Carpenter ant Temporal range: Eocene – Recent | |
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Camponotus cf. eugeniae (minor worker) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Hymenoptera |
Family: | Formicidae |
Subfamily: | Formicinae |
Tribe: | Camponotini |
Genus: | Camponotus Mayr, 1861 |
Type species | |
Formica ligniperda Latreille, 1802 | |
Diversity [1] | |
1,533 species | |
Synonyms [2] | |
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Carpenter ants (Camponotus spp.) are large ants (workers 7 to 13 mm or 1⁄4 to 1⁄2 in) indigenous to many forested parts of the world. [3]
They build nests inside wood, consisting of galleries chewed out with their mandibles or jaws, preferably in dead, damp wood. However, unlike termites, they do not consume wood, [4] but instead discard a material that resembles sawdust outside their nest. Sometimes, carpenter ants hollow out sections of trees. They also commonly infest wooden buildings and structures, causing a widespread problem: they are a major cause of structural damage. Nevertheless, their ability to excavate wood helps in forest decomposition. The genus includes over 1,000 species. [5] They also farm aphids. In their farming, the ants protect the aphids from predators (usually other insects) while they excrete a sugary fluid called honeydew, which the ants get by stroking the aphids with their antennae.
Carpenter ants are generally large ants: workers are 4–7 mm long in small species and 7–13 mm in large species, queens are 9–20 mm long and males are 5–13 mm long. The bases of the antennae are separated from the clypeal border by a distance of at least the antennal scape's maximum diameter. The mesosoma in profile usually forms a continuous curve from the pronotum through to the propodeum. [6] [7]
Carpenter ant species reside both outdoors and indoors in moist, decaying, or hollow wood, most commonly in forest environments. They cut "galleries" into the wood grain to provide passageways to allow for movement between different sections of the nest. Certain parts of a house, such as around and under windows, roof eaves, decks and porches, are more likely to be infested by carpenter ants because these areas are most vulnerable to moisture. [8]
Carpenter ants have been known to construct extensive underground tunneling systems. These systems often end at some food source – often aphid colonies, where the ants extract and feed on honeydew. These tunneling systems also often exist in trees. The colonies typically include a central "parent" colony surrounded and supplemented by smaller satellite colonies. [9]
Carpenter ants are considered both predators and scavengers. These ants are foragers that typically eat parts of other dead insects or substances derived from other insects. Common foods for them include insect parts, "honeydew" produced by aphids, and extrafloral nectar from plants. They are also known for eating other sugary liquids such as honey, syrup, or juices. Carpenter ants can increase the survivability of aphids when they tend them.
Most species of carpenter ants forage at night. When foraging, they usually collect and consume dead insects. Some species less commonly collect live insects. When they discover a dead insect, workers surround it and extract its body fluids to be carried back to the nest. The remaining chitin-based shell is left behind. Occasionally, the ants bring the chitinous head of the insect back to the nest, where they also extract its inner tissue. [10] The ants can forage individually or in small or large groups, though they often opt to do so individually. Different colonies in close proximity may have overlapping foraging regions, although they typically do not assist each other in foraging. Their main food sources normally include proteins and carbohydrates. [11] Instances of carpenter ants bleeding Chinese elm trees for the sap have been observed in northern Arizona. These instances may be rare, as the colonies vastly exceeded the typical size of carpenter ant colonies elsewhere. [12] When workers find food sources, they communicate this information to the rest of the nest. They use biochemical pheromones to mark the shortest path that can be taken from the nest to the source. When a sizable number of workers follows this trail, the strength of the cue increases and a foraging trail is established. This ends when the food source is depleted. The workers will then feed the queen and the larvae by consuming the food they have found, and regurgitating the food at the nest. Foraging trails can be either under or above ground. [13]
Although carpenter ants do not tend to be extremely aggressive, they have developed mechanisms to maximize what they take from a food source when that same food source is also visited by competing organisms. This is accomplished in different ways. Sometimes they colonize an area near a relatively static food supply. More often, they develop a systemic way to visit the food source, with alternating trips by different individual ants or groups. This allows them to decrease the gains of intruders because the intruders tend to visit in a scattered, random, and unorganized manner. The ants, however, visit the sources systematically so that they reduce the average crop remaining. They tend to visit more resource-dense food areas in an attempt to minimize resource availability for others. That is, the more systematic the foraging behavior of the ants, the more random that of its competitors. [14]
Contrary to popular belief, carpenter ants do not actually eat wood, as they are unable to digest cellulose. They only create tunnels and nests within it. [15]
Some carpenter ant species can obtain nitrogen by feeding on urine or urine-stained sand. This may be beneficial in nitrogen-limited environments. [16]
All ants in this genus, and some related genera, possess an obligate bacterial endosymbiont called Blochmannia . [17] This bacterium has a small genome, and retains genes to biosynthesize essential amino acids and other nutrients. This suggests the bacterium plays a role in ant nutrition. Many Camponotus species are also infected with Wolbachia , another endosymbiont that is widespread across insect groups. Wolbachia is associated with the nurse cells in the queen's ovaries in the species Camponotus textor , which results in the worker larva being infected. [18]
Carpenter ants work to build the nests that house eggs in environments with usually high humidity due to their sensitivity to environmental humidity. These nests are called primary nests. Satellite nests are constructed once the primary nest is established and has begun to mature. Residents of satellite nests include older larvae, pupae, and some winged individuals, such as male ants (drones), or future queen ants. Only eggs, the newly hatched larvae, workers, and the queen reside in the primary nests. As satellite nests do not have environmentally sensitive eggs, the ants can construct them in rather diverse locations that can actually be relatively dry. [19] Some species, like Camponotus vagus , build the nest in a dry place, usually in wood.
When conditions are warm and humid, winged males and females participate in a nuptial flight. They emerge from their satellite nests and females mate with a number of males while in flight. The males die after mating. These newly fertilized queens discard their wings and search for new areas to establish primary nests. The queens build new nests and deposit around 20 eggs, nurturing them as they grow until worker ants emerge. The worker ants eventually assist her in caring for the brood as she lays more eggs. After a few years, reproductive winged ants are born, allowing for the making of new colonies. Again, satellite nests will be established and the process will repeat itself. [19]
Relatedness is the probability that a gene in one individual is an identical copy, by descent, of a gene in another individual. It is essentially a measure of how closely related two individuals are with respect to a gene. It is quantified by the coefficient of relatedness, which is a number between zero and one. The larger the value, the more two individuals are "related". Carpenter ants are social hymenopteran insects. This means the relatedness between offspring and parents is disproportionate. Females are more closely related to their sisters than they are to their offspring. Between full sisters, the coefficient of relatedness is r > 0.75 (due to their haplodiploid genetic system). [20] Between parent and offspring, the coefficient of relatedness is r = 0.5, because, given the event in meiosis, a certain gene has a 50% chance of being passed on to the offspring.
Eusocial insects tend to present low genetic diversity within colonies, which can increase with the co-occurrence of multiple queens (polygyny) or with multiple mating by a single queen (polyandry). [21] Distinct reproductive strategies may generate similar patterns of genetic diversity in ants. [21]
According to Hamilton's rule for relatedness, for relative-specific interactions to occur, such as kin altruism, a high level of relatedness is necessary between two individuals. Carpenter ants, like many social insect species, have mechanisms by which individuals determine whether others are nestmates or not. They are useful because they explain the presence or absence of altruistic behavior between individuals. They also act as evolutionary strategies to help prevent incest and promote kin selection. [22] Social carpenter ants recognize their kin in many ways. These methods of recognition are largely chemical in nature, and include environmental odors, pheromones, "transferable labels", and labels from the queen that are distributed to and among nest members. [23] Because they have a chemical basis for emission and recognition, odors are useful because many ants can detect such changes in their environment through their antennae. [24]
The process of recognition for carpenter ants requires two events. First, a cue must be present on a "donor animal". These cues are called "labels". Next, the receiving animal must be able to recognize and process the cue. In order for an individual carpenter ant to be recognized as a nestmate, it must, as an adult, go through specific interactions with older members of the nest. [23] This process is also necessary in order for the ant to recognize and distinguish other individuals. If these interactions do not occur in the beginning of adult life, the ant will be unable to be distinguished as a nestmate and unable to distinguish nestmates. [25]
Recognition allows for the presence of kin-specific interactions, such as kin altruism. Altruistic individuals increase other individuals' fitness at the expense of their own. Carpenter ants perform altruistic actions toward their nestmates so that their shared genes are propagated more readily or more often. In many social insect species like these ants, many worker animals are sterile and do not have the ability to reproduce. As a result, they forgo reproduction to donate energy and help the fertile individuals reproduce. [26]
As in most other social insect species, individual interaction is heavily influenced by the queen. The queen can influence individuals with odors called pheromones, which can have different effects. Some pheromones have been known to calm workers, while others have been known to excite them. Pheromonal cues from ovipositing queens have a stronger effect on worker ants than those of virgin queens. [27]
In many social insect species, social behavior can increase the disease resistance of animals. This phenomenon, called social immunity, exists in carpenter ants. It is mediated through the feeding of other individuals by regurgitation. The regurgitate can have antimicrobial activity, which would be spread amongst members of the colony. Some proteases with antimicrobial activity have been found to exist in regurgitated material. Communal sharing of immune response capability is likely to play a large role in colonial maintenance during highly pathogenic periods. [28]
Polygyny often is associated with many social insect species, and usually is characterized by limited mating flights, small queen size, and other characteristics. However, carpenter ants have "extensive" mating flights and relatively large queens, distinguishing them from polygynous species. Carpenter ants are described as oligogynous because they have a number of fertile queens which are intolerant of each other and must therefore spread to different areas of the nest. Some aggressive interactions have been known to take place between queens, but not necessarily through workers. Queens become aggressive mainly to other queens if they trespass on a marked territory. Queens in a given colony can work together in brood care [9] and the workers tend to experience higher rates of survival in colonies with multiple queens. Some researchers still subscribe to the notion that carpenter ant colonies are only monogynous. [29]
In at least nine Southeast Asian species of the cylindricus complex, including Colobopsis saundersi , workers feature greatly enlarged mandibular glands that run the entire length of the ant's body. They can release their contents suicidally by performing autothysis, thereby rupturing the ant's body and spraying a toxic substance from the head, which gives these species the common name "exploding ants". [30] [31] [32] The enlarged mandibular gland, which is many times the size of that of a normal ant, produces a glue. The glue bursts out and entangles and immobilizes all nearby victims. [33] [34] However, all exploding ant species have been moved to Colobopsis based on recent taxonomy. [35]
The termite species Globitermes sulphureus has a similar defensive system. [36]
Camponotus currently has 43 subgenera. [1]
Carpenter ants can damage wood used in the construction of buildings. They can leave behind a sawdust-like material called frass that provides clues to their nesting location. Carpenter ant galleries are smooth and very different from termite-damaged areas, which have mud packed into the hollowed-out areas. Carpenter ants can be identified by the general presence of one upward protruding node, looking like a spike, at the "waist" attachment between the thorax and abdomen (petiole). [38] Control involves application of insecticides in various forms including dusts and liquids. The dusts are injected directly into galleries and voids where the carpenter ants are living. The liquids are applied in areas where foraging ants are likely to pick the material up and spread the poison to the colony upon returning. [39]
Carpenter ants and their larvae are eaten in various parts of the world. In Australia, the Honeypot ant ( Camponotus inflatus ) is regularly eaten raw by Indigenous Australians. [40] It is a particular favourite source of sugar for Australian Aborigines living in arid regions, partially digging up their nests instead of digging them up entirely, in order to preserve this food source. [41] [42] The honey also has antimicrobial properties which the aboriginal population use to their advantage to cure colds. [43]
In North America, lumbermen during the early years in Maine would eat carpenter ants to prevent scurvy, [44] and in John Muir's publication, First Summer in the Sierra , Muir notes that the Northern Paiute people of California ate the tickling, acid gasters of the large jet-black carpenter ants. [45] In Africa, carpenter ants are among a vast number of species that are consumed by the San people. [46]
Lasius is a genus of formicine ants. The type species for this genus is the black garden ant, Lasius niger. Other major members, which live in drier heathland, are the cornfield ant, L. neoniger, and L. alienus. Other species include the temporary social parasites of the L. mixtus group and the hyper-social parasite Lasius fuliginosus. Lasius flavus is also a commonly seen species, building grassy hillocks in undisturbed pasture. In the Alps, these mounds – always aligned east to catch the first rays of the rising sun – have been traditionally used by goatherds as natural compasses. Species in the subgenus Acanthomyops, in particular L. interjectus and L. claviger, are commonly known as citronella ants due to their citronella-like smell.
Ponerinae, the ponerine ants, is a subfamily of ants in the Poneromorph subfamilies group, with about 1,600 species in 47 extant genera, including Dinoponera gigantea - one of the world's largest species of ant. Mated workers have replaced the queen as the functional egg-layers in several species of ponerine ants. In such queenless species, the reproductive status of workers can only be determined through ovarian dissections.
Messor is a genus of myrmicine ants with more than 100 species, all of which are harvester ants; the generic name comes from the Roman god of crops and harvest, Messor. The subterranean colonies tend to be found in open fields and near roadsides, openings are directly to the surface.
Pogonomyrmex is a genus of harvester ants, occurring primarily in the deserts of North, Central, and South America, with a single endemic species from Haiti.
Dorylus, also known as driver ants, safari ants, or siafu, is a large genus of army ants found primarily in central and east Africa, although the range also extends to southern Africa and tropical Asia. The term siafu is a loanword from Swahili, and is one of numerous similar words from regional Bantu languages used by indigenous peoples to describe various species of these ants. Unlike the New World members of the former subfamily Ecitoninae, members of this genus form temporary subterranean bivouacs in underground cavities which they excavate and inhabit - either for a few days or up to three months. Also, unlike some New World army ants, driver ants are not specialized predators of other species of ant, instead being more generalistic with a diet consisting of a diversity of arthropods. Their colonies are enormous compared to other ant species, and can contain over 20 million individuals. As with their American counterparts, workers exhibit caste polymorphism with the soldiers having particularly large heads that power their scissor-like mandibles. They are capable of stinging, but very rarely do so, relying instead on their powerful shearing jaws. A large part of their diet consists of earthworms. Driver ant queens are the largest living ants known, with the largest measuring between 40 - 63 millimeters in total body length depending on their physiological condition.
The pharaoh ant is a small (2 mm) yellow or light brown, almost transparent ant notorious for being a major indoor nuisance pest, especially in hospitals. A cryptogenic species, it has now been introduced to virtually every area of the world, including Europe, the Americas, Australasia and Southeast Asia. It is a major pest in the United States, Australia, and Europe. The ant's common name is possibly derived from the mistaken belief that it was one of the Egyptian (pharaonic) plagues.
The banded sugar ant, also known as the sugar ant, is a species of ant native to Australia. A member of the genus Camponotus in the subfamily Formicinae, it was described by German entomologist Wilhelm Ferdinand Erichson in 1842. Its common name refers to the ant's liking for sugar and sweet food, as well as the distinctive orange-brown band that wraps around its gaster.
Iridomyrmex is a genus of ants called rainbow ants first described by Austrian entomologist Gustav Mayr in 1862. He placed the genus in the subfamily Dolichoderinae of the family Formicidae. It has 79 described species and five fossil species. Most of these ants are native to Australia; others are found in Asia and Oceania, and they have been introduced to Brazil, New Zealand, and the United Arab Emirates. Fossil species are known from China, France, and the United States.
The black carpenter ant is one of the largest and most common species of carpenter ant native to the central and eastern United States as well as eastern Canada.
Aphaenogaster is a genus of myrmicine ants. About 200 species have been described, including 18 fossil species. They occur worldwide except in South America south of Colombia, sub-Saharan Africa, and Antarctica.
Crematogaster is an ecologically diverse genus of ants found worldwide, which are characterised by a distinctive heart-shaped gaster (abdomen), which gives them one of their common names, the Saint Valentine ant. Members of this genus are also known as cocktail ants because of their habit of raising their abdomens when alarmed. Most species are arboreal (tree-dwelling). These ants are sometimes known as acrobat ants.
Acromyrmex is a genus of New World ants of the subfamily Myrmicinae. This genus is found in South America and parts of Central America, México and the Caribbean Islands, and contains 33 known species. Commonly known as "leafcutter ants" they comprise one of the two genera of advanced attines within the tribe Attini, along with Atta.
Camponotus vagus is a species of large, black, Palaearctic carpenter ant with a wide range that includes much of Europe, a large area of Asia, and part of Africa.
Tetraponera is a genus of ants in the subfamily Pseudomyrmecinae that are commonly known as slender ants and are characterized by their arboreal nature and slender bodies. The 96 described species of Tetraponera all of which live in hollow structures of plants and trees, such as thorns or branches; these hosts are known as myrmecophytes. Tetraponera species are closely related to the New World genus of ants Pseudomyrmex, but differ in their relationships with host plants.
Dolichoderus is a genus of ants found worldwide.
Neivamyrmex is a genus of army ants in the subfamily Dorylinae.
Nylanderia is a large genus of ants in the subfamily Formicinae. The genus has a nearly cosmopolitan distribution with species inhabiting a wide array of habitats in almost all geographic regions. Nylanderia, currently containing over 110 species, is an ecologically important genus, with some species reported as being invasive. The ants are small to medium in size and range in color from pale yellow to black.
Blochmannia is a genus of symbiotic bacteria found in carpenter ants and their allies in the tribe Camponotini. As of 2014, Blochmannia has been discovered in the guts of over 60 species across 6 genera within the Camponotini, and is predicted to be pervasive throughout the tribe. Blochmannia was first discovered by zoologist Friedrich Blochmann in 1887, who described "bacteria-like structures" in the ovaries and midgut of Camponotus ligniperdus in 1887. In 2000, Candidatus Blochmannia was proposed as its own genus.
Camponotus floridanus, or Florida carpenter ant, is a species of ant in the genus Camponotus. First described as Formica floridana by Buckley in 1866, the species was moved to Camponotus by Mayr in 1886. The ant is widespread in Florida and occurs as far north as North Carolina and as far west as Mississippi.
Camponotus fellah is a species of carpenter ant found across the Middle East and North Africa. This species was formally described by Dalla Torre in 1893. A C. fellah queen holds the record for Israeli ant longevity, surviving for 26 years (1983-2009) in a laboratory environment.