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Linepithema humile

What do they look like?

Argentine ants are a small species of ants, with one bump, called a petiole, on the skinny part between their abdomen and thorax. They also have a short, solid thorax, a flattened head, and thin legs. Unlike other ant species, all workers are about the same size, they are usually about 0.3 mm long and weigh about 0.43 mg. Males are about the same size or a little larger than workers, weighing about 1.0 mg. Sexual females, also known as queens, are much larger; they are about 4.5 to 5.0 mm long and weigh about 3.6 mg. All Argentine ants are dull brown, though males and queens are usually darker than workers. Their whole body is the same color, except small areas like their mandibles (appendages near their mouth) and antennae, which can have 3 different shades. Workers have yellowish mandibles and their legs are a lighter shade of brown than their body. Sexual females have yellow mandibles and reddish legs and antennae, while males have yellowish legs, antennae, and mandibles. Sexual females are also more opaque than worker, with more hairs on their body. Males have wings throughout their adult lifespan, while sexual females lose their wings after mating. (Markin, 1970; Newell and Barber, 1913; Passera and Aron, 1996; "Argentine ant — Linepithema humile", 2009)

Eggs are pearly white and elliptical in shape; they are usually 0.3 mm long by 0.2 mm wide and weigh about 0.02 mg. When they are close to hatching, the egg takes on a dull appearance. Newly hatched larvae are about 0.5 mm long, male larvae can grow to 2.5 mm long. Larvae are white and curved, straightening as they grow. They weigh 0.1 to 0.5 mg. Male and worker larvae are shiny and can be distinguished from queen larvae because queens are matte and opaque in comparison. Pupae are naked, but their features are visible. While there are three different color forms, all Argentine ants are white after pupation and transition through several shades of cream and brown until they reach a similar adult shade. Worker pupae are about 2.0 mm long; their head is about 50% of their length, with noticeably visible eyes. Male pupae are about 3.0 mm long, with a large thorax. Queen pupae are much larger than male or worker pupae, with prominent wing pads and proportionate body parts. (Markin, 1970; Newell and Barber, 1913; Passera and Aron, 1996; "Argentine ant — Linepithema humile", 2009)

  • Sexual Dimorphism
  • female larger
  • Average mass
    workers: 0.0043 g, males: 0.001 g, queens: 0.0036 g
    oz
  • Average length
    workers and males: 0.3 cm, queens: 4.5 to 5.0 cm
    in

Where do they live?

Argentine ants (Linepithema humile) are a major invasive species found across the globe. They are originally from South America and are still widespread across the region. Since the late 1800s, they have been spreading due to human activities, mostly to areas with a Mediterranean climate. They are now found on all continents except Antarctica and some Oceanic islands. Their range is restricted by temperature; they cannot usually survive in areas with an average daily temperature below 7 to 14 degrees Celsius. In the United States, they are found as far north as North Carolina and California; they are more common in coastal areas than in the interior parts the country. They are found across Europe, including the Iberian peninsula, France, and of course, the Mediterranean. They are also found in Japan, South Africa, New Zealand, and Australia. (Abril, et al., 2008; Aron, et al., 2001; Brightwell and Silverman, 2010; Brightwell, et al., 2010; Inoue, et al., 2013; Lach, 2013; McGrannachan and Lester, 2013)

What kind of habitat do they need?

Argentine ants are able to survive in a variety of habitats; they are only limited by temperature and water sources. They can survive in temperatures from -5 to 45 degrees Celsius, although they do not thrive in areas with an average daily temperature below 7 to 14 degrees Celsius. Water sources are also key to their survival, including natural sources such as rivers and man-made sources such as urban water runoff. Colonies nest in forests, agricultural fields, shrublands, fields, near rivers, and in other areas disturbed by human activity. Likewise, nests are also found in urban and suburban areas and these ants may be found in homes and other buildings. Nests are shallow, about 1 to 2 inches into the soil. Argentine ants also nest under wood and debris, in sandy soil, under rocks, in cracks in pavement, and in buildings. (Brightwell and Silverman, 2010; Enzmann, et al., 2012; Fitzgerald and Gordon, 2012; Keller, et al., 1989; McGrannachan and Lester, 2013; Rice and Silverman, 2013; "Argentine ant — Linepithema humile", 2009)

How do they grow?

Argentine ants have complete metamorphosis, which means they go through an egg, larvae, pupae, and adult stage. They lay their eggs from early spring to late fall. Their eggs usually hatch in about 28 days, although it can take anywhere from 12 to 55 days. Larvae grow quickly the first 5 days after hatching and develop in 11 to 61 days, 31 days on average. Workers pupate for about 20 days, males pupate for 19 to 28 days, and queens pupate for 2 to 4 weeks. They develop faster in warmer temperatures. Before they become adults, their legs, mouth parts, and antennae become prominent, worker ants help remove their pupal skin and straighten their body parts. This is known as the callow stage, at this time, newly molted ants look like all other adults except they are colorless and clumsy. After 48 to 72 hours, the ant is a fully functioning adult. At least 33 days are needed to develop into an adult, though 74 days seems average. The first batch of eggs that are laid in early spring develop into the breeding group, they are found in the nest by late spring or early summer. Eggs laid later in the season develop into workers. The type of ants produced depends on the presence of queens and the pheromones they produce. The population of worker ants peaks in late summer and fall and brood production decreases steadily in late fall. (Aron, et al., 2001; Keller, et al., 1989; Libbrecht, et al., 2011; Newell and Barber, 1913)

How do they reproduce?

Argentine ants mate in their nests during late spring and summer. Before mating, males may join in on nuptial flights, though queens stay in their nest. Males may return to their original nest or join a new nest. When males join new nests, the workers from that nest may be aggressive toward them, however, when winged females are present; workers are much less aggressive toward new males. Mating takes place a few days after hatching. After about 6 minutes, females bite the males to end mating. In rare cases, female bites can cause the males to die. Males may fight over females or disturb other mating pairs, which likely decreases the amount of sperm transferred. Mating pairs may move to new locations to avoid these disruptive males. Females are fertilized by only one male, although they may mate with several males. Males mate with several females, but they may use all of their sperm in one breeding event. Males die shortly after mating, while queens do not lay eggs until the following spring, if they survive until then. Workers are sterile and do not mate. Nuptial flights help prevent inbreeding, but Argentine ants also mate in the nest. However, because there are many different queens in a nest, they produce genetically different offspring and inbreeding does not occur. (Aron, et al., 2001; Keller and Passera, 1992; Libbrecht, et al., 2011; Passera and Aron, 1996; Passera and Keller, 1994)

Nests of Argentine ants have multiple queens. After mating, queens stay in the nest and do not lay eggs until the following spring. However, 90% of queens are killed by workers before they lay their eggs. Workers attack queens at night on foraging trails, the same way they attack prey. Workers grab the queens' legs while others attack the body. Pieces of the queens' body either remain on the ground or workers carry them as they would prey. It is not known why the workers kill the queens, although it likely frees up food and it may play a role in the gender of their offspring, as the presence of the queens and their pheromones have an effect. Queens that survive lay their first batch of eggs in the spring and continue laying throughout the summer. Queens store sperm from their first mating for the rest of their lives and typically have more sperm stored than they could ever use. They can lay up to 50 or 60 eggs per day, with an average of 20 to 30 eggs. The rate of egg laying is effected by temperature, 28 degrees Celsius is optimal. It is also affected by the number of queens in the colony. A colony can have hundreds of queens. The more queens present, the fewer eggs laid by each queen. With more queens, worker care per queen also decreases. Less food is brought to each queen, which decreases their fitness and egg laying rate. Because queens secrete pheromones that attract workers, more queens mean more pheromones, which become muddled and less distinct, attracting fewer workers to any one specific queen. (Abril, et al., 2008; Keller and Passera, 1992; Keller, et al., 1989; Markin, 1970; Newell and Barber, 1913)

  • How often does reproduction occur?
    Argentine ants may mate several times, but usually only transfer sperm once.
  • Breeding season
    Mating takes place in the late spring and early summer.
  • Average time to independence
    74 days

Argentine ants give significant care to the queens' offspring. Queens themselves supply nutrients in the eggs, but do not give any other care. Workers carry off the eggs as soon as they are laid and constantly care for them, moving them continuously, possibly to regulate humidity. Sometimes eggs and larvae are kept together, sometimes they are separated. Eggs that are not tended by workers do not hatch, so their care is very important. Larvae are fed constantly by workers through mouth to mouth regurgitation. Workers also groom and transport larvae. When the nest is in danger, workers pick up the immature individuals and move them to safer locations. Workers help male pupae remove pupal skin when they are molting into adulthood, they also help other pupae molt by straightening their legs and antennae. In the 48 to 72 hours before their exoskeleton hardens, newly molted ants are in the callow stage and they are unstable and wobbly. Workers still aid them if necessary, or if the nest is in danger, workers pick up callow ants and move them. After their exoskeleton hardens, ants join the colony and care stops. Workers also help determine the adult role of larvae. Workers underfeed female larvae throughout most of the year, causing them to develop into workers as adults. In the spring, workers feed female larvae more, causing them to develop into queens. This change in feed is triggered by a pheromone produced by the queens. (Libbrecht, et al., 2011; Newell and Barber, 1913; Passera, et al., 1995)

  • Parental Investment
  • pre-hatching/birth
    • provisioning
      • female
    • protecting
      • female
  • pre-weaning/fledging
    • provisioning
      • female
    • protecting
      • female
  • pre-independence
    • provisioning
      • female
    • protecting
      • female

How long do they live?

Males typically live a few days to a month or two after reaching adulthood, and usually die shortly after mating. Those that participate in nuptial flights usually live longer. Most queens are executed by workers in the spring at the age of 10 months. Queens that aren't executed can live for over a year, likely several years. Workers live about 10 to 12 months. (Keller, et al., 1989; Newell and Barber, 1913; Passera and Keller, 1994)

  • Range lifespan
    Status: wild
    1+ (high) years
  • Typical lifespan
    Status: wild
    10 to 12 months

How do they behave?

Argentine ants are successful as an invasive species due to their behavior. These ants find food sources much faster than many native species and are able to find food in mazes very quickly. They decrease their foraging time by creating short paths to their food. This allows them to consume food sources before native ant species find them. These ants are territorial and they are aggressive towards non-colony members. Argentine ants are aggressive predators, groups of ants attack larger prey; one group of ants will hold the prey's legs, while another group attacks the prey's body. Unlike many ant species, workers of this species have no castes or divisions of labor. All workers contribute in foraging, brood care, and other tasks. Foraging occurs on trails, typically during the day, mostly in the morning. They also perform some nighttime activities, such as queen executions, which occur in the spring. New nests are formed by budding, where one or several females leave their home nest with a group of workers and form a new nest. Argentine ants are semi-nomadic, they move to new nests when they need to, such as during floods, dry soil, or other disturbances. They may also move seasonally, due to changes in temperature. While they are usually found in Mediterranean climates, colonies can survive in areas with colder winters by nesting in the top few inches of soil or in piles of decaying matter. During cold temperatures, workers are sluggish, foraging stops, and development times for eggs and larvae are significantly longer. (Brightwell, et al., 2010; Enzmann, et al., 2012; Keller and Passera, 1992; Markin, 1970; McGrannachan and Lester, 2013; Newell and Barber, 1913; Torres, et al., 2007)

Argentine ants live in colonies. In their native range of South America, these ants form large colonies that extend from one meter to hundreds of meters. In their introduced range, they form supercolonies, which include many nests over an area of 1,000 to 4,000 km. Ants from other nests in these supercolonies are all genetically similar and are able to move between nests without encountering any aggression. This sort of cooperation over such a large area is part of the reason why Argentine ants are such a successful invasive species. Argentine ants have actually formed a global supercolony across Europe, North America, Australia, and Japan. This supercolony likely formed because these ants were unintentionally transported by humans. (Inoue, et al., 2013; Torres, et al., 2007)

Home Range

Argentine ants form large colonies, ants move freely from nest to nest, without aggression from other colony members. Their home range is only limited to how far they can travel and the location of their nests. Ants likely move between nests separated by several meters, though colonies can extend for thousands of meters. (Passera and Keller, 1994)

How do they communicate with each other?

Argentine ants form giant supercolonies, so recognizing other ants from their colony is important. Nest mates know each other by a shared colony odor on the cuticle of their exoskeleton. By identifying nest mates, ants know when to attack and show aggression. The hydrocarbon signature in their introduced range is similar between several continents, indicating a global supercolony. When Argentine ants are brought together from different continents, they do not show aggression towards each other and identify each other as nest mates. Chemicals on their cuticle also signal when an ant has died. The disappearance of some chemicals after death alerts other ants in the nest, who collect the dead ants and move them to a waste pile. Living ants can detect a dead ant in less than an hour after death. (Abril, et al., 2008; Choe, et al., 2009; Inoue, et al., 2013; Newell and Barber, 1913; Passera, et al., 1995; Reid, et al., 2012; Torres, et al., 2007)

Ants use touch to communicate. Workers often groom each other, keeping their body, mandibles, and antennae clean. Touch is also important for sensing their environment, as Argentine ants often do not notice objects unless their antennae or other body part comes into contact. Their vision might not be strong, although they can detect light. Pheromones are also important for communication. While foraging, Argentine ants lay down pheromones to create trails that other ants follow. Since pheromones evaporate quickly, foraging ants make u-turns to reinforce the trails. By creating such a strong trail, large groups of Argentine ants are able to quickly find the food source. This has helped Argentine ants beat native ants to food sources and deplete them. Queens produce pheromones to attract workers to provide care and bring food. Another pheromone emitted by queens helps to determine the gender of their offspring. This pheromone causes workers to overfeed or underfeed larvae. When there are many queens, workers underfeed female larvae to produce other workers. After queens are executed in the spring, there are fewer pheromones and workers increase feedings, causing breeding ants to develop. (Abril, et al., 2008; Choe, et al., 2009; Inoue, et al., 2013; Newell and Barber, 1913; Passera, et al., 1995; Reid, et al., 2012; Torres, et al., 2007)

What do they eat?

Argentine ants are omnivorous. These ants prey on many different insect species. They also eat nectar, bird's eggs, and dead arthropods and other carrion. A major part of their diet comes from honeydew farmed from aphids and scale insects. The diet of a colony can change over time. New colonies tend to eat protein-rich insect prey, while more established super colonies primarily eat carbohydrate-rich honeydew. This may be because it takes time to establish long-term relationships with honeydew producing insects. Argentine ants also eat any available human food, particularly sweets. There is also evidence that colonies cannibalize eggs and larvae. (Brightwell and Silverman, 2010; Lach, 2013; Newell and Barber, 1913; Shik and Silverman, 2013)

  • Animal Foods
  • eggs
  • carrion
  • insects
  • terrestrial non-insect arthropods
  • Plant Foods
  • nectar

What eats them and how do they avoid being eaten?

Argentine ants are highly aggressive towards other ant species and potential predators. They also use chemicals to defend themselves. They are preyed on by many spider species, including Zodarion cesari in the Mediterranean. Larval antlions are also known predators. Antlions catch ants that fall into their pitfall traps. Other larger insects, such as cockroaches, also feed on these ants. Larger animals, such as amphibians and reptiles, including Japanese tree frogs, as well as several bird species, including northern flickers and house sparrows also prey on Argentine ants. (Buczkowski and Bennett, 2008; Glenn and Holway, 2008; Ito, et al., 2009; Monzo, et al., 2013; Newell and Barber, 1913; Torres, et al., 2007)

What roles do they have in the ecosystem?

As one of the most invasive ant species in the world, Argentine ants have a huge impact on the ecosystems they invade. As their range expands, Argentine ants displace native ant species and other insect populations. Argentine ants out-compete native species because they find food sources faster, forage longer, send many individuals to the food sources, and live in a wide range of habitats. Their ability to form supercolonies also allows for large scale cooperation within the species. By displacing native ant species, Argentine ants disrupt many ant-plant seed dispersal relationships, and do not appear to disperse seeds themselves. Pollinators may also be displaced by the ants, causing problems for native plants. By changing the insect community, Argentine ants also affect other parts of the community. Eliminating native insect prey may cause a decrease in bird populations and nesting habits. Likewise, displacing native species also affects reptile, amphibian, and mammal species. One example is the decreasing population size of coastal horned lizards in California, likely due to Argentine ants out-competing native ant species. Asian needle ants, another invasive ant species, are displacing Argentine ants from their invasive range in the eastern United States. (Abril, et al., 2008; Brightwell and Silverman, 2010; Buczkowski and Bennett, 2008; Inoue, et al., 2013; Lach, 2013; McGrannachan and Lester, 2013; Rice and Silverman, 2013; Rodriguez-Cabal, et al., 2012; Suarez and Case, 2002; Suarez, et al., 2005)

Argentine ants form mutualistic relationships with honeydew-producing insects, such as aphids and coccids. The ants tend the insects and eat the honeydew they produce; in exchange they protect these populations from predators and parasitoids. Since honeydew is an important part of their diet, it is critical that Argentine ants form these relationships. By tending and protecting honeydew producers, these ants allow pest populations to flourish, decreasing the survival of the plants on which the pests live. These ants have mutualisms with scale insects, including terrapin scales, Mediterranean black scales, California red scales, and Virginia pine scales, many species of aphids including melon aphids and oleander aphids, and mealybugs including obscure mealybugs, grape mealybugs, vine mealybugs, citrus mealybugs, and citrophilus mealybugs. Argentine ants also farm honeydew produced by gall wasp larvae. Phorid flies are parasitoids that lay their eggs on or inside Argentine ants. The larvae eat the ants' tissues, eventually killing the ant. Parasitic Wolbachia bacteria are common in native populations of Argentine ants, though infections are less common in introduced populations. (Brightwell and Silverman, 2010; Brightwell, et al., 2010; Bristow, 1991; Inouye and Agrawal, 2004; Markin, 1970; Mgocheki and Addison, 2009; Newell and Barber, 1913; Orr and Seike, 1998; Powell and Silverman, 2010; Reuter, et al., 2005)

Species (or larger taxonomic groups) that are mutualists with this species
Commensal or parasitic species (or larger taxonomic groups) that use this species as a host

Do they cause problems?

Argentine ants are one of the most invasive ant species in the world; they have been distributed worldwide by human activities and travel. In the areas they invade, they impact crops, human households, and native species. Their invasion has resulted in a loss of biodiversity, as they out-compete native ant species, causing negative effects on many other animal populations. While not directly damaging crops, their ability to form relationships with aphids and other honeydew producing insects allow these crop pests to flourish, causing damage to crop yields. Argentine ants also cause infrastructure damage when they invade buildings. In addition to being a nuisance, these ants can transfer diseases such as Escherichia coli, Enterococcus, Streptococcus, and Staphylococcus. Researchers have searched for ways to manage and end ant infestations. Due to their large numbers and supercolony structure, it is very difficult to completely remove these ants from any one area. If one nest is removed, ants from other nearby nests will likely re-colonize the area. Although, removing or re-locating water sources can help keep ants out of buildings and homes. Argentine ants are limited by cold temperatures, but due to global climate change and warming temperatures, their range could expand even further. (Abril, et al., 2008; Brightwell, et al., 2010; Enzmann, et al., 2012; Lowe, et al., 2000; dos Santos, et al., 2009)

  • Ways that these animals might be a problem for humans
  • injures humans
    • carries human disease
  • household pest

How do they interact with us?

There are no known positive effects of Argentine ants on humans.

Are they endangered?

As a major invasive species; Argentine ants have no special conservation status. Instead, they are one of the 100 most invasive alien species in the world according to the Invasive Species Specialist Group (ISSG). (Lowe, et al., 2000)

Some more information...

Argentine ants (Linepithema humile) were previously known as Linepithema humile. According to a government publication from 1913, there are reported cases of human infants covered in these swarming ants, which reportedly even caused several infant deaths, but these accounts were not verified and were likely sensationalized, though the ability of Argentine ants to recruit large numbers is well documented. There is a massive amount of literature and research available concerning their impact on the ecosystems they invade, the mutualisms they establish, their supercolony structure, the damage they can do to human populations, and ways to halt their expansion and curb any further ecosystem disturbances. (Newell and Barber, 1913; Passera and Keller, 1994)

Contributors

Angela Miner (author), Animal Diversity Web Staff, Leila Siciliano Martina (editor), Animal Diversity Web Staff.

References

University of California Agriculture and Natural Resources. 2009. "Argentine ant — Linepithema humile" (On-line). UC - IPM online. Accessed September 07, 2013 at http://www.ipm.ucdavis.edu/TOOLS/ANTKEY/argentine.html.

Abril, S., J. Oliveras, C. Gomez. 2008. Effect of temperature on the oviposition rate of Argentine ant queens (Linepithema humile Mayr) under monogynous and polygynous experimental conditions. Journal of Insect Physiology, 54/1: 265-272.

Aron, S., L. Keller, L. Passera. 2001. Role of resource availability on sex, caste and reproductive allocation ratios in the Argentine ant Linepithema humile. Journal of Animal Ecology, 70/5: 831-839.

Brightwell, R., P. Labadie, J. Silverman. 2010. Northward Expansion of the Invasive Linepithema humile (Hymenoptera: Formicidae) in the Eastern United States is Constrained by Winter Soil Temperatures. Environmental Entomology, 39/5: 1659-1665.

Brightwell, R., J. Silverman. 2010. Invasive Argentine ants reduce fitness of red maple via a mutualism with an endemic coccid. Biological Invasion, 12/7: 2051-2057.

Bristow, C. 1991. Are ant-aphid associations a tritrophic interaction - oleander aphids and Argentine ants. Oecologia, 87/4: 514-521.

Buczkowski, G., G. Bennett. 2008. Aggressive interactions between the introduced Argentine ant, Linepithema humile and the native odorous house ant, Tapinoma sessile. Biological Invasions, 10/7: 1001-1011.

Choe, D., J. Millar, M. Rust. 2009. Chemical signals associated with life inhibit necrophoresis in Argentine ants. Proceedings of the National Academy of Sciences of the United States of America, 106/20: 8251-8255.

Enzmann, B., K. Kapheim, T. Wang, P. Nonacs. 2012. Giving them what they want: manipulating Argentine ant activity patterns with water. Journal of Applied Entomology, 136/8: 588-595.

Fitzgerald, K., D. Gordon. 2012. Effects of Vegetation Cover, Presence of a Native Ant Species, and Human Disturbance on Colonization by Argentine Ants. Conservation Biology, 26/3: 525-538.

Glenn, S., D. Holway. 2008. Consumption of introduced prey by native predators: Argentine ants and pit-building ant lions. Biological Invasions, 10: 273-280.

Inoue, M., E. Sunamura, E. Suhr, F. Ito, S. Tatsuki, K. Goka. 2013. Recent range expansion of the Argentine ant in Japan. Diversity and Distributions, 19/1: 29-37.

Inouye, B., A. Agrawal. 2004. Ant mutualists alter the composition and attack rate of the parasitoid community for the gall wasp Disholcaspis eldoradensis (Cynipidae). Ecological Entomology, 29/6: 692-696.

Ito, F., M. Okaue, T. Ichikawa. 2009. A note on prey composition of the Japanese treefrog, Hyla japonica, in an area invaded by Argentine ants, Linepithema humile, in Hiroshima Prefecture, western Japan (Hymenoptera: Formicidae). Myrmecological News, 12: 35-39.

Keller, L., L. Passera. 1992. Mating System, Optimal Number of Matings, and Sperm Transfer in the Argentine Ant Linepithema humile. Behavioral Ecology and Sociobiology, 31/5: 359-366.

Keller, L., L. Passera, J. Suzzoni. 1989. Queen execution in the Argentine Ant, Linepithema humile. Physiological Entomology, 14/2: 157-163.

Lach, L. 2013. A comparison of floral resource exploitation by native and invasive Argentine ants. Arthropod-Plant Interactions, 7/2: 177-190.

Libbrecht, R., T. Schwander, L. Keller. 2011. Genetic components to caste allocation in a multiple-queen ant species. Evolution, 65/10: 2907-2915.

Lowe, S., M. Browne, S. Boudjelas. 2000. "100 of the World's Worst Invasive Alien Species" (On-line pdf). IUCN/SSC Invasive Species Specialist Group (ISSG). Accessed August 31, 2013 at http://www.issg.org/pdf/publications/worst_100/english_100_worst.pdf.

Markin, G. 1970. Seasonal life cycle of Argentine ant, Linepithema humile (Hymenoptera - Formicidae), in southern California. Annals of the Entomological Society of America, 63/5: 1238-1242.

McGrannachan, C., P. Lester. 2013. Temperature and starvation effects on food exploitation by Argentine ants and native ants in New Zealand. Journal of Applied Entomology, 137/7: 550-559.

Mgocheki, N., P. Addison. 2009. Interference of ants (Hymenoptera: Formicidae) with biological control of the vine mealybug Planococcus ficus (Signoret) (Hemiptera: Pseudococcidae). Biological Control, 49/2: 180-185.

Monzo, C., M. Juan-Blasco, S. Pekar, O. Molla, P. Castanera, A. Urbaneja. 2013. Pre-adaptive shift of a native predator (Araneae, Zodariidae) to an abundant invasive ant species (Hymenoptera, Formicidae). Biological Invasions, 15/1: 91-96.

Newell, W., T. Barber. 1913. The Argentine Ant. Washington D.C.: U.S. Department of Agriculture - Bureau of Entomology.

Orr, M., S. Seike. 1998. Parasitoids deter foraging by Argentine ants (Linepithema humile) in their native habitat in Brazil. Oecologia, 117/3: 420-425.

Passera, L., S. Aron. 1996. Early Sex Discrimination and Male Brood Elimination by Workers of the Argentine Ant. Proceedings of the Royal Society B-Biological Sciences, 263: 1041-1046.

Passera, L., S. Aron, D. Bach. 1995. Elimination of sexual brood in the Argentine ant Linepithema humile - queen effect and brood recognition. Entomologia Experimentalis et Applicata, 75/3: 203-212.

Passera, L., L. Keller. 1994. Mate availability and male dispersal in the Argentine ant Linepithema humile (Mayr) (Linepithema humile). Animal Behaviour, 48: 361-369.

Powell, B., J. Silverman. 2010. Impact of Linepithema humile and Tapinoma sessile (Hymenoptera: Formicidae) on three natural enemies of Aphis gossypii (Hemiptera: Aphididae). Biological Control, 54/3: 285-291.

Reid, C., T. Latty, M. Beekman. 2012. Making a trail: informed Argentine ants lead colony to the best food by U-turning coupled with enhanced pheromone laying. Animal Behaviour, 84/6: 1579-1587.

Reuter, M., J. Pederson, L. Keller. 2005. Loss of Wolbachia infection during colonization in the invasive Argentine ant Linepithema humile. Heredity, 94/3: 364-369.

Rice, E., J. Silverman. 2013. Propagule Pressure and Climate Contribute to the Displacement of Linepithema humile by Pachycondyla chinensis. PLOS ONE, 8/2: e56281. Accessed August 31, 2013 at http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0056281.

Rodriguez-Cabal, M., K. Stuble, B. Guenard, R. Dunn, N. Sanders. 2012. Disruption of ant-seed dispersal mutualisms by the invasive Asian needle ant (Pachycondyla chinensis). Biological Invasions, 14/3: 557-565.

Shik, J., J. Silverman. 2013. Towards a nutritional ecology of invasive establishment: aphid mutualists provide better fuel for incipient Argentine ant colonies than insect prey. Biological Invasions, 15/4: 829-836.

Suarez, A., T. Case. 2002. Bottom-up effects on persistence of a specialist predator: Ant invasions and horned lizards. Ecological Applications, 12/1: 291-298.

Suarez, A., P. Yeh, T. Case. 2005. Impacts of Argentine ants on avian nesting success. Insectes Sociaux, 52/4: 378-382.

Torres, C., M. Brandt, N. Tsutsui. 2007. The role of cuticular hydrocarbons as chemical cues for nestmate recognition in the invasive Argentine ant (Linepithema humile). Insectes Sociaux, 54/4: 363-373.

dos Santos, P., A. Fonseca, M. Sanches. 2009. Ants (Hymenoptera: Formicidae) as vectors for bacteria in two hospitals in the municipality of Divinopolis, State of Minas Gerais. Revista da Sociedade Brasileira de Medicina Tropical, 42/5: 565-569.

 
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