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The yellow crazy ant (''Anoplolepis gracilipes''), also known as the long-legged ant or Maldive ant, is a species of
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22,0 ...
, thought to be native to
West Africa West Africa or Western Africa is the westernmost region of Africa. The United Nations geoscheme for Africa#Western Africa, United Nations defines Western Africa as the 16 countries of Benin, Burkina Faso, Cape Verde, The Gambia, Ghana, Guinea, ...
or
Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...
. They have been accidentally introduced to numerous places in the world's tropics. The yellow crazy ant has colloquially been given the modifier "crazy" on account of the ant's erratic movements when disturbed. Its long legs and antennae make it one of the largest invasive ant species in the world.''Anoplolepis gracilipes''.
Global Invasive Species Database. ISSG.
Like several other invasive ants, such as the red imported fire ant (''
Solenopsis invicta The red imported fire ant (''Solenopsis invicta''), also known as the fire ant or RIFA, is a species of ant native to South America. A member of the genus '' Solenopsis'' in the subfamily Myrmicinae, it was described by Swiss entomologist Feli ...
''), the big-headed ant (''
Pheidole megacephala ''Pheidole megacephala'' is a species of ant in the family Formicidae. It is commonly known as the big-headed ant in the USA and the coastal brown ant in Australia. It is a very successful invasive species and is considered a danger to native an ...
''), the little fire ant (''
Wasmannia auropunctata The little fire ant (''Wasmannia auropunctata''), also known as the electric ant, is a small (approx long), light to golden brown (ginger) social ant native to Central and South America, now spread to parts of Africa (including Gabon and Camero ...
''), and the Argentine ant ('' Linepithema humile''), the yellow crazy ant is a "tramp ant", a species that easily becomes established and dominant in new habitat due to traits such as
aggression Aggression is overt or covert, often harmful, social interaction with the intention of inflicting damage or other harm upon another individual; although it can be channeled into creative and practical outlets for some. It may occur either reacti ...
toward other ant species, little aggression toward members of its own species, efficient
recruitment Recruitment is the overall process of identifying, sourcing, screening, shortlisting, and interviewing candidates for jobs (either permanent or temporary) within an organization. Recruitment also is the processes involved in choosing individual ...
, and large
colony In modern parlance, a colony is a territory subject to a form of foreign rule. Though dominated by the foreign colonizers, colonies remain separate from the administration of the original country of the colonizers, the '' metropolitan state'' ...
size. It is on a list of "one hundred of the world's worst invasive species" formulated by the
International Union for Conservation of Nature The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of nat ...
(IUCN), having invaded ecosystems from
Hawaii Hawaii ( ; haw, Hawaii or ) is a U.S. state, state in the Western United States, Western United States, located in the Pacific Ocean about from the U.S. mainland. It is the only U.S. state outside North America, the only state that is ...
to the
Seychelles Seychelles (, ; ), officially the Republic of Seychelles (french: link=no, République des Seychelles; Creole: ''La Repiblik Sesel''), is an archipelagic state consisting of 115 islands in the Indian Ocean. Its capital and largest city, ...
, and formed supercolonies on
Christmas Island Christmas Island, officially the Territory of Christmas Island, is an states and territories of Australia, Australian external territory comprising the island of the same name. It is located in the Indian Ocean, around south of Java and Su ...
in the
Indian Ocean The Indian Ocean is the third-largest of the world's five oceanic divisions, covering or ~19.8% of the water on Earth's surface. It is bounded by Asia to the north, Africa to the west and Australia to the east. To the south it is bounded by t ...
.


Physiology

''Anoplolepis gracilipes'' is a relatively large, yellow to orange ant with long legs, large eyes and extremely long antennal scapes. Although ''A. gracilipes'' is the only invasive species in the genus ''Anoplolepis'', there are several other genera for which it can be mistaken. Both ''
Leptomyrmex A video on how Spider Ant colonies function) ''Leptomyrmex'', or spider ants, is a genus of ants and a distinctive member of the ant subfamily Dolichoderinae. Commonly known as "spider ants" for their long legs and spider-like movements, these oran ...
'' and '' Oecophylla'' can be confused with ''Anoplolepis'' because of their similar sizes and very long limbs. ''Anoplolepis'' can be distinguished from ''Leptomyrmex'' by the presence of an
acidopore The acidopore is a component of ant anatomy; a round orifice located on the abdomen encircled by hairs which typifies the subfamily Formicinae. The conical structure connects to the posterior end of an ant's gaster where formic acid and other hy ...
, while ''Anoplolepis'' can be distinguished from ''Oecophylla'' by the more compact petiole. Although both of these genera occur in the Pacific, neither contain any invasive species. Several species of invasive ants belonging to the genera '' Camponotus'' and '' Paratrechina'' can appear similar to ''A. gracilipes''. Although several invasive species of ''
Pheidole ''Pheidole'' is a genus of ants that belongs to the ant subfamily Myrmicinae. The genus is widespread and ecologically dominant. It probably includes more than 1000 species. The genus first evolved in the Americas, eventually spreading across th ...
'' can also be slender-bodied with long legs and long antennal scapes, they can be separated from ''A. gracilipes'' by their two-segmented waists. ''A. gracilipes'' is widespread across the tropics, and populations are especially dense in the Pacific region. The species is most infamous for causing the ecological "meltdown" of
Christmas Island Christmas Island, officially the Territory of Christmas Island, is an states and territories of Australia, Australian external territory comprising the island of the same name. It is located in the Indian Ocean, around south of Java and Su ...
. Although widespread across the Pacific, ''A. gracilipes'' can cause significant damage to native biological diversity. Strong
quarantine A quarantine is a restriction on the movement of people, animals and goods which is intended to prevent the spread of disease or pests. It is often used in connection to disease and illness, preventing the movement of those who may have been ...
measures are encouraged to keep it from spreading to new localities.


Geographical range and dispersal

The yellow crazy ant's natural habitats are the moist tropical lowlands of
Southeast Asia Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia or SEA, is the geographical United Nations geoscheme for Asia#South-eastern Asia, south-eastern region of Asia, consistin ...
, and surrounding areas and islands in the Indian and Pacific Oceans. It has been introduced into a wide range of tropical and subtropical environments including northern Australia, some of the Caribbean islands, some Indian Ocean islands (
Seychelles Seychelles (, ; ), officially the Republic of Seychelles (french: link=no, République des Seychelles; Creole: ''La Repiblik Sesel''), is an archipelagic state consisting of 115 islands in the Indian Ocean. Its capital and largest city, ...
,
Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Africa ...
,
Mauritius Mauritius ( ; french: Maurice, link=no ; mfe, label=Mauritian Creole, Moris ), officially the Republic of Mauritius, is an island nation in the Indian Ocean about off the southeast coast of the African continent, east of Madagascar. It incl ...
,
Réunion Réunion (; french: La Réunion, ; previously ''Île Bourbon''; rcf, label= Reunionese Creole, La Rényon) is an island in the Indian Ocean that is an overseas department and region of France. It is located approximately east of the island o ...
, the
Cocos Islands ) , anthem = "''Advance Australia Fair''" , song_type = , song = , image_map = Australia on the globe (Cocos (Keeling) Islands special) (Southeast Asia centered).svg , map_alt = Location of the Cocos (Keeling) Islands , map_caption = ...
and the Christmas Islands) and some Pacific islands (
New Caledonia ) , anthem = "" , image_map = New Caledonia on the globe (small islands magnified) (Polynesia centered).svg , map_alt = Location of New Caledonia , map_caption = Location of New Caledonia , mapsize = 290px , subdivision_type = Sovereign st ...
,
Hawaii Hawaii ( ; haw, Hawaii or ) is a U.S. state, state in the Western United States, Western United States, located in the Pacific Ocean about from the U.S. mainland. It is the only U.S. state outside North America, the only state that is ...
,
French Polynesia )Territorial motto: ( en, "Great Tahiti of the Golden Haze") , anthem = , song_type = Regional anthem , song = " Ia Ora 'O Tahiti Nui" , image_map = French Polynesia on the globe (French Polynesia centered).svg , map_alt = Location of Frenc ...
,
Okinawa is a Prefectures of Japan, prefecture of Japan. Okinawa Prefecture is the southernmost and westernmost prefecture of Japan, has a population of 1,457,162 (as of 2 February 2020) and a geographic area of 2,281 Square kilometre, km2 (880 sq mi). ...
,
Vanuatu Vanuatu ( or ; ), officially the Republic of Vanuatu (french: link=no, République de Vanuatu; bi, Ripablik blong Vanuatu), is an island country located in the South Pacific Ocean. The archipelago, which is of volcanic origin, is east of no ...
,
Micronesia Micronesia (, ) is a subregion of Oceania, consisting of about 2,000 small islands in the western Pacific Ocean. It has a close shared cultural history with three other island regions: the Philippines to the west, Polynesia to the east, an ...
,
Johnston Atoll Johnston Atoll is an unincorporated territory of the United States, currently administered by the United States Fish and Wildlife Service (USFWS). Johnston Atoll is a National Wildlife Refuge and part of the Pacific Remote Islands Marine Nationa ...
, and the Galapagos archipelago). The species has been known to occupy such agricultural systems as cinnamon, citrus, coffee and coconut plantations. Because yellow crazy ants have generalized nesting habits, they are able to disperse via trucks, boats and other forms of human transport. Crazy ant colonies naturally disperse through "budding", i.e. when mated queens and workers leave the nest to establish a new one, and only rarely through flight via female winged reproductive forms. Generally, colonies that disperse through budding have a lower rate of dispersal, requiring human intervention to reach distant areas. It has been recorded that ''A. gracilipes'' moves as much as a year in the
Seychelles Seychelles (, ; ), officially the Republic of Seychelles (french: link=no, République des Seychelles; Creole: ''La Repiblik Sesel''), is an archipelagic state consisting of 115 islands in the Indian Ocean. Its capital and largest city, ...
. A survey on
Christmas Island Christmas Island, officially the Territory of Christmas Island, is an states and territories of Australia, Australian external territory comprising the island of the same name. It is located in the Indian Ocean, around south of Java and Su ...
, however, yielded an average spreading speed of per day, the equivalent of one kilometer (0.6 mile) per year.


Diet

''A. gracilipes'' has been described as a "scavenging predator" exhibiting a broad diet, a characteristic of many invasive species. It consumes a wide variety of foods, including grains, seeds, arthropods, and decaying matter such as vertebrate corpses. They have been reported to attack and dismember invertebrates such as small
isopods Isopoda is an order of crustaceans that includes woodlice and their relatives. Isopods live in the sea, in fresh water, or on land. All have rigid, segmented exoskeletons, two pairs of antennae, seven pairs of jointed limbs on the thorax, an ...
,
myriapods Myriapods () are the members of subphylum Myriapoda, containing arthropods such as millipedes and centipedes. The group contains about 13,000 species, all of them terrestrial. The fossil record of myriapods reaches back into the late Silurian, a ...
,
molluscs Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000 extant species of molluscs are recognized. The number of fossil species is estim ...
,
arachnids Arachnida () is a class of joint-legged invertebrate animals (arthropods), in the subphylum Chelicerata. Arachnida includes, among others, spiders, scorpions, ticks, mites, pseudoscorpions, harvestmen, camel spiders, whip spiders and vinegaro ...
, land crabs,
earthworm An earthworm is a terrestrial invertebrate that belongs to the phylum Annelida. They exhibit a tube-within-a-tube body plan; they are externally segmented with corresponding internal segmentation; and they usually have setae on all segments. Th ...
s and insects. Like all ants, ''A. gracilipes'' requires a protein-rich food source for the queen to lay eggs and carbohydrates as energy for the workers. They get their carbohydrates from plant
nectar Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualis ...
and honeydew producing insects, especially
scale insect Scale insects are small insects of the order Hemiptera, suborder Sternorrhyncha. Of dramatically variable appearance and extreme sexual dimorphism, they comprise the infraorder Coccomorpha which is considered a more convenient grouping than the ...
s,
aphid Aphids are small sap-sucking insects and members of the superfamily Aphidoidea. Common names include greenfly and blackfly, although individuals within a species can vary widely in color. The group includes the fluffy white woolly aphids. A t ...
s, and other
Sternorrhyncha The Sternorrhyncha suborder of the Hemiptera contains the aphids, whiteflies, and scale insects, groups which were traditionally included in the now-obsolete order "Homoptera". "Sternorrhyncha" refers to the rearward position of the mouthparts rel ...
. Studies indicate that crazy ants rely so much on scale insects that a scarcity of them can actually limit ant population growth.


Mutualism

Crazy ants obtain much of their food requirements from
scale insects Scale insects are small insects of the order Hemiptera, suborder Sternorrhyncha. Of dramatically variable appearance and extreme sexual dimorphism, they comprise the infraorder Coccomorpha which is considered a more convenient grouping than th ...
, serious plant pests that feed on sap of trees and release honeydew, a sugary liquid. Ants eat honeydew, and in return protect the scale from their enemies and spread them among trees, an example of mutualism. The honeydew not eaten by the ants drips onto the trees and encourages the growth of sooty mold over the leaves and stems. This gives plants an ugly black appearance and reduces their health and vigor. The ants protect the insects by "nannying" the mobile crawler stages and protecting them against their natural enemies. Experiments have shown that this connection is so strong that, in environments where ''A. gracilipes'' was removed, the density of scale insects dropped by 67% within 11 weeks, and to zero after 12 months.


In Australia

In Australia, yellow crazy ants have been found at more than 30 sites in
Queensland ) , nickname = Sunshine State , image_map = Queensland in Australia.svg , map_caption = Location of Queensland in Australia , subdivision_type = Country , subdivision_name = Australia , established_title = Before federation , established_ ...
, and in
Arnhem Land Arnhem Land is a historical region of the Northern Territory of Australia, with the term still in use. It is located in the north-eastern corner of the territory and is around from the territory capital, Darwin. In 1623, Dutch East India Compan ...
in the
Northern Territory The Northern Territory (commonly abbreviated as NT; formally the Northern Territory of Australia) is an Australian territory in the central and central northern regions of Australia. The Northern Territory shares its borders with Western Aus ...
, where a large scattered population exists.Invasive Species Council; (2016
Yellow Crazy Ants In Australia
Invasive Species Council Fact Sheet.
A single
New South Wales ) , nickname = , image_map = New South Wales in Australia.svg , map_caption = Location of New South Wales in AustraliaCoordinates: , subdivision_type = Country , subdivision_name = Australia , established_title = Before federation , e ...
infestation was detected and eradicated, and, in
Western Australia Western Australia (commonly abbreviated as WA) is a state of Australia occupying the western percent of the land area of Australia excluding external territories. It is bounded by the Indian Ocean to the north and west, the Southern Ocean to t ...
, yellow crazy ants have been intercepted in shipping freight arriving at
Fremantle Fremantle () () is a port city in Western Australia, located at the mouth of the Swan River in the metropolitan area of Perth, the state capital. Fremantle Harbour serves as the port of Perth. The Western Australian vernacular diminutive for ...
.Csurhes, Steve and Hankamer, Clare; 2012 (updated 2016
"Invasive Animal Risk Assessment Yellow Crazy Ant"
Queensland Government of Australia. Retrieved 2017-03-17.
Queensland's main infestation is in and around
Wet Tropics of Queensland The Wet Tropics of Queensland World Heritage Site consists of approximately 8,940 km2 of Australian wet tropical forests growing along the north-east Queensland portion of the Great Dividing Range. The Wet Tropics of Queensland meets all f ...
rainforest Rainforests are characterized by a closed and continuous tree canopy, moisture-dependent vegetation, the presence of epiphytes and lianas and the absence of wildfire. Rainforest can be classified as tropical rainforest or temperate rainforest, ...
, a
World Heritage Site A World Heritage Site is a landmark or area with legal protection by an international convention administered by the United Nations Educational, Scientific and Cultural Organization (UNESCO). World Heritage Sites are designated by UNESCO for h ...
. The Northern Territory infestation covers , an area larger than the
Australian Capital Territory The Australian Capital Territory (commonly abbreviated as ACT), known as the Federal Capital Territory (FCT) until 1938, is a landlocked federal territory of Australia containing the national capital Canberra and some surrounding townships. I ...
. Climate modelling indicates yellow crazy ants could spread across northern Australia from Queensland to Western Australia, across much of Queensland and into coastal and inland parts of New South Wales. Areas with the most ideal habitat and climatic conditions, such as the Wet Tropics of Queensland rainforests, are likely to experience the highest impacts. A costbenefit analysis by the Queensland government undertaken in 2012 found that yellow crazy ants could cost Australia's economy over A$3 billion if the ants were not treated. This analysis did not take potential impacts on Australia's biodiversity into account. The known impacts of crazy ants in tropical rainforests overseas may provide useful insights into these impacts, bearing in mind that the most significant impacts are associated with relatively small islands, such as Christmas Island.


Impact on Christmas Island

Crazy ants have had a profound impact on the biodiversity of Christmas Island. The crazy ant has a significant destructive impact on the island's ecosystem, killing and displacing crabs on the forest floor. The supercolonies also devastate crab numbers migrating to the coast. This has seen a rapid depletion in the number of land crabs — killing up to 20 million of them — which are vital to Christmas Island's biodiversity; land crabs are a keystone species in the forest ecology: they dig burrows, turn over the soil, and fertilize it with their droppings. Seedlings that were previously eaten by crabs started to grow and, as a result, changed the structure of the forest. Weeds have spread into the rainforest because there are no crabs to control them. One of the most noticeable changes in the forest is the increased numbers of the stinging tree '' Dendrocnide peltata'', which now flourishes in many areas frequently visited by humans. The forest canopy also changed as the scale insects tended by yellow crazy ants multiplied and killed mature trees.
Christmas Island red crab The Christmas Island red crab (''Gecarcoidea natalis'') is a species of land crab that is endemic to Christmas Island and Cocos (Keeling) Islands in the Indian Ocean. Although restricted to a relatively small area, an estimated 43.7 million ad ...
s are completely wiped out in infested areas. Populations of other ground and canopy dwelling animals, such as reptiles and other leaf litter fauna, have also decreased. During crab migrations, many crabs move through areas infested with ants and are killed. Studies show that the ant has displaced an estimated 15–20 million crabs by occupying their burrows, killing and eating resident crabs, and using their burrows as nest sites. This factor has greatly depleted red crabs, and made their annual land migrations far more perilous. Although crazy ants do not bite or sting, they spray
formic acid Formic acid (), systematically named methanoic acid, is the simplest carboxylic acid, and has the chemical formula HCOOH and structure . It is an important intermediate in chemical synthesis and occurs naturally, most notably in some ants. E ...
as a defence mechanism and to subdue their prey. In areas of high ant density, the movement of a land crab disturbs the ants and, as a result, the ants instinctively spray formic acid as a form of defence. The high levels of formic acid at ground level eventually overwhelm the crabs, and they are usually blinded then eventually die from dehydration (while attempting to flush off the
formic acid Formic acid (), systematically named methanoic acid, is the simplest carboxylic acid, and has the chemical formula HCOOH and structure . It is an important intermediate in chemical synthesis and occurs naturally, most notably in some ants. E ...
) and exhaustion. As the dead crabs decay, the protein becomes available to the ants. Crazy ants kill fauna, but encourage scale insects. Increased densities of scale insects cause forest die back, and even the death of large forest trees. These changes create a cascade of negative impacts, including weed invasion, significantly altering the forest landscape.


Supercolonies

Christmas Island is a focal point for international control efforts. These supercolonies spread farther and cause more damage than single colonies, and they pose the single greatest known threat to the island's biodiversity. Staff from Christmas Island National Park have worked in recent years to keep ant numbers in check. With help from the Christmas Island Crazy Ant Scientific Advisory Panel and support from the Australian Government they are holding ground. Another supercolony nearly devastated the bird fauna of
Johnston Atoll Johnston Atoll is an unincorporated territory of the United States, currently administered by the United States Fish and Wildlife Service (USFWS). Johnston Atoll is a National Wildlife Refuge and part of the Pacific Remote Islands Marine Nationa ...
in the Pacific Ocean. The single massive colony was found to occupy nearly a quarter of the island, with up to 1,000 queens in a plot of land wide. The infestation is thought to have been eradicated.


Control measures

To reduce the impacts of crazy ants on red crabs and Christmas Island's ecosystems the Parks Australia carried out a major aerial baiting program in 2009, to follow up the first aerial baiting conducted in 2002. The first step was conducting an extensive island-wide survey to determine the exact locations of the supercolonies. For several months, staff traversed the island surveying over 900 sites. The result was a map of crazy ant supercolonies and red crab burrow densities, together with other biodiversity data. In September 2009, a helicopter was used to precisely bait crazy ant supercolonies, which covered of the island. A very low concentration of
fipronil Fipronil is a broad-spectrum insecticide that belongs to the phenylpyrazole chemical family. Fipronil disrupts the insect central nervous system by blocking the ligand-gated ion channel of the GABAA receptor and glutamate-gated chloride (GluCl ...
bait (0.1%) was used to control the ants. Monthly monitoring of these baited supercolony sites shows that crazy ant densities were reduced by 99%. Park staff placed a high emphasis on minimising non-target impact of baiting. Food lures were dropped from a helicopter to attract robber crabs away from areas that were about to be baited. This technique, combined with the low concentration fipronil bait, proved to be highly successful with extremely low numbers of robber crabs and no red crabs known to be killed by the baiting. While baiting has slowed the decline of the red crab, its effects on the crazy ant populations are only temporary, as escaping colonies invade the treated areas again, and it is expensive, requiring much man power. In an effort to find a better control, after research, Australian Parks in December 2016 imported '' Tachardiaephagus somervillei'', a small () wasp and began breeding them for release. The wasp, which attacks only scale insects, is a voracious predator of what is believed to be one of the crazy ant's largest source of honeydew on Christmas Island, the yellow lac scale insect. Researchers from
La Trobe University La Trobe University is a public university, public research university based in Melbourne, Victoria (Australia), Victoria, Australia. Its main campus is located in the suburb of Bundoora, Victoria, Bundoora. The university was established in 196 ...
in Melbourne, funded by Parks Australia, began looking for biological controls in 2009. While the ants are omnivores, studies have shown honeydew is an important part of the diet of Christmas Island crazy ants. Samples of ants taken from colonies that are growing rapidly have more honeydew in their diet than when the colonies decline. Further, restricting access to honeydew, by binding trees where the scale insects feed, dramatically reduced the colony as ant activity on the ground fell by 95% in just four weeks. In the laboratory, colonies with limited sources of sugar were compared to colonies with access to abundant sugar. Those with abundant sugar had more fertile queens and lower death rates among workers. The workers were also more aggressive toward other ant species and explored their environments more. This is believed to show why the ants decline when deprived of access to scale insects in the field, and confirm reduced honeydew will greatly reduce the ants' ability to form super colonies. While controlling the yellow lac scale insect is expected to control the yellow crazy ant on Christmas Island, on mainland Australia it is thought this would not help. There are at least a dozen honeydew producing insects as well as extrafloral nectar from native acacia trees, all of which fuel yellow crazy ants. Experts continue to call for a fully funded, long term baiting program on mainland Australia.


References


External links

* {{Taxonbar, from=Q935471 Fauna of Christmas Island Formicinae Insect pests of tropical forests Insects described in 1857 Insects of India Taxa named by Frederick Smith (entomologist)