species in 1993.

In the early 1970s, scientists discovered different forms of meats ants, forming the Iridomyrmex purpureus species group; three forms were identified (the regular I. purpureus and the other two forms sanguineus and viridiaeneus), differing in colour and nest structure. However, as there were no clear morphological differences among these variations, the taxonomic status and evolutionary relationship of these ants remained uncertain.[20][21] Colour varies from having a red head and thorax with a blue metallic sheen, to a yellowish body with a bluish-purple gaster. Others may be darker, appearing metallic-bluish to purplish-black.[20] Owing to their different geographical ranges, the behaviour and ecology among these forms differentiated, based on examinations where multiple forms were sympatric.[21] Each form had its own geographical limit, although the regular I. purpureus form is frequently encountered (inhabiting at least one-third of the continent). Some forms may be dominant in certain habitats; for example, one form may be widespread throughout moist environments in dry areas, and others in cool and dry areas.[20]

In 1974, I. sanguineus and I. viridiaeneus were classified as two subspecies of the meat ant by P.J. Greenslade. He also recognised two new variants, viridiaeneus in a strict sense and an undescribed blue form that were first studied several years eaelier.[22][23][24] Greenslade did not propose any taxonomic recognition of the newly discovered taxon, but instead he would still refer it as a "form".[9] In 1975, a study examined the distinctiveness of three forms by comparing the amylase enzyme allele frequencies. The blue form was found to be genetically isolated and its allele frequencies differed significantly from the forms purpureus and viridiaeneus, but the same study concluded that these two forms were similar.[25] During this time, Greenslade would continue to study these variants to include a dark yellow, yellow, orange and a small purple variant.[9] In addition, two new forms were recognised in later studies, but these two forms were given no formal species description. Instead, they were only recognised from their colour and the genitalia of the males. Although these forms most likely represented undescribed, yet distinct species, the I. purpureus species group only contained a single valid species, three subspecies and seven different forms.[9] In 1993, all known forms and subspecies were either revived or classified at species level in an article revising the group, concluding the separation of the forms from each other.

Etymology and common names

The specific epithet of the meat ant, purpureus, derives from Latin, meaning "purple", "purple-coloured" or "dark-red", referencing the coloured appearance of the ant.[26] In classical Latin, purpureus primarily translated to "dyed purple", while the word purpura, originally used as the specific name for the meat ant (Formica purpura), translates to "purpled-dyed cloth".[2][27] The name of the genus of the meat ant, Iridomyrmex, translates to "rainbow ants", another reference which points to its appearance; this in particular is due to its blue-green iridescence colour.[28] The word Irido, meaning "rainbow", derives from Ancient Greek,[29] and myrmex, another Greek word, means "ant".[30] The ant is commonly known as the meat ant because of its habit of stripping meat off dead vertebrates, but they are also known as meat-eater ants, gravel ants, Greenslade's meat ants or southern meat ants.[31][32][33][34] The name "southern meat ant" is due to its extreme abundance in the southern regions of Australia.

Description

Worker specimen of I. purpureus

In general, meat ants are medium to large in size, measuring 6–12 mm (0.24–0.47 in) and can be easily recognised by their coloured appearance, appearing with a dark-bluish body and red head.[35] Most of the time their heads and pronotums share similar colours and are lighter than the mesothorax and the propodeum, which are reddish brown.[9] However, the head may sometimes be lighter and the pronotum and mesothorax share similar colours. The mesosomal setae are dark and sometimes translucent. The iridescence between the compound eyes and the lateral portion of the head ranges from slightly purple to strong and dark purple. The colour of the legs and coxae (the basal segment of the insect leg, which attaches to the body) are darker than the mesothorax, and the petiole is reddish brown and also darker than the mesothorax.[9] In the lateral portion of the second gastral tergite, it is shiny and the iridescence varies among workers; the iridescence varies from green/blue to plain green and purple. The pilosity on the head is frequent around the occipital margin, and around the mandibular insertion, three to eight pale setae are usually seen. Examined specimens show no known ocelli. Erect setae on the pronotum are abundant, and the pilosity is common around the first gastral tergite. The anterodorsal portion of the propodeum is arched and flat.[9]

There are no allometric differences among workers.[3] On average, workers measure around 6–7 mm (0.24–0.28 in).[36] The head and pronotum range from orange to brick-red, and the mesonotum and propodeum are either light, concolorous or darker than the head. The gaster can be brown or black with blue or purple iridescence and the legs are either orange or brown. The iridescence around the foreparts are blue, pink, pale greenish yellow and purple. The erect setae are brown. The head has a concaved posterior margin with erect setae abundant infront of the face and the sides of the head are convex shaped.[3] Fully erect genal setae are either seen on the sides or the head or absent, although a small number of setae may be seen around the mandibular insertion. The eyes are semicircular and positioned around the midpoint of the ants' head capsule. The frontal carinae is convex and the antennal scapes extend beyond the head's posterior margin by two or three times the diameter. Erect setae are found all over the antennal scape and noticeably prominent on the clypeal margin. The mandibles are elongated and triangular in shape with long curved setae around the head capsule.[3] The pronotum is evenly curved with at least 12 or more pronotal setae present. This setae are mostly short and bristly. The mesonotum is sinuous (meaning it has many curves), and like the pronotum, has 12 or more mesonotal setae. The mesothoracic spiracles are very small and the propodeal dorsum is smooth or convex. There are also a number of propodeal setae seen.[3] The dorsum of the node (a segment between the mesosoma and gaster) is thin, scale-like and sometimes vertical. Around the gaster there are both non-marginal and marginal setae present on the first gastral tergite.

Specimen image of a male

Queens are easily distinguishable from workers, measuring 12.7 mm (0.50 in).[2] Queens are black and fuscous, being mostly dark and sombre. The antennae and legs are ferruginous, the head is fusco-ferruginous, and the sides beneath the face and mandibles are ferruginous. The head is wider than the thorax and emarginate. There is an impressed line that runs from the anterior stemma to the base of the clypeus.[2] The thorax is ovate-shaped and thinly covered with short reddish brown pubescence (soft short hair). The wings are subhyaline, exhibiting a glassy appearance. The wings are yellowish along the anterior margin of the superior pair and also around the base; the nervures (the veins of the wings) are rufo-fuscous.[2] Like the thorax, the abdomen is ovate and several abdominal segments appear rufo-piceous.[2] Males are smaller than the queens, measuring 8 mm (0.31 in). Males are bright violet, and the antennae (except for the first joint) and tarsi are ferruginous.[10] The first pair of the legs almost look ferruginous, and the head, legs and thorax are covered in black pubescence. Like the queen, the wings are subhyaline and the nervuvres are rufo-fuscous. The abdomen shows a bright green tinge when they are seen under certain lights.[10]

Microscopic view of an eye

The larvae measure 2.7–2.9 mm (0.11–0.11 in).[37] The body is stout-shaped and the dorsal side is longer whereas the ventral side is shorter and more straight. The head and anus are ventral. The integument is covered in spinules that are either isolated from each other or in short rows on the posterior somite and on the ventral surface. The body hairs are very short, measuring 0.008–0.016 mm (0.00031–0.00063 in).[37] The cranium on its dorsal side outline a curve which is smooth, and the spinules are moderately large. These spinules are either isolated or seen in sub-parallel rows. Several head hairs are present but they are small at 0.013–0.025 mm (0.00051–0.00098 in) in length. The labrum is narrow and bilobed, consisting of two lobes. Each lobe has spinules and three sensilla around the anterior surface. The ventral border only has two sensilla and a number of spinules, and on the posterior surface there are several rows of spinules and three sensilla. The mandibles have a central apical tooth which are clearly noticeable and sharp. The maxilae are lobose, and the labial palps are knob-shaped.[37]

Workers of the meat ant may be confused with I. lividus, as the two appear similar and are grouped in the same species complex.[3] Like I. lividus and the much more localised I. spadius, these ants can be distinguished from other members of the I. purpureus group by the shape of the pronotum when specimens are being examined. Aside from colour differentiation that was a key morphological character to distinguish I. purpureus and the synonym I. greensladei from each other, some meat ant populations exhibit polymorphism despite being monomorphic. Examined workers once referred to as I. greensladei from the south-west of Western Australia have erect setae on the genae, whereas those studied elsewhere have glabrous genae.[3] Such patterns are most likely clinal, where several characteristics may gradually differentiate over a geographical area.[9] The colour of the setae found throughout the ants body, and to some extent, the iridescence, may also vary. For example, populations restricted to the southern coasts of Western Australia usually have pale setae, compared to most colonies throughout the country having the common blackish setae.[3] However, Shattuck (1993) could not separate populations with pale setae when other and more important key characteristics were used to separate species.[9] Meat ants from the Western Australian wheatbelt and goldfields show different iridescence; the iridescence in some collected specimens ranged from pale greenish-blue to yellowish-green, especially around the humeri and frons. The variation of the iridescence is, however, a consistent pattern found in other Iridomyrmex species with little distinction, making it a subtle character.[3] The colour variation is less marked in all collected specimens of I. purpureus, as well as its close relative I. viridiaeneus, which is found in dry regions around the south-western regions. Shattuck further notes that populations found throughout the Northern Territory and South Australia have reduced pubescence on the first gastral tergite, but this varies elsewhere.[3]

Distribution and habitat

Reported occurrences of meat ant populations according to the Atlas of Living Australia

The meat ant is one of the best known species of ant endemic to Australia with an enormous geographical range, covering at least one third of the entire continent.[20][38] It spans an east to west range of 4,000 kilometres (2,500 mi), and 3,000 kilometres (1,900 mi) from north to south.[39] This large range has allowed the meat ant to form large nesting grounds in areas where no development has occurred, and the large amount of gravel and open space has led to an abundant supply of materials used to construct nests. Its isolation has also allowed meat ants to form associations with neighbouring nests.[40] The ant is particularly dominant and frequently seen across the coastal and inland regions of southeastern Australia.[41][42] Based on examined material, the meat ant is common in the southwestern regions of Western Australia but in the north it is less common.[3] However, The CSIRO Division of Entomology claims the ant's presence in the state has not been verified.[43] In the Northern Territory, specimens have been collected in the north and south regions, but compared to other jurisdictions the ant is uncommon.[3] Most specimens collected in South Australia are from the southeast, but some populations are known in the northwest and northeast regions of the state. In Queensland, they are frequently encountered along the eastern areas of the state, whereas their abundance is limited around the north and central parts. They are widespread throughout New South Wales, the Australian Capital Territory and Victoria.[3][9] No specimens have been collected from Tasmania.

Meat ants can thrive in many habitats, especially where it is open and warm.[44] These ants are adapted to warm climates because of their large distribution and the fact that temperatures are always warm. The meat ant shares its distribution with many other animals and insects, some of which may cause harm to the ants or form rivalries with other ant species, such as the banded sugar ant (Camponotus consobrinus).[40][45] Body size among meat ants may vary depending on their location; particularly, those that are found in very hot regions tend to be larger, whereas others are smaller in regions with high humidity.[39] Nests are seen in Box-pine scrubs, Callitris forests, dry sclerophyll woodland, Eucalyptus woodland and eucalypt open woodland, in farm pastures, flat savannah woodland, mallee woodland, heath, mulga, riparian woodland, around roads and cracks in sidewalks, urban areas such as urban gardens and parkland and wet sclerophyll woodland.[32][41][46] They are also common in lateritic ridges, granite outcrops and clay formations. Meat ants are able to survive in dry areas if there is a rich supply of water and food resources, especially along river banks, station properties and irrigated areas. Meat ants are mostly found at altitudes of between 5 and 1,170 m (16 and 3,839 ft) above sea level, but most of the time they are not found above 915 m (3,002 ft).[20][46] Those that are found at such altitudes are always associated with Eucalyptus rubida, and colonies situated in eastern New South Wales tend to nest near Eucalyptus melliodora and Eucalyptus blakelyi. In the south coast of New South Wales, meat ants are mainly found in heath land, but they are absent around heavily timbered slopes. They also do not occur in dense pasture. For example, the Canberra suburb of Turner was constructed on subterranean clover pasture, of which meat ants do not nest around. Their populations would later flourish and nests were numerous around houses after shrubs and trees were planted.[20]

Nests

Meat ant mound near Bungendore, New South Wales

Meat ants are well known for their large, oval-shaped and obvious mounds (normally one or two metres in diameter) that are frequently encountered.[22][32][47][48] These nests are often accompanied with many entrance holes; most studied nests usually have 20 to 35 holes. On the surface on the nest, workers clear the area of vegetation and cover the mound with gravel but may use other materials that are available, including sand, pebbles dead vegetation, eucalyptus fruits and twig fragments.[32][47] Nests are often found in the sun and never in shaded areas, which allow the nests to warm up. The ant is a polydomous species, meaning that they live in more than one nest.[49] Some colonies are known to create "super-nests": workers construct many nests connected through established paths, extending up to 650 metres (2,130 ft) in length.[19] In one extreme case, a single colony was found to occupy over 10 hectares of land with 85 individual nests and 1,500 entrance holes.[50] While meat ants are never aggressive to their nest mates, they will be aggressive to those who live in different nests within the same colony.[49]

At any size, a meat ant nest always has a possibility of dying out in any year, but the site may be reoccupied by another colony. Such scenario leads to the extreme ageing of nests as suggested in one study.[51] Some nests may never be reoccupied, although it is unknown why they are not. The regrowth of vegetation which shades the nest, soil damage or even a disease may wipe out a colony and leave the nest site completely abandoned.[47][52] The death of a colony may be obvious when competing colonies increase in population and size or satellite nests are well established in order exploit the food sources once used by a former colony. Satellite nests may diverge from their parent nests to become independent, as suggested by the antagonism of worker ants from different nests or when others are uninhabitable by insecticide treatment. After the eradication of a nest, satellite nests emerge nearby and may sever their connections with the parent nest.[53][54] The separation of a satellite nest is an effective way for a nest to exploit food sources, and a colony with less than 11 entrance holes are recognised as non-mature (a nest is mature at 12). As most satellite nests have 11 holes and accepted a queen of their own, a satellite nest may easily develop maturely in one year.[47]

Ant mound holes prevent water from entering the nest during rain

All entrance holes lead into a separate set of galleries.[55] Entrance holes tend to be very small with only enough space for a single worker to fit through, but others may be 1 centimetre (0.4 in) wide. Beneath the surface, there are widened circular vertical shafts which are 1.5 centimetres (0.6 in) wide. Below these shafts, they turn into irregular gallery systems with paths going outward and downward which form more galleries. Almost all of these galleries are clustered together 15 to 20 centimetres (5.9 to 7.9 in) beneath the soil, of which the area is extensively reworked on by workers.[55] These sets of galleries are isolated from others connecting to neighbouring holes, although some may come close. However, there is no known physical connection. Each gallery has a flat floor, a domed roof and irregularly oval-shaped. A gallery is typically 1.5 centimetres (0.6 in) high and 5 to 7 centimetres (2.0 to 2.8 in) wide. Below the galleries are a small number of shafts in undisturbed soil with large, yet scattered chambers where the population remains during the winter. In the upper areas of the colony, the galleries and shafts only account for 7-10% of nest space. For example, the galleries of excavated colonies occupy 420 cubic centimetres (26 cu in) whereas the total volume is around 5,000 cubic centimetres (310 cu in). Overall, a nest may dwell extremely deep beneath the soil, as excavated nests are as deep as 3 metres (9.8 ft).[55]

Behaviour and ecology

Foraging and dominance

Worker foraging on vegetation

The meat ant is a diurnal species and there is little overlap when they and nocturnal ants are active at the same location.[56] The meat ant and other members of the I. purpureus species group are regarded as a dominant group of diurnal species in Australia, as they are highly active, aggressive, and have an enormous geographical range.[57] During the day, the bulk of the workers attend and feed from secreting insects that give them honeydew on the foliage of Eucalyptus trees.[47] These trees are connected by foraging trails formed by meat ants, and back to other nests that are apart of a single colony (satellite nests);[47] these trails are easily visible due to the absence of vegetation, and are important for food sources and water, especially on hot and dry days.[47] Other workers in smaller numbers will often scavenge around the nest area for dead insects, or other proteinaceous foods to bring back to their colony.[47] Their dominance is clear when meat ants are capable of finding food sources more successfully than any other species, capable of displacing many ants and other insects while they themselves cannot.[58][59] Food sources are also discovered more rapidly and greatly reduces the foraging efficiently of other ants, including those of Iridomyrmex. However, some ants such as the green-head ant (Rhytidoponera metallica) are not affected by the presence of meat ants and are still successful with finding food sources.[59] As green-head ants are general predators and forage singularly and cannot recruit other nestmates, they are unable to defend food sources from dominant ants. They heavily rely on any food source and the impossibility of successfully defending it from other ants may have led to its peaceful coexistence with dominant species, including meat ants.[59]

Nests do not occur in shaded areas and are located in open areas so the nest can warm up in the morning.[60] The rate of movement of workers and temperature is associated with each other, and foraging workers leave the nest after sunrise when the nest is warmed in the early morning.[47] Meat ants emerge and start foraging when temperatures are approximately 14 °C (57 °F), but they are capable of withstanding extremely hot temperatures, as workers have been seen foraging in 43 °C (109 °F) heat.[61] Workers may only forage for several minutes when soil temperature is well above their thermal limit at 45.8 °C (114– °F). At 50 °C (122 °F) (soil surface temperature), foraging activity is reduced even further. As these ants exhibit fast running speeds during hot days, their speed deceases after 50 °C due to the physiological stress put onto the ants. Workers can withstand such temperatures by exhibiting opportunistic thermal responses to adjust their behaviour in order to handle high trail temperatures and alter their thermal biology for low temperatures. They will also deliberately expose themselves closely to their thermal maxima.[62]

Foraging trail with a satellite nest vaguely seen (top left)

The impact of meat ants among other species varies; this depends on the habitat and type of food source and the ecological role the ants plays.[59] The deliberate exclusion of meat ants show that not only they can alter the amount of sources available, but the dynamic of resource use on entire outcrops. If present, meat ants rely on rapidly discovering food sources and retain their dominance so other ants cannot collect them, as well as exploitation and interference which helps displace other ants. The habitat meat ants live in may affect their dominance among the fauna. They are less successful in complex habitats and more successful in open areas, allowing workers to forage efficiently; for example, workers forage around rocks and collect food sources more successfully in contrast to those in vegetation.[59] Meat ants are even more dominant than other I. purpureus species group members in non-complex habitats. Meat ants are normally larger than other Iridomyrmex species and can displace or even kill those trying to compete for food sources. The similar behaviours among meat ants and certain Iridomyrmex species has increased the level of conflict among each other, especially those in open habitats.[59] In foraging areas where meat ants are excluded or not present, the number of ants observed are far greater and activity dramatically increases. However, the presence of meats ants are unable to affect abundant species, and most of the time it is only Iridomyrmex ants that increase their activity rate.[58][57][63]

The success of meat ants also depends on what resource they're dealing with. In one study, the removal of meat ants allowed the successful foraging of Iridomyrmex ants at carbohydrate sources, but not for protein baits.[59] The success for the ants also did not increase with mealworm baits, suggesting that competition does not always occur with meat ants and other factors may be the reason (i.e. food preference). In contrast with camponotines and some myrmicines, their success was affected at numerous food sources with meat ants present. Camponotines exhibit no aggression and cannot defend themselves, whereas the myrmicines, which are highly competitive, are capable of defending themselves from foraging workers; the rapid recruitment of nestmates and the use of chemicals for defence allows them to withstand attacks from meat ants. This is not the case with mealworms, but they have semi-success with honey baits.[59]

Diet

Worker feeding on honey

Like other Iridomyrmex species, the meat ant is an omnivore, retrieving food sources from various insects they tend to including caterpillars and various sorts of butterflies, particularly the larvae of the Waterhouse's hairstreak (Jalmenus lithochroa).[32][64] Meat ants usually feed on honeydew from sap-sucking insects, flower nectar, sugar and other sweet substances.[41][65] These ants prey on various insects and animals, collecting both live and dead invertebrates and acquire meat from dead vertebrates.[65] They will prey on crucifix toads going through metamorphism.[66] Insects the meat ants prey on include giant lacewings of the genus Ithone, where foraging workers will swarm up on trees to kill them,[67] Ogyris genoveva butterflies,[68][69] Indian mealmoths, almond moths, the Western Australian jarrah leafminer and Trichogramma wasp larvae.[70][71][72] Large and developed larvae of the cabbage butterfly (Pieris rapae) are often attacked by workers in an effective manner, whereas the larvae are usually left alone by smaller Iridomyrmex species.[73] On sandy beaches, this species is observed preying on the polychaete annelid worm, Armandia intermedia, causing high mortality rates on them (rates of 30 percent).[74] These ants will feed on a number of dead animals, including snakes, lizards, birds and on some occasions, swarms of works have been found on dead foxes.[75][41] The meat ant is the only known ant in Australia that feeds on fresh guano.[76] The collection of guano by a nearby meat ant colony shows the opportunistic nature of the species. Observations show that trails of workers in groups of two to four were found collecting the guano under an active bat roost within an abandoned mine and proceeded to return it to the nest. The collection of guano by any Australian ant was never recorded prior to these observations, but it is unknown why meat ants collect fresh guano.[76]

Meat ants cooperating to devour a cicada

Meat ants are among the very few native species of Australia that are not harmed by the toxins of the cane toad, an invasive species. Most of the time foraging workers target metamorph toads.[77][78] Observations show that workers would forage around ponds and seize any toadlet. This normally starts with a single worker making contact with an individual and tracing its movements, followed by three or four workers capturing it. All the participating ants would grab a limb and sever the legs, returning the prey item back to their nest. Most of the time these tactics do not work. For example, most toadlets are able to escape the ants by displaying aversion-like behaviour; an individual may escape by struggling erratically or forcing the ants to release their grip by returning to water. This aversion behaviour leads to most toadlets remain in the water or stay on objects such as water lilies, pieces of bark that the ants cannot access to or move around on moist substrate.[77] It is unknown if meat ant predation on the toadlets affected the population, but based on the population density of the toadlets within the studied site and the foraging time and efficiency, approximately 2,700 toadlets could be removed per day. As the toadlet population density is extremely high, the impact of meat ant predation is minor. However, the survivability of the toadlets may be affected if the ants prevent the toadlets from foraging into many areas of moist substrate.[77]

Water is an important resource for colonies living in dry and arid environments, but sometimes it may not be available. To counter this, workers are able to extract significant amount of water from sand with 2% to 4% water content and 4% from soil.[79] Meat ants are unable to retrieve as much water from soil, whereas with sand they are able to attain a greater amount of water; however, the soil contains a wide variety of particles, including clay and coarse sand which causes water to be bound firmly. They may retrieve it by digging or directly suck on the soil itself at a low metabolic cost. This may be an evolved response based on tested ants, but no observations show meat ants doing this. Meat ants are known to dig into moist soil to gain access to water or where water has been spilled, either if the site is nearby their nest or far away.[79]

Predators

While the meat ant is a dominant species, a number of animals are known to prey on them. The short-beaked echidna (Tachyglossus aculeatus) is a prominent predator of the meat ant, mostly due to the virgin queen ants present containing high levels of fat.[80] These queens can almost contain 47% fat, and so when no queens are available after an attack, an echidna may stop attacking the nest.[81] However, these ants are normally consumed either in low numbers or avoided entirely.[80] Attacking echidnas would burrow down in the hole they have made and consume them (mostly for the queens) while handling the bites from the ants, as they frequently scratch themselves on the head and chest.[82][83] The echidna does not consume meat ants throughout the whole year;[83] Instead, echidnas in Canberra would only attack meat ant nests from August to October, which is when nuptial flight usually occurs.[83] This particular time period makes it much easier for echidnas to prey on the winged females since they are directly above on the nest.

Nest damage from a feeding echidna

Several birds prey on meat ants. The masked woodswallow (Artamus personatus) and the white-browed woodswallow (Artamus superciliosus) will gather around on their nests and swoop them, catching several ants before eating them.[84] Pieces of meat ants have been found in the red-capped robin (Petroica goodenovii), rufous whistler (Pachycephala rufiventris), hooded robin (Melanodryas cucullata) and the red-browed treecreeper (Climacteris erythrops).[85] Meat ants who foraged on Ventilago viminalis trees were often eaten by the apostlebird.[86] Some large ground-feeding birds, such as currawongs, magpies and ravens will dig out newly established colonies after a queen has found a suitable spot to nest. Since small domes of excavated soil is present, this will reveal their presence to these birds.[47] Due to this, many queens will be eaten by these birds, leaving many abandoned nest chambers.

The blind snake Ramphotyphlops nigrescens follow trails laid by meat ants to possibly to locate them as a potential prey species, and they are also known to eat on the brood of this species.[87] Various species of spiders prefer to prey on meat ants, which are mainly attracted by the alarm pheromone the ants release.[88] One spider in particular, the cursorial spider Habronestes bradleyi, is a specialist predator against these ants and will use their alarm pheromones that is released during territorial disputes to locate them.[89][90][91]

Moths of the genus Cyclotorna, particulartly Cyclotorna monocentra feeds on meat ant brood.[92] The larvae of these moths are parasites to leafhoppers, and will move to meat ant colonies to complete their development, where they will proceed to consume the brood.[92][93] Females will lay many eggs near ant trails which are close to the leafhoppers which the ants attend to.[94] Other observations show that the larvae of the moth species Iphierga macarista is a scavenger in meat ant nests,[92][95] while Sphallomorpha beetles live in burrows near nests of meat ants. Sphallomorpha larvae can be found in nests which capture and prey on workers passing by.[96] The larvae of the spitfire sawfly and Pseudoperga guerini are able to regurgitate a fluid against the meat ant if they are getting attacked by them; depending on how much is regurgitated, an ant will either walk away and clean itself, or become fatally affected by it.[97] Lizards such as the thorny dragon is a sit and wait predator that consumes meat ants,[98] but other lizards which eat Iridomyrmex ants usually reject this species.[99]

Life cycle and reproduction

Meat ant alates swarming for nuptial flight

Nuptial flight usually occurs during spring in October.[100] Reproductive females will only mate with a single male and begin establishing her own colony afterwards.[101] Nuptial flight occurs after rain, where the males would emerge from their nest first, followed by the virgin queens.[47] Groups of 20 to 40 females occur after the males have flown away. They would position themselves on top of the nest in order to heat themselves, and would all fly at the same time once they are warm. This process would happen multiple times unless the weather had changed, otherwise the queens would return to their nest.[47] Nuptial flight would continue for days until all virgin queens have withdrawn from the nest. Most of the time, a single queen will start her own colony and lay eggs that will take around 44 to 61 days to fully develop and emerge as adults,[102][103] but can also be founded through multiple queens cooperating with each other, adoption into an existing colony, or "budding" (also called "satelliting" or "fractionating"), where a subset of the colony including queens, workers and brood (eggs, larvae and pupae) leave the main colony for an alternative nest site.[103] Around 10% of queens will have at least another queen with them during colony foundation.[104] Many queens are killed during colony founding; major aspects include predation by birds and other ants, even those of the same species, due to the fact they attempt to establish their nests near large colonies.[100] However, some queens are successful with this, and sometimes they would even be attended and protected by neighbouring workers, and would also help the queen dig some chambers. Other indications of queen deaths include disease and starvation.[47] A queens ovaries may take four weeks to mature, and she will lay around 20 eggs that may develop into larvae in less than a month.[47] Workers have been observed laying eggs, presumably trophic eggs.[105][106]

A queen meat ant burrowing a hole after her nuptial flight

A mature nest of several years of age can hold up to 11,000 - 64,000 ants, while other estimates are around 300,000.[107][51] During this time, some nests are known to contain nearly 70,000 larvae, 64,000 workers, 20,000 males and over 1,000 virgin queens.[47] However, some colonies contain more virgin queens than males. The ratio of worker ants and number of larvae in colonies ranges from one for every two, or two for every one.[47]

Most colonies are monogyne, where a colony will only have a single queen,[108] but based on observations, nests can contain more than a single queen.[47] Some nests are known to contain two queens, with some even having as many as four in a single colony. Some colonies are oligogynous, which means that multiple queens are present in a colony, but they are tolerated by all workers birthed from different queens and treated equally.[103][109] Tolerance still occurred even when new reproductive females and males are born,[103] but recognition based on kin from queens and workers is known, hinting brood discrimination when larvae are fed or groomed;[110] queens would only take care of their own brood and neglect to look after brood laid by other queens.[111] The queens on the other hand will only cooperate with each other during nest founding, but will be antagonistic once there are workers present in the colony.[112] Queens will become more intolerant to each other as the colony grows, and eventually separate within the nest, resulting in the queen laying more eggs.[103] Such cases usually happen when pleometrotic founding occurs, or if a queen ant is adopted by a colony, setting up aggressive relationships.[103][113] Physical fights between queens in the same colony is rare.

Ritualised fighting

Two workers engaged in ritualised fighting

Meat ants are highly territorial and aggressive ants which establish firm borders around neighbouring colonies.[42] While the boundaries are not physical, worker ants maintain this by engaging in ritualised fighting with opponent ants, an interaction most colonies engage in. Fatalities are rare on both sides but sometimes it may cause an occasional injury to several workers. The meat ant has evolved over time where it will mainly engage in ritualised fighting to resolve territorial disputes with nearby or neighbouring colonies, as this enables territory to be contested between opposing colonies without them killing each other, costing many casualties on both sides.[42] Because of this, it is a method of avoiding casualties and promotes intercolony communication and assessment.[114] A drainage of the work force would occur if these boundaries acted as a conflict zone if ritualised fighting does not take place.[114] Lethal fighting will occur if the colony is under attack.

Encounters between workers lasts for 15 seconds.[42] Ritualised fight only occurs with two worker ants who come into contact with each other, but this will break if both ants are from the same colony, followed by them grooming themselves. After, they would later walk around until they make contact with another ant.[42] A meat ant detects a foreign worker by intense antennation and gaping of the mandibles, and will also stretch themselves upward to appear taller and larger, suggesting that meat ants will do this in a display of size-matching.[114] Workers will perform a behaviour known as "front leg boxing"; both workers will have their front legs sweeping up and down, where it would flex at the coxal joint in a paddling motion.[42] This paddling motion was both aimed at each other during this behaviour, going on for three to five seconds on average. From this point on, this would determine who is a "loser" and who is the "winner".

The ant who lost the ritual fight will lower its body, where it would lean sideways from the victorious ant. The victorious ant will remain raised upward and would reach down to the worker and open its mandibles wider, where it would grasp on the ants mandibles, and then tugs and shakes its head slightly for a brief moment of time.[42] However, ritual fights may continue if neither worker backs down, and would commence a side to side posture.[42] They would raise their bodies and circle each other and present their gasters directed to their opponent, and on some occasions, either one or both of the ants would kick outward using their legs at each other. Eventually, they will break contact and groom once appeasement has been reached and continue to search for another ant.[42]

Relationship with other organisms

Meat ants with leafhoppers. These insects excrete a sugary sap that is collected by the ants, which protect this valuable food resource.

Meat ants have been observed blocking banded sugar ant nesting holes with pebbles and soil to prevent them from leaving their nest during the early hours of the day. Banded sugar ants counter this by preventing meat ants from leaving their nest by blocking their nesting holes with debris, a behaviour known as nest-plugging.[45][115] If meat ant nests are encroached by trees or other shade, banded sugar ants may invade and take over the nest, since the health of the colony may deteriorate from overshadowing.[52] Members of an affected meat ant colony later move to a nearby satellite nest that is placed in a suitable area, while invading banded sugar ants fill nest galleries up with a black resinous material.[55]

Meat ants are generally intolerant of myrmecophiles living in their colonies, but Cyclotorna larvae[47] are known dwell in colonies.[116] While Sphallomorpha larvae sometimes prey on workers as discussed earlier, some are inquilines and live in the nest commensally. Unused or abandoned areas inside colonies are sometimes occupied by other species of ants and in some cases, termites.[47] Meat ants play an important role in seed dispersal. A meat ant colony is capable of dispersing 334,000 individual bellyache bush seeds per hectare, which shows a strong ant-seed relationship among the two.[117]

Relationship with humans

Meat ants are able to kill poisonous cane toads, an introduced pest, as the toxins that usually kill cane toad's predators do not affect the meat ants.[118] Due to this, scientists have considered using meat ants as a form of pest control to reduce the cane toad population.[119] One way of doing this is by establishing meat ant nests in habitats where cane toad numbers are high.[120] In rural Australia, meat ants are important to farmers as they place animal carcasses on their nests. In a matter of weeks, the entire carcass will be consumed and reduced to bones.[32]

As pests

Despite their beneficial importance to humans, meat ants are sometimes considered pests, due to these ants entering human houses occasionally to feed on food and soil disturbance. The nests may cause annoyance if nests are formed around gravel paths, tennis courts and other cleared spaces.[121][122] Although meat ants enter houses occasionally, they have adapted rather well to urbanisation and populations can flourish in urban areas. During the early days in Canberra, newly constructed suburbs provided new nest sites for meat ants. Unpaved streets, gravel paths and drive ways are among the many new sites meat ants could nest in, making them a commonly encountered ant that may pose as a considerable pest to many. The abundance of food supplies from parks, plantations and home gardens also attracted meat ants to urban areas.[48][53] They pose as a serious problem for citrus growers in eastern Australia by affecting the biological control of Hemiptera insects, specifically those who produce honeydew.[123] Meat ants cannot sting, but they can induce irritating bites and secrete a defensive fluid from the end of their abdomen.[31]

Many ways of eradicating meat ants have proven unsuccessful. The first investigations were made in the 1930s to control meat ant populations in urban areas by fumigation, using carbon disulfide and calcium cyanide. Although these methods successfully eradicated some nests, they were rapidly reoccupied by those living in satellite or rival nests. However, the reoccupation of these nests by incipient colonies (young colonies beginning to develop) does not occur. This is because colony founding is often difficult and rare when most areas are nested by mature colonies.[53] This behaviour has led to long term maintenance and repeated treatments to ensure the nests are not resettled. As well as that, not all nests could be affected and others could remain undetected from pest controllers.[124] Greaves (1973) notes that the poor penetration of insecticides into all nest galleries may be the reason why nests are reoccupied, but owing to the nest structure, the insecticides have to be poured into each individual hole as no gallery connects to one another. Missing a single gallery can mostly lead to the reoccupation of the nest site.[53] Greaves concludes that dieldrin is the most effective insecticide to control meat ants, capable of killing ants quickly and being the most long-lasting chemical used.

The difficulty of eradicating meat ants have led to further studies. Two studies between 1996 and 2002 studied the the effects of granular baits on meat ant colonies and some of the baits (specifically those containing hydramethylnon and fipronil) effectively reduced the number of workers foraging.[125][126] This was only effective when ten grams of bait was placed on citrus trees or onto mounds. The ant bait Amdro was used in a recent study to identify an effective method of eradicating meat ants, but results showed that the bait failed to reduce ant populations significantly. There was evidence that the amount of active mounds declined, but this effect was only temporary. This is due to colonies only having five grams of bait which was insufficient to eliminate further nests, but the effect may have been more dramatic if extra bait was used.[122]

See also

References

  1. ↑ Johnson, Norman F. (19 December 2007). "Iridomyrmex purpureus (Smith)". Hymenoptera Name Server version 1.5. Columbus, Ohio, USA: Ohio State University. Retrieved 1 April 2015.
  2. 1 2 3 4 5 6 Smith, F. (1858). Catalogue of hymenopterous insects in the collection of the British Museum part VI (PDF). London: British Museum. pp. 36–40.
  3. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 Shattuck, S.; Heterick, B.E. (2011). "Revision of the ant genus Iridomyrmex (Hymenoptera : Formicidae)" (PDF). Zootaxa 2845: 1–74. ISBN 978-1-86977-676-3. ISSN 1175-5334.
  4. ↑ Mayr, G. (1862). "Myrmecologische Studien" (PDF). Verhandlungen der Zoologisch-Botanischen Gesellschaft in Wien 12: 649–776. doi:10.5281/zenodo.25912.
  5. ↑ Mayr, G. (1863). "Formicidarum index synonymicus" (PDF). Verhandlungen der Zoologisch-Botanischen Gesellschaft in Wien 13: 385–460. doi:10.5281/zenodo.25913.
  6. ↑ André, E. (1896). "Fourmis nouvelles d'Asie et d'Australie" (PDF). Revue d'Entomologie (Caen) 15: 251–265. doi:10.5281/zenodo.27114.
  7. ↑ Forel, A.H. (1902). "Fourmis nouvelles d'Australie" (PDF). Revue Suisse de Zoologie 10: 405–548. doi:10.5281/ZENODO.14495.
  8. ↑ Bingham, C.T. (1903). The fauna of British India, including Ceylon and Burma. Hymenoptera, Vol. II. Ants and Cuckoo-wasps (PDF). London: Taylor and Francis. p. 297.
  9. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 Shattuck, S.O. (1993). "Revision of the Iridomyrmex purpureus species-group (Hymenoptera: Formicidae)". Invertebrate Systematics 7 (1): 113–149. doi:10.1071/IT9930113.
  10. 1 2 3 Lowne, B.T. (1865). "Contributions to the natural history of Australian ants" (PDF). The Entomologist 2: 275–280. doi:10.5281/zenodo.25931.
  11. ↑ Taylor, Robert W.; Brown, D.R.; Cardale, Josephine C. (1985). Hymenoptera, Formicoidea, Vespoidea and Sphecoidea. Zoological catalogue of Australia 2. Canberra: Australian Government Publishing Service. p. 102. ISBN 978-0-644-03922-2.
  12. ↑ Mayr, G. (1876). "Die australischen Formiciden" (PDF). Journal des Museum Goddefroy 12 (4): 56–115. doi:10.5281/zenodo.25857.
  13. ↑ "Specimen: CASENT0909519 Iridomyrmex purpureus". AntWeb. The California Academy of Sciences. Retrieved 21 January 2016.
  14. ↑ Halliday, R.B. (1978). "Esterase variation at three loci in meat ants.". The Journal of heredity 70 (1): 57–61. PMID 469224.
  15. ↑ Halliday, R. B. (1981). "Heterozygosity and Genetic Distance in Sibling Species of Meat Ants (Iridomyrmex purpureus Group)". Evolution 35 (2): 234–242. doi:10.2307/2407834.
  16. ↑ Greenslade, P.J.M. (1987). "Environment and Competition as Determinants of Local Geographical-Distribution of 5 Meat Ants, Iridomyrmex-purpureus and Allied Species (Hymenoptera, Formicidae)". Australian Journal of Zoology 35 (3): 259–273. doi:10.1071/ZO9870259.
  17. ↑ Bolton, B. (2014). "Iridomyrmex purpureus". AntCat. Retrieved 19 December 2015.
  18. ↑ Ward, P.S.; Brady, S.G; Fisher, B.L.; Schultz, T.R. (2010). "Phylogeny and biogeography of dolichoderine ants: effects of data partitioning and relict taxa on historical inference". Systematic Biology 59: 342–362. doi:10.1093/sysbio/syq012. PMID 20525640.
  19. 1 2 Shattuck, S.O.; Barnett, N. (2010). "Iridomyrmex Mayr, 1862". Ants Down Under. CSIRO Entomology. Archived from the original on 24 May 2015. Retrieved 23 May 2015.
  20. 1 2 3 4 5 6 Greaves, T. (1971). "The distribution of the three forms of the meat ant Iridomyrmex purpureus (Hymenoptera: Formicidae) in Australia". Australian Journal of Entomology 10 (1): 15–21. doi:10.1111/j.1440-6055.1971.tb00004.x.
  21. 1 2 Halliday, R.B. (1979). Genetic studies of meat ants (Iridomyrmex purpureus) (PDF) (Ph.D.). University of Adelaide. pp. 1–3.
  22. 1 2 Greenslade, P.J.M. (1974). "Distribution of two forms of the Meat ant, Iridomyrmex purpureus (Hymenoptera: Formicidae), in parts of South Australia". Australian Journal of Zoology 22 (4): 489–504. doi:10.1071/ZO9740489.
  23. ↑ Greenslade, P.J.M. (1974). "The identity of Iridomyrmex purpureus form viridiaeneus Viehmeyer (Hymenoptera: Formicidae)". Australian Journal of Entomology 13 (3): 247–248. doi:10.1111/j.1440-6055.1974.tb02181.x.
  24. ↑ Greenslade, P. J. M. (1970). "Observations on the inland variety (v. viridiaeneus Viehmeyer) of the meat ant Iridomyrmex purpureus (Frederick Smith) (Hymenoptera: Formicidae)". Australian Journal of Entomology 9 (3): 227–231. doi:10.1111/j.1440-6055.1970.tb00796.x.
  25. ↑ Halliday, R.B. (1975). "Electrophoretic variation of amylase in Meat ants, Iridomyrmex purpureus, and its taxonomic significance". Australian Journal of Zoology 23 (2): 271. doi:10.1071/ZO9750271.
  26. ↑ purpureus. Charlton T. Lewis and Charles Short. A Latin Dictionary on Perseus Project.
  27. ↑ Karkov, C.E.; Brown, G.H. (2012). Anglo-Saxon Styles. SUNY Press. p. 231. ISBN 978-0-7914-8614-6.
  28. ↑ AntWeb. "Genus: Iridomyrmex". The California Academy of Sciences. Retrieved 12 April 2015.
  29. ↑ Rosenthal, G.A.; Berenbaum, M.R. (2012). Herbivores: Their Interactions with Secondary Plant Metabolites: The Chemical Participants (2nd ed.). Academic Press. p. 299. ISBN 978-0-32-313940-3.
  30. ↑ Harper, Douglas. "Formica". Online Etymology Dictionary.
  31. 1 2 "Iridomyrmex purpureus (Smith, 1858): Greenslade's Meat Ant". Atlas of Living Australia. Government of Australia. Retrieved 18 December 2015.
  32. 1 2 3 4 5 6 Australian Museum. "Animal Species: Meat Ant". Retrieved 16 April 2015.
  33. ↑ Andersen, A.N. (2002). "Common names for Australian ants (Hymenoptera: Formicidae)" (PDF). Australian Journal of Entomology 41 (4): 285–293. doi:10.1046/j.1440-6055.2002.00312.x.
  34. ↑ Kastelic, P.; Koeshall, J. "Iridomyrmex purpureus: interactions". University of Wisconsin System. Retrieved 7 February 2016.
  35. ↑ The concise encyclopedia of Australia 2. Australia: Horowitz. 1979. p. 620.
  36. ↑ Thomas, M.L.; Parry, L.J.; Allan, R.A.; Elgar, M.A. (1999). "Geographic Affinity, Cuticular Hydrocarbons and Colony Recognition in the Australian Meat Ant Iridomyrmex purpureus". Naturwissenschaften 86 (2): 87–92. doi:10.1007/s001140050578.
  37. 1 2 3 Wheeler, G.C.; Wheeler, J. (1974). "Ant larvae of the subfamily Dolichoderinae: second supplement (Hymenoptera: Formicidae)". Pan-Pacific Entomologist 49: 396–401. doi:10.5281/zenodo.25094.
  38. ↑ "Formicidae Family". CSIRO Entomology. Retrieved 26 January 2016.
  39. 1 2 Gollan, J. R.; Ashcroft, M. B.; Ramp, D. (2013). "Fine-grained climate data alters the interpretation of a trait-based cline" (PDF). Ecosphere 4 (12): 154. doi:10.1890/ES13-00275.1.
  40. 1 2 Kastelic, P.; Koeshall, J. "Iridomyrmex purpureus: habitat". University of Wisconsin System. Retrieved 27 January 2016.
  41. 1 2 3 4 Shattuck, S.; Barnett, N. (2011). "Iridomyrmex purpureus (Smith, 1858)". Ants Down Under. CSIRO Entomology. Archived from the original on 16 April 2015. Retrieved 15 April 2015.
  42. 1 2 3 4 5 6 7 8 9 Ettershank, G.; Ettershank, J. A. (1982). "Ritualised fighting in the meat ant Iridomyrmex purpureus (Smith) (Hymenoptera: Formicidae)". Australian Journal of Entomology 21 (2): 97–102. doi:10.1111/j.1440-6055.1982.tb01772.x.
  43. ↑ "Southern meat ants - Iridomyrmex purpureus (Smith)". CSIRO Division of Entomology. 18 September 2004. Retrieved 27 January 2016.
  44. ↑ Andersen, A.N. (1995). "A Classification of Australian Ant Communities, Based on Functional Groups Which Parallel Plant Life-Forms in Relation to Stress and Disturbance". Journal of Biogeography 22 (1): 15–29. doi:10.2307/2846070. JSTOR 2846070.
  45. 1 2 Hölldobler & Wilson 1990, p. 424.
  46. 1 2 "Species: Iridomyrmex purpureus". AntWeb. The California Academy of Sciences. Retrieved 27 January 2016.
  47. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 Greaves, T.; Hughes, R. D. (1974). "The population biology of the meat ant". Australian Journal of Entomology 13 (4): 329–351. doi:10.1111/j.1440-6055.1974.tb02212.x.
  48. 1 2 "Ants and ant nests". Access Canberra. ACT Government. 2015. Retrieved 25 January 2016.
  49. 1 2 van Wilgenburg, E.; Ryan, D.; Morrison, P.; Marriott, P. J.; Elgar, M. A. (2006). "Nest- and colony-mate recognition in polydomous colonies of meat ants (Iridomyrmex purpureus)". Naturwissenschaften 93 (7): 309–314. Bibcode:2006NW.....93..309W. doi:10.1007/s00114-006-0109-y. ISSN 1432-1904. PMID 16555093.
  50. ↑ Greenslade, P. J. M.; Halliday, R. B. (1983). "Colony dispersion and relationships of meat ants Iridomyrmex purpureus and allies in an arid locality in south Australia". Insectes Sociaux 30 (1): 82–99. doi:10.1007/BF02225659. ISSN 1420-9098.
  51. 1 2 Greenslade, P.J.M. (1974). "Some relations of the meat ant, Iridomyrmex purpureus (hymenoptera: formicidae) with soil in South Australia". Soil Biology and Biochemistry 6 (1): 7–14. doi:10.1016/0038-0717(74)90004-2.
  52. 1 2 Cowan, J. A.; Humphreys, G. S.; Mitchell, P. B.; Murphy, C. L. (1985). "An assessment of pedoturbation by two species of mound-building ants, Camponotus intrepidus (Kirby) and Iridomyrmex purpureus (F. Smith)". Australian Journal of Soil Research 23 (1): 95. doi:10.1071/SR9850095.
  53. 1 2 3 4 Greaves, T. (1973). "Biological problems in the control of the meat ant, Iridomyrmex purpureus (Hymenoptera: Formicidae )". Australian Journal of Entomology 12 (4): 284–288. doi:10.1111/j.1440-6055.1973.tb01674.x.
  54. ↑ Duncan-Weatherley, A.H. (1953). "Some aspects of the biology of the mound ant Iridomyrmex detectus (Smith)". Australian Journal of Zoology 1 (2): 178–192. doi:10.1071/ZO9530178.
  55. 1 2 3 4 Ettershank, G. (1968). "The three dimensional gallery structure of the nest of the meat ant Iridomyrmex purpureus (SM.) (Hymenoptera : Formicidae)". Australian Journal of Zoology 16 (4): 715–723. doi:10.1071/ZO9680715.
  56. ↑ Greenslade, P.J.M. (1976). "The meat ant Iridomyrmex purpureus (Hymenoptera: Formicidae) as a dominant member of ant communities". Australian Journal of Entomology 15 (2): 237–240. doi:10.1111/j.1440-6055.1976.tb01700.x.
  57. 1 2 Andersen, A.N.; Patel, A.D. (1994). "Meat ants as dominant members of Australian ant communities: an experimental test of their influence on the foraging success and forager abundance of other species". Oecologia 98 (1): 15–24. doi:10.1007/BF00326085. ISSN 1432-1939.
  58. 1 2 Gibb, H.; Hochuli, D.F. (2004). "Removal experiments reveal limited effects of a dominant species on ant assemblages". Ecology 85 (3): 648–657. doi:10.1890/03-0007.
  59. 1 2 3 4 5 6 7 8 Gibb, H. (2005). "The effect of a dominant ant, Iridomyrmex purpureus, on resource use by ant assemblages depends on microhabitat and resource type". Austral Ecology 30 (8): 856–867. doi:10.1111/j.1442-9993.2005.01528.x.
  60. ↑ Andrews, E.A. (1927). "Ant-mounds as to temperature and sunshine". Journal of Morphology 44 (1): 1–20. doi:10.1002/jmor.1050440102.
  61. ↑ Hölldobler & Wilson 1990, p. 381.
  62. ↑ Andrew, N.R.; Hart, R.A.; Jung, M.P.; Hemmings, Z.; Terblanche, J.S. (2013). "Can temperate insects take the heat? A case study of the physiological and behavioural responses in a common ant, Iridomyrmex purpureus (Formicidae), with potential climate change". Journal of insect physiology 59 (9): 870–880. doi:10.1016/j.jinsphys.2013.06.003. PMID 23806604.
  63. ↑ Gibb, H.; Hochuli, D.F. (2003). "Colonisation by a dominant ant facilitated by anthropogenic disturbance: effects on ant assemblage composition, biomass and resource use". Oikos 103 (3): 469–478. doi:10.1034/j.1600-0706.2003.12652.x.
  64. ↑ Braby, M.F. (2004). The complete field guide to butterflies of Australia. Collingwood, Vic.: CSIRO Publishing. p. 246. ISBN 978-0-643-09027-9.
  65. 1 2 "Southern Meat Ant". Queensland Museum. Government of Queensland. Retrieved 7 February 2016.
  66. ↑ Sharman, M.; Williamson, I.; Ramsey, D.S.L. (1995). "Observations on the early life history stages of Notaden bennettii in the Chinchilla area of southern Queensland". Memoirs Of The Queensland Museum 38 (2): 667–669.
  67. ↑ Tillyard, R. J. (1922). "The Life-history of the Australian Moth-lacewing, Ithone fusca, Newman (Order Neuroptera Planipennia)". Bulletin of Entomological Research 13 (2): 205–223. doi:10.1017/S000748530002811X. ISSN 0007-4853. OCLC 1537749.
  68. ↑ Hölldobler & Wilson 1990, p. 518.
  69. ↑ Samson, P.R.; O'Brien, C.F. (1981). "predation on Ogyris genoveva (Lepidoptera: Lycaenidae) by meat ants". Australian Entomological Magazine 8 (2-3): 21.
  70. ↑ Good, N.E.; Cotton, R.T. (1937). Annotated List of the Insects and Mites Associated with Stored Grain and Cereal Products, and of Their Arthropod Parasites and Predators. U.S. Department of Agriculture. p. 44.
  71. ↑ Common 1990, p. 60.
  72. ↑ Kitching, R.L. (1999). Biology of Australian Butterflies 6. CSIRO Publishing. p. 140. ISBN 978-0-643-05027-3.
  73. ↑ Jones, R. E. (1987). "Ants, Parasitoids, and the Cabbage Butterfly Pieris rapae". The Journal of Animal Ecology 56 (3): 739–749. doi:10.2307/4945. JSTOR 4945.
  74. ↑ Palomo, G.; Martinetto, P.; Perez, C.; Iribarne, O. (2003). "Ant predation on intertidal polychaetes in a SW Atlantic estuary" (PDF). Marine Ecology Progress Series 253: 165–173. doi:10.3354/meps253165. Retrieved 15 April 2015.
  75. ↑ "The Australian Naturalist". Journal and Magazine of the Naturalists' Society of New South Wales (University of Minnesota) 1–2: 187. 1909.
  76. 1 2 Moulds, T. (2006). "The first Australian record of subterranean guano-collecting ants" (PDF). Helictite 39 (1): 3–4.
  77. 1 2 3 Clerke, R. B.; Williamson, I. (1992). "A note on the predation of Bufo marinus juveniles by the ant Iridomyrmex purpureus". Australian Zoologist 28 (1-4): 64–67. doi:10.7882/AZ.1992.015.
  78. ↑ Cabrera-Guzmán, E.; Crossland, M.R.; Shine, R. (2015). "Invasive cane toads as prey for native arthropod predators in tropical Australia". Herpetological Monographs 29 (1): 28–39. doi:10.1655/HERPMONOGRAPHS-D-13-00007.
  79. 1 2 Ettershank, G. (1978). "Acquisition of water from soil by the meat-ant, Iridomyrmex purpureus (Smith) (Hymenoptera: Formicidae)". Insectes Sociaux 25 (1): 31–38. doi:10.1007/BF02224483.
  80. 1 2 Abensperg-Traun, M.A. (1988). "Food preference of the echidna, Tachyglossus aculeatus (Monotremata: Tachyglossidae), in the wheatbelt of Western Australia". Australian Mammal Society 11: 117–123. ISSN 0310-0049.
  81. ↑ Griffiths, M.; Simpson, K.G. (1966). "A seasonal feeding habit of Spiny Ant-eaters". CSIRO Wildlife Research 11 (1): 137–143. doi:10.1071/CWR9660137.
  82. ↑ Augee, M.; Gooden, B.; Musser, A. (2006). Echidna: Extraordinary Egg-Laying Mammal. CSIRO Publishing. p. 97. ISBN 978-0-643-09885-5.
  83. 1 2 3 Griffiths, M. (2012). The Biology of the Monotremes. Elsevier. pp. 21, 84–85. ISBN 978-0-323-15331-7.
  84. ↑ "Stray feathers". The Emu: official organ of the Australasian Ornithologists' Union 19: 244–247. 1919. ISSN 0046-192X. OCLC 1567848.
  85. ↑ Lea, M.L. (1915). "Stomach contents of birds". Transactions of the Royal Society of South Australia 39: 760–766. LCCN 95641248. OCLC 7487493.
  86. ↑ Jacskon, S.W. (1912). "Haunts of the Spotted Bowerbird (Chlamydodera maculata, Gld.)". The Emu: official organ of the Australasian Ornithologists' Union 12 (2): 65–104. doi:10.1071/MU912065. ISSN 0046-192X. OCLC 1567848.
  87. ↑ Webb, J.K.; Shine, R. (1992). "To find an ant: trail-following in Australian blindsnakes (Typhlopidae)". Animal Behaviour 43 (6): 941–948. doi:10.1016/S0003-3472(06)80007-2.
  88. ↑ Drijfhout, F. (2010). "Chemical Ecology". eLS. doi:10.1002/9780470015902.a0003265.pub2.
  89. ↑ Herberstein, M.E. (2011). Spider Behaviour: Flexibility and Versatility. Cambridge University Press. p. 139. ISBN 978-1-139-49478-6.
  90. ↑ Capinera, J.L. (2008). Encyclopedia of Entomology 4. Springer Science & Business Media. p. 93. ISBN 978-1-4020-6242-1.
  91. ↑ Litwack, G. (2010). Pheromones 83. Academic Press. p. 227. ISBN 978-0-12-381533-0. ISSN 0083-6729.
  92. 1 2 3 Pierce, N.E. (1995). "Predatory and parasitic Lepidoptera: Carnivores living on plants". Journal of The Lepidopterists' Society 49: 412–453. ISSN 0024-0966.
  93. ↑ Common 1990, p. 308.
  94. ↑ Dodd, F.P. (1912). "Some remarkable ant-friend Lepidoptera". Transactions of the Entomological Society of London 1911: 577–589.
  95. ↑ Common 1990, p. 179.
  96. ↑ Moore, B.P. (1974). "The larval habits of two species of Sphallomorpha Westwood (Coleoptera: Carabidae: Pseudomorphinae)". Australian Journal of Entomology 13 (3): 179–183. doi:10.1111/j.1440-6055.1974.tb02171.x.
  97. ↑ Hairston, N.G. (1989). Ecological Experiments: Purpose, Design and Execution. Cambridge, Massachusetts: Cambridge University Press. p. 96. ISBN 978-0-521-34692-4.
  98. ↑ Withers, P.C; Dickman, C.R (1995). "The role of diet in determining water, energy and salt intake in the thorny devil Moloch horridus (Lacertilia: Agamidae)" (PDF). Journal of the Royal Society of Western Australia, 7 78 (3).
  99. ↑ Tyler, .M.J. (1960). "Observations on the diet and size variation of Amphibolurus adelaidensis (Gray) (Reptilia-Agamidae) on the Nullarbor Plain". Transactions of The Royal Society of South Australia 83: 111–117.
  100. 1 2 Hölldobler & Wilson 1990, p. 210.
  101. ↑ Hölldobler & Wilson 1990, p. 156.
  102. ↑ Hölldobler & Wilson 1990, p. 170.
  103. 1 2 3 4 5 6 Hölldobler, B.; Carlin, N.F. (1985). "Colony founding, queen dominance and oligogyny in the Australian meat ant Iridomyrmex purpureus". Behavioral Ecology and Sociobiology 18 (1): 45–58. doi:10.1007/BF00299237. ISSN 1432-0762. JSTOR 4599861.
  104. ↑ Hölldobler & Wilson 1990, p. 217.
  105. ↑ Trager 1988, p. 168.
  106. ↑ Trager 1988, p. 187.
  107. ↑ Hölldobler & Wilson 1990, p. 163.
  108. ↑ Trager 1988, p. 203.
  109. ↑ Michelucci, P. (2013). Handbook of Human Computation. New York, NY: Springer New York. p. 931. ISBN 978-1-4614-8806-4.
  110. ↑ Hölldobler & Wilson 1990, p. 202.
  111. ↑ Trager 1988, pp. 271–272.
  112. ↑ Hölldobler & Wilson 1990, p. 159.
  113. ↑ Hölldobler & Wilson 1990, p. 201.
  114. 1 2 3 Elgar, M.A.; Wilgenburg, E. van; Lieshout, E. van (2005). "Conflict resolution strategies in meat ants (Iridomyrmex purpureus): ritualised displays versus lethal fighting". Behaviour 142 (6): 701–716. doi:10.1163/1568539054729150. JSTOR 4536265.
  115. ↑ Gordon, D.M. (1988). "Nest-plugging: interference competition in desert ants (Novomessor cockerelli and Pogonomyrmex barbatus)". Oecologia 75 (1): 114–118. doi:10.1007/BF00378823. ISSN 1432-1939.
  116. ↑ Moore, B.P. (1964). "Australian larval carabidae of the subfamilies Broscinae, Poydrinae and Pseudomorphinae (Coleoptera)." (PDF). Pacific Insects 6 (2): 242–246.
  117. ↑ Lundgren, J.G. (2009). Relationships of natural enemies and non-prey foods. Dordrecht: Springer. p. 220. ISBN 978-1-4020-9235-0.
  118. ↑ Sweeney, Claire (31 March 2009). "Killer ants are weapons of mass toad destruction". Times Online. Archived from the original on 18 December 2015. Retrieved 31 March 2009.
  119. ↑ "Meat ant the new weapon against cane toads". The Sydney Morning Herald. 31 March 2009. Retrieved 18 December 2015.
  120. ↑ Grebner, D.L.; Bettinger, P.; Siry, J.P. (2013). Introduction to forestry and natural resources (1st ed.). Academic Press. p. 142. ISBN 978-0-12-386902-9.
  121. ↑ "The Farm Home. Control of Ants.". The Grenfell Record and Lachlan District Advertiser (NSW: National Library of Australia). 20 August 1934. p. 4. Retrieved 21 July 2015.
  122. 1 2 Webb, G.; Mayer, R.; Thomson, R. (2014). "Control of meat ants (Iridomyrmex sanguineus Forel) in a Western Australian sandalwood plantation using bait technology" (PDF). General and Applied Entomology 43 (1): 43–49. Archived from the original (PDF) on 6 February 2016.
  123. ↑ James, D. G.; Stevens, M. M.; O'Malley, K. J. (1998). "Prolonged exclusion of foraging ants (Hymenoptera: Formicidae) from citrus trees using controlled-release chlorpyrifos trunk bands". International Journal of Pest Management 44 (2): 65–69. doi:10.1080/096708798228338.
  124. ↑ Greaves, T. (1939). "The control of meat ants (Iridomyrmex detectus Sm.)". Journal of the Council for Scientific and Industrial Research 12: 109–114.
  125. ↑ James, D.G.; Stevens, M.M.; O'Malley, K.; Heffer, R. (1996). "Activity of a hydramethylnon-based bait against the citrus ant pests, Iridomyrmex rufoniger gp. spp. and I. purpureus". Plant Protection Quarterly 11 (3): 122–125.
  126. ↑ Stevens, M. M.; James, D. G.; Schiller, L. J. (2002). "Attractiveness of bait matrices and matrix/toxicant combinations to the citrus pests Iridomyrmex purpureus (F.Smith) and Iridomyrmex rufoniger gp sp. (Hym., Formicidae)". Journal of Applied Entomology 126 (9): 490–496. doi:10.1046/j.1439-0418.2002.00699.x.

Cited literature

External links

This article is issued from Wikipedia - version of the Sunday, February 14, 2016. The text is available under the Creative Commons Attribution/Share Alike but additional terms may apply for the media files.