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This article is about social insects. For other uses, see Termite (disambiguation).

Termites are eusocial insects that are classified at the taxonomic rank of infraorder Isoptera, or alternatively as epifamily Termitoidae, within the order Blattodea (along with cockroaches). Termites were once classified in a separate order from cockroaches, but recent phylogenetic studies indicate that they evolved from cockroaches, as they are deeply nested within the group, and the sister group to wood eating cockroaches of the genus Cryptocercus. Previous estimates suggested the divergence took place during the Jurassic or Triassic. More recent estimates suggest that they have an origin during the Late Jurassic, with the first fossil records in the Early Cretaceous. About 3,106 species are currently described, with a few hundred more left to be described. Although these insects are often called "white ants", they are not ants, and are not closely related to ants.

Termite
Temporal range:Early Cretaceous–Recent
Formosan subterranean termite (Coptotermes formosanus)
Soldiers (red-coloured heads)
Workers (pale-coloured heads)
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Cohort: Polyneoptera
Superorder: Dictyoptera
Order: Blattodea
Infraorder: Isoptera
Brullé, 1832
Families

Cratomastotermitidae
Mastotermitidae
Termopsidae
Archotermopsidae
Hodotermitidae
Stolotermitidae
Kalotermitidae
Archeorhinotermitidae
Stylotermitidae
Rhinotermitidae
Serritermitidae
Termitidae

Like ants and some bees and wasps from the separate order Hymenoptera, termites divide as "workers" and "soldiers" that are usually sterile. All colonies have fertile males called "kings" and one or more fertile females called "queens". Termites mostly feed on dead plant material and cellulose, generally in the form of wood, leaf litter, soil, or animal dung. Termites are major detritivores, particularly in the subtropical and tropical regions, and their recycling of wood and plant matter is of considerable ecological importance.

Termites are among the most successful groups of insects on Earth, colonising most landmasses except Antarctica. Their colonies range in size from a few hundred individuals to enormous societies with several million individuals. Termite queens have the longest known lifespan of any insect, with some queens reportedly living up to 30 to 50 years. Unlike ants, which undergo a complete metamorphosis, each individual termite goes through an incomplete metamorphosis that proceeds through egg, nymph, and adult stages. Colonies are described as superorganisms because the termites form part of a self-regulating entity: the colony itself.

Termites are a delicacy in the diet of some human cultures and are used in many traditional medicines. Several hundred species are economically significant as pests that can cause serious damage to buildings, crops, or plantation forests. Some species, such as the West Indian drywood termite (Cryptotermes brevis), are regarded as invasive species.

Contents

The infraorder name Isoptera is derived from the Greek words iso (equal) and ptera (winged), which refers to the nearly equal size of the fore and hind wings. "Termite" derives from the Latin and Late Latin word termes ("woodworm, white ant"), altered by the influence of Latin terere ("to rub, wear, erode") from the earlier word tarmes. A termite nest is also known as a termitary or termitarium (plural termitaria or termitariums). In earlier English, termites were known as "wood ants" or "white ants". The modern term was first used in 1781.

The external appearance of the giant northern termite Mastotermes darwiniensis is suggestive of the close relationship between termites and cockroaches.

Termites were formerly placed in the order Isoptera. As early as 1934 suggestions were made that they were closely related to wood-eating cockroaches (genus Cryptocercus, the woodroach) based on the similarity of their symbiotic gut flagellates. In the 1960s additional evidence supporting that hypothesis emerged when F. A. McKittrick noted similar morphological characteristics between some termites and Cryptocercus nymphs. In 2008 DNA analysis from 16S rRNA sequences supported the position of termites being nested within the evolutionary tree containing the order Blattodea, which included the cockroaches. The cockroach genus Cryptocercus shares the strongest phylogenetical similarity with termites and is considered to be a sister-group to termites. Termites and Cryptocercus share similar morphological and social features: for example, most cockroaches do not exhibit social characteristics, but Cryptocercus takes care of its young and exhibits other social behaviour such as trophallaxis and allogrooming. Termites are thought to be the descendants of the genus Cryptocercus. Some researchers have suggested a more conservative measure of retaining the termites as the Termitoidae, an epifamily within the cockroach order, which preserves the classification of termites at family level and below. Termites have long been accepted to be closely related to cockroaches and mantids, and they are classified in the same superorder (Dictyoptera).

The oldest unambiguous termite fossils date to the early Cretaceous, but given the diversity of Cretaceous termites and early fossil records showing mutualism between microorganisms and these insects, they possibly originated earlier in the Jurassic or Triassic. Possible evidence of a Jurassic origin is the assumption that the extinct Fruitafossor consumed termites, judging from its morphological similarity to modern termite-eating mammals. The oldest termite nest discovered is believed to be from the Upper Cretaceous in West Texas, where the oldest known faecal pellets were also discovered. Claims that termites emerged earlier have faced controversy. For example, F. M. Weesner indicated that the Mastotermitidae termites may go back to the Late Permian, 251 million years ago, and fossil wings that have a close resemblance to the wings of Mastotermes of the Mastotermitidae, the most primitive living termite, have been discovered in the Permian layers in Kansas. It is even possible that the first termites emerged during the Carboniferous. The folded wings of the fossil wood roach Pycnoblattina, arranged in a convex pattern between segments 1a and 2a, resemble those seen in Mastotermes, the only living insect with the same pattern. Krishna et al., though, consider that all of the Paleozoic and Triassic insects tentatively classified as termites are in fact unrelated to termites and should be excluded from the Isoptera. Other studies suggest that the origin of termites is more recent, having diverged from Cryptocercus sometime during the Early Cretaceous.

Macro image of a worker.

The primitive giant northern termite (Mastotermes darwiniensis) exhibits numerous cockroach-like characteristics that are not shared with other termites, such as laying its eggs in rafts and having anal lobes on the wings. It has been proposed that the Isoptera and Cryptocercidae be grouped in the clade "Xylophagodea". Termites are sometimes called "white ants" but the only resemblance to the ants is due to their sociality which is due to convergent evolution with termites being the first social insects to evolve a caste system more than 100 million years ago. Termite genomes are generally relatively large compared to those of other insects; the first fully sequenced termite genome, of Zootermopsis nevadensis, which was published in the journal Nature Communications, consists of roughly 500Mb, while two subsequently published genomes, Macrotermes natalensis and Cryptotermes secundus, are considerably larger at around 1.3Gb.

External phylogeny

Internal phylogeny

As of 2013, about 3,106 living and fossil termite species are recognised, classified in 12 families; reproductive and/or soldier castes are usually required for identification. The infraorder Isoptera is divided into the following clade and family groups, showing the subfamilies in their respective classification:

Basal termite families

Infraorder Isoptera (= Epifamily Termitoidae)
Family Cratomastotermitidae
Family Mastotermitidae
Parvorder Euisoptera
Family Arceotermitidae
Family Archotermopsidae
Family Hodotermitidae
Family Kalotermitidae
Family Krishnatermitidae
Family Melqartitermitidae
Family Mylacrotermitidae
Family Stolotermitidae
Family Tanytermitidae
Family Termopsidae

Neoisoptera

The Neoisoptera, literally meaning "newer termites" (in an evolutionary sense), are a recently coined nanorder that include families commonly referred-to as "higher termites", although some authorities only apply this term to the largest family Termitidae. The latter characteristically do not have Pseudergate nymphs (many "lower termite" worker nymphs have the capacity to develop into reproductive castes: see below). Cellulose digestion in "higher termites" has co-evolved with eukaryotic gut microbiota and many genera have symbiotic relationships with fungi such as Termitomyces; in contrast, "lower termites" typically have flagellates and prokaryotes in their hindguts. Five families are now included here:

Termites are found on all continents except Antarctica. The diversity of termite species is low in North America and Europe (10 species known in Europe and 50 in North America), but is high in South America, where over 400 species are known. Of the 3,000 termite species currently classified, 1,000 are found in Africa, where mounds are extremely abundant in certain regions. Approximately 1.1 million active termite mounds can be found in the northern Kruger National Park alone. In Asia, there are 435 species of termites, which are mainly distributed in China. Within China, termite species are restricted to mild tropical and subtropical habitats south of the Yangtze River. In Australia, all ecological groups of termites (dampwood, drywood, subterranean) are endemic to the country, with over 360 classified species. Because termites are highly social and abundant, they represent a disproportionate amount of the world's insect biomass. Termites and ants comprise about 1% of insect species, but represent more than 50% of insect biomass.

Due to their soft cuticles, termites do not inhabit cool or cold habitats. There are three ecological groups of termites: dampwood, drywood and subterranean. Dampwood termites are found only in coniferous forests, and drywood termites are found in hardwood forests; subterranean termites live in widely diverse areas. One species in the drywood group is the West Indian drywood termite (Cryptotermes brevis), which is an invasive species in Australia.

Diversity of Isoptera by continent:
Asia Africa North America South America Europe Australia
Estimated number of species 435 1,000 50 400 10 360
Close-up view of a worker's head

Termites are usually small, measuring between 4 to 15 millimetres (316 to916 in) in length. The largest of all extant termites are the queens of the species Macrotermes bellicosus, measuring up to over 10 centimetres (4 in) in length. Another giant termite, the extinct Gyatermes styriensis, flourished in Austria during the Miocene and had a wingspan of 76 millimetres (3 in) and a body length of 25 millimetres (1 in).

Most worker and soldier termites are completely blind as they do not have a pair of eyes. However, some species, such as Hodotermes mossambicus, have compound eyes which they use for orientation and to distinguish sunlight from moonlight. The alates (winged males and females) have eyes along with lateral ocelli. Lateral ocelli, however, are not found in all termites, absent in the families Hodotermitidae, Termopsidae, and Archotermopsidae. Like other insects, termites have a small tongue-shaped labrum and a clypeus; the clypeus is divided into a postclypeus and anteclypeus. Termite antennae have a number of functions such as the sensing of touch, taste, odours (including pheromones), heat and vibration. The three basic segments of a termite antenna include a scape, a pedicel (typically shorter than the scape), and the flagellum (all segments beyond the scape and pedicel). The mouth parts contain a maxillae, a labium, and a set of mandibles. The maxillae and labium have palps that help termites sense food and handling.

Consistent with all insects, the anatomy of the termite thorax consists of three segments: the prothorax, the mesothorax and the metathorax. Each segment contains a pair of legs. On alates, the wings are located at the mesothorax and metathorax, which is consistent with all four-winged insects. The mesothorax and metathorax have well-developed exoskeletal plates; the prothorax has smaller plates.

Diagram showing a wing, along with the clypeus and leg

Termites have a ten-segmented abdomen with two plates, the tergites and the sternites. The tenth abdominal segment has a pair of short cerci. There are ten tergites, of which nine are wide and one is elongated. The reproductive organs are similar to those in cockroaches but are more simplified. For example, the intromittent organ is not present in male alates, and the sperm is either immotile or aflagellate. However, Mastotermitidae termites have multiflagellate sperm with limited motility. The genitals in females are also simplified. Unlike in other termites, Mastotermitidae females have an ovipositor, a feature strikingly similar to that in female cockroaches.

The non-reproductive castes of termites are wingless and rely exclusively on their six legs for locomotion. The alates fly only for a brief amount of time, so they also rely on their legs. The appearance of the legs is similar in each caste, but the soldiers have larger and heavier legs. The structure of the legs is consistent with other insects: the parts of a leg include a coxa, trochanter, femur, tibia and the tarsus. The number of tibial spurs on an individual's leg varies. Some species of termite have an arolium, located between the claws, which is present in species that climb on smooth surfaces but is absent in most termites.

Unlike in ants, the hind-wings and fore-wings are of equal length. Most of the time, the alates are poor flyers; their technique is to launch themselves in the air and fly in a random direction. Studies show that in comparison to larger termites, smaller termites cannot fly long distances. When a termite is in flight, its wings remain at a right angle, and when the termite is at rest, its wings remain parallel to the body.

Caste system

Caste system of termites
A – King
B – Queen
C – Secondary queen
D – Tertiary queen
E – Soldiers
F – Worker

Worker termites undertake the most labour within the colony, being responsible for foraging, food storage, and brood and nest maintenance. Workers are tasked with the digestion of cellulose in food and are thus the most likely caste to be found in infested wood. The process of worker termites feeding other nestmates is known as trophallaxis. Trophallaxis is an effective nutritional tactic to convert and recycle nitrogenous components. It frees the parents from feeding all but the first generation of offspring, allowing for the group to grow much larger and ensuring that the necessary gut symbionts are transferred from one generation to another. Some termite species may rely on nymphs to perform work without differentiating as a separate caste. Workers may be male or female and are usually sterile, especially in termites that have a nest site that is separate from their foraging site. Sterile workers are sometimes termed as true workers while those that are fertile, as in the wood-nesting Archotermopsidae, are termed as false workers.

The soldier caste has anatomical and behavioural specialisations, and their sole purpose is to defend the colony. Many soldiers have large heads with highly modified powerful jaws so enlarged they cannot feed themselves. Instead, like juveniles, they are fed by workers. Fontanelles, simple holes in the forehead that exude defensive secretions, are a feature of the family Rhinotermitidae. Many species are readily identified using the characteristics of the soldiers' larger and darker head and large mandibles. Among certain termites, soldiers may use their globular (phragmotic) heads to block their narrow tunnels. Different sorts of soldiers include minor and major soldiers, and nasutes, which have a horn-like nozzle frontal projection (a nasus). These unique soldiers are able to spray noxious, sticky secretions containing diterpenes at their enemies. Nitrogen fixation plays an important role in nasute nutrition. Soldiers are usually sterile but some species of Archotermopsidae are known to have neotenic forms with soldier-like heads while also having sexual organs.

The reproductive caste of a mature colony includes a fertile female and male, known as the queen and king. The queen of the colony is responsible for egg production for the colony. Unlike in ants, the king mates with her for life. In some species, the abdomen of the queen swells up dramatically to increase fecundity, a characteristic known as physogastrism. Depending on the species, the queen starts producing reproductive alates at a certain time of the year, and huge swarms emerge from the colony when nuptial flight begins. These swarms attract a wide variety of predators.

A young termite nymph. Nymphs first moult into workers, but others may further moult to become soldiers or alates.
Termite, and shed wings from other termites, on an interior window sill. Shedding of wings is associated with reproductive swarming.

Termites are often compared with the social Hymenoptera (ants and various species of bees and wasps), but their differing evolutionary origins result in major differences in life cycle. In the eusocial Hymenoptera, the workers are exclusively female. Males (drones) are haploid and develop from unfertilised eggs, while females (both workers and the queen) are diploid and develop from fertilised eggs. In contrast, worker termites, which constitute the majority in a colony, are diploid individuals of both sexes and develop from fertilised eggs. Depending on species, male and female workers may have different roles in a termite colony.

The life cycle of a termite begins with an egg, but is different from that of a bee or ant in that it goes through a developmental process called incomplete metamorphosis, with egg, nymph and adult stages. Nymphs resemble small adults, and go through a series of moults as they grow. In some species, eggs go through four moulting stages and nymphs go through three. Nymphs first moult into workers, and then some workers go through further moulting and become soldiers or alates; workers become alates only by moulting into alate nymphs.

The development of nymphs into adults can take months; the time period depends on food availability, temperature, and the general population of the colony. Since nymphs are unable to feed themselves, workers must feed them, but workers also take part in the social life of the colony and have certain other tasks to accomplish such as foraging, building or maintaining the nest or tending to the queen. Pheromones regulate the caste system in termite colonies, preventing all but a very few of the termites from becoming fertile queens.

Queens of the eusocial termite Reticulitermes speratus are capable of a long lifespan without sacrificing fecundity. These long-lived queens have a significantly lower level of oxidative damage, including oxidative DNA damage, than workers, soldiers and nymphs. The lower levels of damage appear to be due to increased catalase, an enzyme that protects against oxidative stress.

Reproduction

Alates swarming during nuptial flight after rain

Termite alates only leave the colony when a nuptial flight takes place. Alate males and females pair up together and then land in search of a suitable place for a colony. A termite king and queen do not mate until they find such a spot. When they do, they excavate a chamber big enough for both, close up the entrance and proceed to mate. After mating, the pair never go outside and spend the rest of their lives in the nest. Nuptial flight time varies in each species. For example, alates in certain species emerge during the day in summer while others emerge during the winter. The nuptial flight may also begin at dusk, when the alates swarm around areas with many lights. The time when nuptial flight begins depends on the environmental conditions, the time of day, moisture, wind speed and precipitation. The number of termites in a colony also varies, with the larger species typically having 100–1,000 individuals. However, some termite colonies, including those with many individuals, can number in the millions.

The queen only lays 10–20 eggs in the very early stages of the colony, but lays as many as 1,000 a day when the colony is several years old. At maturity, a primary queen has a great capacity to lay eggs. In some species, the mature queen has a greatly distended abdomen and may produce 40,000 eggs a day. The two mature ovaries may have some 2,000 ovarioles each. The abdomen increases the queen's body length to several times more than before mating and reduces her ability to move freely; attendant workers provide assistance.

Egg grooming behaviour of Reticulitermes speratus workers in a nursery cell

The king grows only slightly larger after initial mating and continues to mate with the queen for life (a termite queen can live between 30 to 50 years); this is very different from ant colonies, in which a queen mates once with the males and stores the gametes for life, as the male ants die shortly after mating. If a queen is absent, a termite king produces pheromones which encourage the development of replacement termite queens. As the queen and king are monogamous, sperm competition does not occur.

Termites going through incomplete metamorphosis on the path to becoming alates form a subcaste in certain species of termite, functioning as potential supplementary reproductives. These supplementary reproductives only mature into primary reproductives upon the death of a king or queen, or when the primary reproductives are separated from the colony. Supplementaries have the ability to replace a dead primary reproductive, and there may also be more than a single supplementary within a colony. Some queens have the ability to switch from sexual reproduction to asexual reproduction. Studies show that while termite queens mate with the king to produce colony workers, the queens reproduce their replacements (neotenic queens) parthenogenetically.

The neotropical termite Embiratermes neotenicus and several other related species produce colonies that contain a primary king accompanied by a primary queen or by up to 200 neotenic queens that had originated through thelytokous parthenogenesis of a founding primary queen. The form of parthenogenesis likely employed maintains heterozygosity in the passage of the genome from mother to daughter, thus avoiding inbreeding depression.

Diet

Termite faecal pellets

Termites are detritivores, consuming dead plants at any level of decomposition. They also play a vital role in the ecosystem by recycling waste material such as dead wood, faeces and plants. Many species eat cellulose, having a specialised midgut that breaks down the fibre. Termites are considered to be a major source (11%) of atmospheric methane, one of the prime greenhouse gases, produced from the breakdown of cellulose. Termites rely primarily upon symbiotic protozoa (metamonads) and other microbes such as flagellate protists in their guts to digest the cellulose for them, allowing them to absorb the end products for their own use. The microbial ecosystem present in the termite gut contains many species found nowhere else on Earth. Termites hatch without these symbionts present in their guts, and develop them after fed a culture from other termites. Gut protozoa, such as Trichonympha, in turn, rely on symbiotic bacteria embedded on their surfaces to produce some of the necessary digestive enzymes. Most higher termites, especially in the family Termitidae, can produce their own cellulase enzymes, but they rely primarily upon the bacteria. The flagellates have been lost in Termitidae. Researches have found species of spirochetes living in termite guts capable of fixing atmospheric nitrogen to a form usable by the insect. Scientists' understanding of the relationship between the termite digestive tract and the microbial endosymbionts is still rudimentary; what is true in all termite species, however, is that the workers feed the other members of the colony with substances derived from the digestion of plant material, either from the mouth or anus. Judging from closely related bacterial species, it is strongly presumed that the termites' and cockroach's gut microbiota derives from their dictyopteran ancestors.

Certain species such as Gnathamitermes tubiformans have seasonal food habits. For example, they may preferentially consume Red three-awn (Aristida longiseta) during the summer, Buffalograss (Buchloe dactyloides) from May to August, and blue grama Bouteloua gracilis during spring, summer and autumn. Colonies of G. tubiformans consume less food in spring than they do during autumn when their feeding activity is high.

Various woods differ in their susceptibility to termite attack; the differences are attributed to such factors as moisture content, hardness, and resin and lignin content. In one study, the drywood termite Cryptotermes brevis strongly preferred poplar and maple woods to other woods that were generally rejected by the termite colony. These preferences may in part have represented conditioned or learned behaviour.

Some species of termite practice fungiculture. They maintain a "garden" of specialised fungi of genus Termitomyces, which are nourished by the excrement of the insects. When the fungi are eaten, their spores pass undamaged through the intestines of the termites to complete the cycle by germinating in the fresh faecal pellets. Molecular evidence suggests that the family Macrotermitinae developed agriculture about 31 million years ago. It is assumed that more than 90 percent of dry wood in the semiarid savannah ecosystems of Africa and Asia are reprocessed by these termites. Originally living in the rainforest, fungus farming allowed them to colonise the African savannah and other new environments, eventually expanding into Asia.

Depending on their feeding habits, termites are placed into two groups: the lower termites and higher termites. The lower termites predominately feed on wood. As wood is difficult to digest, termites prefer to consume fungus-infected wood because it is easier to digest and the fungi are high in protein. Meanwhile, the higher termites consume a wide variety of materials, including faeces, humus, grass, leaves and roots. The gut of the lower termites contains many species of bacteria along with protozoa and Holomastigotoides, while the higher termites only have a few species of bacteria with no protozoa.

Predators

Crab spider with a captured alate

Termites are consumed by a wide variety of predators. One termite species alone, Hodotermes mossambicus, was found in the stomach contents of 65 birds and 19 mammals. Arthropods such as ants, centipedes, cockroaches, crickets, dragonflies, scorpions and spiders, reptiles such as lizards, and amphibians such as frogs and toads consume termites, with two spiders in the family Ammoxenidae being specialist termite predators. Other predators include aardvarks, aardwolves, anteaters, bats, bears, bilbies, many birds, echidnas, foxes, galagos, numbats, mice and pangolins. The aardwolf is an insectivorous mammal that primarily feeds on termites; it locates its food by sound and also by detecting the scent secreted by the soldiers; a single aardwolf is capable of consuming thousands of termites in a single night by using its long, sticky tongue. Sloth bears break open mounds to consume the nestmates, while chimpanzees have developed tools to "fish" termites from their nest. Wear pattern analysis of bone tools used by the early hominin Paranthropus robustus suggests that they used these tools to dig into termite mounds.

A Matabele ant (Megaponera analis) kills a Macrotermes bellicosus termite soldier during a raid.

Among all predators, ants are the greatest enemy to termites. Some ant genera are specialist predators of termites. For example, Megaponera is a strictly termite-eating (termitophagous) genus that perform raiding activities, some lasting several hours. Paltothyreus tarsatus is another termite-raiding species, with each individual stacking as many termites as possible in its mandibles before returning home, all the while recruiting additional nestmates to the raiding site through chemical trails. The Malaysian basicerotine ants Eurhopalothrix heliscata uses a different strategy of termite hunting by pressing themselves into tight spaces, as they hunt through rotting wood housing termite colonies. Once inside, the ants seize their prey by using their short but sharp mandibles. Tetramorium uelense is a specialised predator species that feeds on small termites. A scout recruits 10–30 workers to an area where termites are present, killing them by immobilising them with their stinger. Centromyrmex and Iridomyrmex colonies sometimes nest in termite mounds, and so the termites are preyed on by these ants. No evidence for any kind of relationship (other than a predatory one) is known. Other ants, including Acanthostichus, Camponotus, Crematogaster, Cylindromyrmex, Leptogenys, Odontomachus, Ophthalmopone, Pachycondyla, Rhytidoponera, Solenopsis and Wasmannia, also prey on termites. In contrast to all these ant species, and despite their enormous diversity of prey, Dorylus ants rarely consume termites.

Ants are not the only invertebrates that perform raids. Many sphecoid wasps and several species including Polybia and Angiopolybia are known to raid termite mounds during the termites' nuptial flight.

Parasites, pathogens and viruses

Termites are less likely to be attacked by parasites than bees, wasps and ants, as they are usually well protected in their mounds. Nevertheless, termites are infected by a variety of parasites. Some of these include dipteran flies, Pyemotes mites, and a large number of nematode parasites. Most nematode parasites are in the order Rhabditida; others are in the genus Mermis, Diplogaster aerivora and Harteria gallinarum. Under imminent threat of an attack by parasites, a colony may migrate to a new location. Certain fungal pathogens such as Aspergillus nomius and Metarhizium anisopliae are, however, major threats to a termite colony as they are not host-specific and may infect large portions of the colony; transmission usually occurs via direct physical contact. M. anisopliae is known to weaken the termite immune system. Infection with A. nomius only occurs when a colony is under great stress. Over 34 fungal species are known to live as parasites on the exoskeleton of termites, with many being host-specific and only causing indirect harm to their host.

Termites are infected by viruses including Entomopoxvirinae and the Nuclear Polyhedrosis Virus.

Locomotion and foraging

Because the worker and soldier castes lack wings and thus never fly, and the reproductives use their wings for just a brief amount of time, termites predominantly rely upon their legs to move about.

Foraging behaviour depends on the type of termite. For example, certain species feed on the wood structures they inhabit, and others harvest food that is near the nest. Most workers are rarely found out in the open, and do not forage unprotected; they rely on sheeting and runways to protect them from predators. Subterranean termites construct tunnels and galleries to look for food, and workers who manage to find food sources recruit additional nestmates by depositing a phagostimulant pheromone that attracts workers. Foraging workers use semiochemicals to communicate with each other, and workers who begin to forage outside of their nest release trail pheromones from their sternal glands. In one species, Nasutitermes costalis, there are three phases in a foraging expedition: first, soldiers scout an area. When they find a food source, they communicate to other soldiers and a small force of workers starts to emerge. In the second phase, workers appear in large numbers at the site. The third phase is marked by a decrease in the number of soldiers present and an increase in the number of workers. Isolated termite workers may engage in Lévy flight behaviour as an optimised strategy for finding their nestmates or foraging for food.

Competition

Competition between two colonies always results in agonistic behaviour towards each other, resulting in fights. These fights can cause mortality on both sides and, in some cases, the gain or loss of territory. "Cemetery pits" may be present, where the bodies of dead termites are buried.

Studies show that when termites encounter each other in foraging areas, some of the termites deliberately block passages to prevent other termites from entering. Dead termites from other colonies found in exploratory tunnels leads to the isolation of the area and thus the need to construct new tunnels. Conflict between two competitors does not always occur. For example, though they might block each other's passages, colonies of Macrotermes bellicosus and Macrotermes subhyalinus are not always aggressive towards each other. Suicide cramming is known in Coptotermes formosanus. Since C. formosanus colonies may get into physical conflict, some termites squeeze tightly into foraging tunnels and die, successfully blocking the tunnel and ending all agonistic activities.

Among the reproductive caste, neotenic queens may compete with each other to become the dominant queen when there are no primary reproductives. This struggle among the queens leads to the elimination of all but a single queen, which, with the king, takes over the colony.

Ants and termites may compete with each other for nesting space. In particular, ants that prey on termites usually have a negative impact on arboreal nesting species.

Communication

Hordes of Nasutitermes on a march for food, following and leaving trail pheromones

Most termites are blind, so communication primarily occurs through chemical, mechanical and pheromonal cues. These methods of communication are used in a variety of activities, including foraging, locating reproductives, construction of nests, recognition of nestmates, nuptial flight, locating and fighting enemies, and defending the nests. The most common way of communicating is through antennation. A number of pheromones are known, including contact pheromones (which are transmitted when workers are engaged in trophallaxis or grooming) and alarm, trail and sex pheromones. The alarm pheromone and other defensive chemicals are secreted from the frontal gland. Trail pheromones are secreted from the sternal gland, and sex pheromones derive from two glandular sources: the sternal and tergal glands. When termites go out to look for food, they forage in columns along the ground through vegetation. A trail can be identified by the faecal deposits or runways that are covered by objects. Workers leave pheromones on these trails, which are detected by other nestmates through olfactory receptors. Termites can also communicate through mechanical cues, vibrations, and physical contact. These signals are frequently used for alarm communication or for evaluating a food source.

When termites construct their nests, they use predominantly indirect communication. No single termite would be in charge of any particular construction project. Individual termites react rather than think, but at a group level, they exhibit a sort of collective cognition. Specific structures or other objects such as pellets of soil or pillars cause termites to start building. The termite adds these objects onto existing structures, and such behaviour encourages building behaviour in other workers. The result is a self-organised process whereby the information that directs termite activity results from changes in the environment rather than from direct contact among individuals.

Termites can distinguish nestmates and non-nestmates through chemical communication and gut symbionts: chemicals consisting of hydrocarbons released from the cuticle allow the recognition of alien termite species. Each colony has its own distinct odour. This odour is a result of genetic and environmental factors such as the termites' diet and the composition of the bacteria within the termites' intestines.

Defence

See also: Insect defences
Termites rush to a damaged area of the nest.

Termites rely on alarm communication to defend a colony. Alarm pheromones can be released when the nest has been breached or is being attacked by enemies or potential pathogens. Termites always avoid nestmates infected with Metarhizium anisopliae spores, through vibrational signals released by infected nestmates. Other methods of defence include intense jerking and secretion of fluids from the frontal gland and defecating faeces containing alarm pheromones.

In some species, some soldiers block tunnels to prevent their enemies from entering the nest, and they may deliberately rupture themselves as an act of defence. In cases where the intrusion is coming from a breach that is larger than the soldier's head, soldiers form a phalanx-like formation around the breach and bite at intruders. If an invasion carried out by Megaponera analis is successful, an entire colony may be destroyed, although this scenario is rare.

To termites, any breach of their tunnels or nests is a cause for alarm. When termites detect a potential breach, the soldiers usually bang their heads, apparently to attract other soldiers for defence and to recruit additional workers to repair any breach. Additionally, an alarmed termite bumps into other termites which causes them to be alarmed and to leave pheromone trails to the disturbed area, which is also a way to recruit extra workers.

Nasute termite soldiers on rotten wood

The pantropical subfamily Nasutitermitinae has a specialised caste of soldiers, known as nasutes, that have the ability to exude noxious liquids through a horn-like frontal projection that they use for defence. Nasutes have lost their mandibles through the course of evolution and must be fed by workers. A wide variety of monoterpene hydrocarbon solvents have been identified in the liquids that nasutes secrete. Similarly, Formosan subterranean termites have been known to secrete naphthalene to protect their nests.

Soldiers of the species Globitermes sulphureus commit suicide by autothysis – rupturing a large gland just beneath the surface of their cuticles. The thick, yellow fluid in the gland becomes very sticky on contact with the air, entangling ants or other insects that are trying to invade the nest. Another termite, Neocapriterme taracua, also engages in suicidal defence. Workers physically unable to use their mandibles while in a fight form a pouch full of chemicals, then deliberately rupture themselves, releasing toxic chemicals that paralyse and kill their enemies. The soldiers of the neotropical termite family Serritermitidae have a defence strategy which involves front gland autothysis, with the body rupturing between the head and abdomen. When soldiers guarding nest entrances are attacked by intruders, they engage in autothysis, creating a block that denies entry to any attacker.

Workers use several different strategies to deal with their dead, including burying, cannibalism, and avoiding a corpse altogether. To avoid pathogens, termites occasionally engage in necrophoresis, in which a nestmate carries away a corpse from the colony to dispose of it elsewhere. Which strategy is used depends on the nature of the corpse a worker is dealing with (i.e. the age of the carcass).

Relationship with other organisms

Rhizanthella gardneri is the only orchid known to be pollinated by termites.

A species of fungus is known to mimic termite eggs, successfully avoiding its natural predators. These small brown balls, known as "termite balls", rarely kill the eggs, and in some cases the workers tend to them. This fungus mimics these eggs by producing a cellulose-digesting enzyme known as glucosidases. A unique mimicking behaviour exists between various species of Trichopsenius beetles and certain termite species within Reticulitermes. The beetles share the same cuticle hydrocarbons as the termites and even biosynthesize them. This chemical mimicry allows the beetles to integrate themselves within the termite colonies. The developed appendages on the physogastric abdomen of Austrospirachtha mimetes allows the beetle to mimic a termite worker.

Some species of ant are known to capture termites to use as a fresh food source later on, rather than killing them. For example, Formica nigra captures termites, and those who try to escape are immediately seized and driven underground. Certain species of ants in the subfamily Ponerinae conduct these raids although other ant species go in alone to steal the eggs or nymphs. Ants such as Megaponera analis attack the outside of mounds and Dorylinae ants attack underground. Despite this, some termites and ants can coexist peacefully. Some species of termite, including Nasutitermes corniger, form associations with certain ant species to keep away predatory ant species. The earliest known association between Azteca ants and Nasutitermes termites date back to the Oligocene to Miocene period.

An ant raiding party collecting Pseudocanthotermes militaris termites after a successful raid

54 species of ants are known to inhabit Nasutitermes mounds, both occupied and abandoned ones. One reason many ants live in Nasutitermes mounds is due to the termites' frequent occurrence in their geographical range; another is to protect themselves from floods. Iridomyrmex also inhabits termite mounds although no evidence for any kind of relationship (other than a predatory one) is known. In rare cases, certain species of termites live inside active ant colonies. Some invertebrate organisms such as beetles, caterpillars, flies and millipedes are termitophiles and dwell inside termite colonies (they are unable to survive independently). As a result, certain beetles and flies have evolved with their hosts. They have developed a gland that secrete a substance that attracts the workers by licking them. Mounds may also provide shelter and warmth to birds, lizards, snakes and scorpions.

Termites are known to carry pollen and regularly visit flowers, so are regarded as potential pollinators for a number of flowering plants. One flower in particular, Rhizanthella gardneri, is regularly pollinated by foraging workers, and it is perhaps the only Orchidaceae flower in the world to be pollinated by termites.

Many plants have developed effective defences against termites. However, seedlings are vulnerable to termite attacks and need additional protection, as their defence mechanisms only develop when they have passed the seedling stage. Defence is typically achieved by secreting antifeedant chemicals into the woody cell walls. This reduces the ability of termites to efficiently digest the cellulose. A commercial product, "Blockaid", has been developed in Australia that uses a range of plant extracts to create a paint-on nontoxic termite barrier for buildings. An extract of a species of Australian figwort, Eremophila, has been shown to repel termites; tests have shown that termites are strongly repelled by the toxic material to the extent that they will starve rather than consume the food. When kept close to the extract, they become disoriented and eventually die.

Relationship with the environment

Termite populations can be substantially impacted by environmental changes including those caused by human intervention. A Brazilian study investigated the termite assemblages of three sites of Caatinga under different levels of anthropogenic disturbance in the semi-arid region of northeastern Brazil were sampled using 65 x 2 m transects. A total of 26 species of termites were present in the three sites, and 196 encounters were recorded in the transects. The termite assemblages were considerably different among sites, with a conspicuous reduction in both diversity and abundance with increased disturbance, related to the reduction of tree density and soil cover, and with the intensity of trampling by cattle and goats. The wood-feeders were the most severely affected feeding group.

Termite workers at work
An arboreal termite nest in Mexico
Termite nest in a Banksia, Palm Beach, Sydney.

A termite nest can be considered as being composed of two parts, the inanimate and the animate. The animate is all of the termites living inside the colony, and the inanimate part is the structure itself, which is constructed by the termites. Nests can be broadly separated into three main categories: subterranean (completely below ground), epigeal (protruding above the soil surface), and arboreal (built above ground, but always connected to the ground via shelter tubes). Epigeal nests (mounds) protrude from the earth with ground contact and are made out of earth and mud. A nest has many functions such as providing a protected living space and providing shelter against predators. Most termites construct underground colonies rather than multifunctional nests and mounds. Primitive termites of today nest in wooden structures such as logs, stumps and the dead parts of trees, as did termites millions of years ago.

To build their nests, termites primarily use faeces, which have many desirable properties as a construction material. Other building materials include partly digested plant material, used in carton nests (arboreal nests built from faecal elements and wood), and soil, used in subterranean nest and mound construction. Not all nests are visible, as many nests in tropical forests are located underground. Species in the subfamily Apicotermitinae are good examples of subterranean nest builders, as they only dwell inside tunnels. Other termites live in wood, and tunnels are constructed as they feed on the wood. Nests and mounds protect the termites' soft bodies against desiccation, light, pathogens and parasites, as well as providing a fortification against predators. Nests made out of carton are particularly weak, and so the inhabitants use counter-attack strategies against invading predators.

Arboreal carton nests of mangrove swamp-dwelling Nasutitermes are enriched in lignin and depleted in cellulose and xylans. This change is caused by bacterial decay in the gut of the termites: they use their faeces as a carton building material. Arboreal termites nests can account for as much as 2% of above ground carbon storage in Puerto Rican mangrove swamps. These Nasutitermes nests are mainly composed of partially biodegraded wood material from the stems and branches of mangrove trees, namely, Rhizophora mangle (red mangrove), Avicennia germinans (black mangrove) and Laguncularia racemosa (white mangrove).

Some species build complex nests called polycalic nests; this habitat is called polycalism. Polycalic species of termites form multiple nests, or calies, connected by subterranean chambers. The termite genera Apicotermes and Trinervitermes are known to have polycalic species. Polycalic nests appear to be less frequent in mound-building species although polycalic arboreal nests have been observed in a few species of Nasutitermes.

Mounds

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Nests are considered mounds if they protrude from the earth's surface. A mound provides termites the same protection as a nest but is stronger. Mounds located in areas with torrential and continuous rainfall are at risk of mound erosion due to their clay-rich construction. Those made from carton can provide protection from the rain, and in fact can withstand high precipitation. Certain areas in mounds are used as strong points in case of a breach. For example, Cubitermes colonies build narrow tunnels used as strong points, as the diameter of the tunnels is small enough for soldiers to block. A highly protected chamber, known as the "queen's cell", houses the queen and king and is used as a last line of defence.

Species in the genus Macrotermes arguably build the most complex structures in the insect world, constructing enormous mounds. These mounds are among the largest in the world, reaching a height of 8 to 9 metres (26 to 29 feet), and consist of chimneys, pinnacles and ridges. Another termite species, Amitermes meridionalis, can build nests 3 to 4 metres (9 to 13 feet) high and 2.5 metres (8 feet) wide. The tallest mound ever recorded was 12.8 metres (42 ft) long found in the Democratic Republic of the Congo.

The sculptured mounds sometimes have elaborate and distinctive forms, such as those of the compass termite (Amitermes meridionalis and A. laurensis), which builds tall, wedge-shaped mounds with the long axis oriented approximately north–south, which gives them their common name. This orientation has been experimentally shown to assist thermoregulation. The north–south orientation causes the internal temperature of a mound to increase rapidly during the morning while avoiding overheating from the midday sun. The temperature then remains at a plateau for the rest of the day until the evening.

Shelter tubes

Nasutiterminae shelter tubes on a tree trunk provide cover for the trail from nest to forest floor.

Termites construct shelter tubes, also known as earthen tubes or mud tubes, that start from the ground. These shelter tubes can be found on walls and other structures. Constructed by termites during the night, a time of higher humidity, these tubes provide protection to termites from potential predators, especially ants. Shelter tubes also provide high humidity and darkness and allow workers to collect food sources that cannot be accessed in any other way. These passageways are made from soil and faeces and are normally brown in colour. The size of these shelter tubes depends on the number of food sources that are available. They range from less than 1 cm to several cm in width, but may be dozens of metres in length.

As pests

Termite mound as an obstacle on a runway at Khorixas (Namibia)
Termite damage on external structure

Owing to their wood-eating habits, many termite species can do significant damage to unprotected buildings and other wooden structures. Termites play an important role as decomposers of wood and vegetative material, and the conflict with humans occurs where structures and landscapes containing structural wood components, cellulose derived structural materials and ornamental vegetation provide termites with a reliable source of food and moisture. Their habit of remaining concealed often results in their presence being undetected until the timbers are severely damaged, with only a thin exterior layer of wood remaining, which protects them from the environment. Of the 3,106 species known, only 183 species cause damage; 83 species cause significant damage to wooden structures. In North America, 18 subterranean species are pests; in Australia, 16 species have an economic impact; in the Indian subcontinent 26 species are considered pests, and in tropical Africa, 24. In Central America and the West Indies, there are 17 pest species. Among the termite genera, Coptotermes has the highest number of pest species of any genus, with 28 species known to cause damage. Less than 10% of drywood termites are pests, but they infect wooden structures and furniture in tropical, subtropical and other regions. Dampwood termites only attack lumber material exposed to rainfall or soil.

Drywood termites thrive in warm climates, and human activities can enable them to invade homes since they can be transported through contaminated goods, containers and ships. Colonies of termites have been seen thriving in warm buildings located in cold regions. Some termites are considered invasive species. Cryptotermes brevis, the most widely introduced invasive termite species in the world, has been introduced to all the islands in the West Indies and to Australia.

Termite damage in wooden house stumps

In addition to causing damage to buildings, termites can also damage food crops. Termites may attack trees whose resistance to damage is low but generally ignore fast-growing plants. Most attacks occur at harvest time; crops and trees are attacked during the dry season.

The damage caused by termites costs the southwestern United States approximately $1.5 billion each year in wood structure damage, but the true cost of damage worldwide cannot be determined. Drywood termites are responsible for a large proportion of the damage caused by termites. The goal of termite control is to keep structures and susceptible ornamental plants free from termites.; Structures may be homes or business, or elements such as wooden fence posts and telephone poles. Regular and thorough inspections by a trained professional may be necessary to detect termite activity in the absence of more obvious signs like termite swarmers or alates inside or adjacent to a structure. Termite monitors made of wood or cellulose adjacent to a structure may also provide indication of termite foraging activity where it will be in conflict with humans. Termites can be controlled by application of Bordeaux mixture or other substances that contain copper such as chromated copper arsenate. In the United states, application of a soil termiticide with the active ingredient Fipronil, such as Termidor SC or Taurus SC, by a licensed professional, is a common remedy approved by the Environmental Protection Agency for economically significant subterranean termites. A growing demand for alternative, green, and "more natural" extermination methods has increased demand for mechanical and biological control methods such as Orange Oil.

To better control the population of termites, various methods have been developed to track termite movements. One early method involved distributing termite bait laced with immunoglobulin G (IgG) marker proteins from rabbits or chickens. Termites collected from the field could be tested for the rabbit-IgG markers using a rabbit-IgG-specific assay. More recently developed, less expensive alternatives include tracking the termites using egg white, cow milk, or soy milk proteins, which can be sprayed on termites in the field. Termites bearing these proteins can be traced using a protein-specific ELISA test.

In 1994 termites, of the species Reticulitermes grassei, were identified in two bungalows in Saunton, Devon. Anecdotal evidence suggests the infestation could date back 70 years before the official identification. There are reports that gardeners had seen white ants and that a greenhouse had had to be replaced in the past. The Saunton infestation was the first and only colony ever recorded in the UK. In 1998 Termite Eradication Programme was set-up, with the intention of containing and eradicating the colony. The TEP was managed by the Ministry of Housing, Communities & Local Government (now the Department for Levelling Up, Housing and Communities.) The TEP used 'insect growth regulators' to prevent the termites from reaching maturity and reproducing. In 2021 the UK’s Termite Eradication Programme announced the eradication of the colony, the first time a country has eradicated termites.

As food

See also: Entomophagy
Mozambican boys from the Yawo tribe collecting flying termites
These flying alates were collected as they came out of their nests in the ground during the early days of the rainy season.

43 termite species are used as food by humans or are fed to livestock. These insects are particularly important in impoverished countries where malnutrition is common, as the protein from termites can help improve the human diet. Termites are consumed in many regions globally, but this practice has only become popular in developed nations in recent years.

Termites are consumed by people in many different cultures around the world. In many parts of Africa, the alates are an important factor in the diets of native populations. Groups have different ways of collecting or cultivating insects; sometimes collecting soldiers from several species. Though harder to acquire, queens are regarded as a delicacy. Termite alates are high in nutrition with adequate levels of fat and protein. They are regarded as pleasant in taste, having a nut-like flavour after they are cooked.

Alates are collected when the rainy season begins. During a nuptial flight, they are typically seen around lights to which they are attracted, and so nets are set up on lamps and captured alates are later collected. The wings are removed through a technique that is similar to winnowing. The best result comes when they are lightly roasted on a hot plate or fried until crisp. Oil is not required as their bodies usually contain sufficient amounts of oil. Termites are typically eaten when livestock is lean and tribal crops have not yet developed or produced any food, or if food stocks from a previous growing season are limited.

In addition to Africa, termites are consumed in local or tribal areas in Asia and North and South America. In Australia, Indigenous Australians are aware that termites are edible but do not consume them even in times of scarcity; there are few explanations as to why. Termite mounds are the main sources of soil consumption (geophagy) in many countries including Kenya, Tanzania, Zambia, Zimbabwe and South Africa. Researchers have suggested that termites are suitable candidates for human consumption and space agriculture, as they are high in protein and can be used to convert inedible waste to consumable products for humans.

In agriculture

Scientists have developed a more affordable method of tracing the movement of termites using traceable proteins.

Termites can be major agricultural pests, particularly in East Africa and North Asia, where crop losses can be severe (3–100% in crop loss in Africa). Counterbalancing this is the greatly improved water infiltration where termite tunnels in the soil allow rainwater to soak in deeply, which helps reduce runoff and consequent soil erosion through bioturbation. In South America, cultivated plants such as eucalyptus, upland rice and sugarcane can be severely damaged by termite infestations, with attacks on leaves, roots and woody tissue. Termites can also attack other plants, including cassava, coffee, cotton, fruit trees, maize, peanuts, soybeans and vegetables. Mounds can disrupt farming activities, making it difficult for farmers to operate farming machinery; however, despite farmers' dislike of the mounds, it is often the case that no net loss of production occurs. Termites can be beneficial to agriculture, such as by boosting crop yields and enriching the soil. Termites and ants can re-colonise untilled land that contains crop stubble, which colonies use for nourishment when they establish their nests. The presence of nests in fields enables larger amounts of rainwater to soak into the ground and increases the amount of nitrogen in the soil, both essential for the growth of crops.

In science and technology

The termite gut has inspired various research efforts aimed at replacing fossil fuels with cleaner, renewable energy sources. Termites are efficient bioreactors, capable of producing two litres of hydrogen from a single sheet of paper. Approximately 200 species of microbes live inside the termite hindgut, releasing the hydrogen that was trapped inside wood and plants that they digest. Through the action of unidentified enzymes in the termite gut, lignocellulose polymers are broken down into sugars and are transformed into hydrogen. The bacteria within the gut turns the sugar and hydrogen into cellulose acetate, an acetate ester of cellulose on which termites rely for energy. Community DNA sequencing of the microbes in the termite hindgut has been employed to provide a better understanding of the metabolic pathway. Genetic engineering may enable hydrogen to be generated in bioreactors from woody biomass.

The development of autonomous robots capable of constructing intricate structures without human assistance has been inspired by the complex mounds that termites build. These robots work independently and can move by themselves on a tracked grid, capable of climbing and lifting up bricks. Such robots may be useful for future projects on Mars, or for building levees to prevent flooding.

Termites use sophisticated means to control the temperatures of their mounds. As discussed above, the shape and orientation of the mounds of the Australian compass termite stabilises their internal temperatures during the day. As the towers heat up, the solar chimney effect (stack effect) creates an updraft of air within the mound. Wind blowing across the tops of the towers enhances the circulation of air through the mounds, which also include side vents in their construction. The solar chimney effect has been in use for centuries in the Middle East and Near East for passive cooling, as well as in Europe by the Romans. It is only relatively recently, however, that climate responsive construction techniques have become incorporated into modern architecture. Especially in Africa, the stack effect has become a popular means to achieve natural ventilation and passive cooling in modern buildings.

In culture

The pink-hued Eastgate Centre

The Eastgate Centre is a shopping centre and office block in central Harare, Zimbabwe, whose architect, Mick Pearce, used passive cooling inspired by that used by the local termites. It was the first major building exploiting termite-inspired cooling techniques to attract international attention. Other such buildings include the Learning Resource Center at the Catholic University of Eastern Africa and the Council House 2 building in Melbourne, Australia.

Few zoos hold termites, due to the difficulty in keeping them captive and to the reluctance of authorities to permit potential pests. One of the few that do, the Zoo Basel in Switzerland, has two thriving Macrotermes bellicosus populations – resulting in an event very rare in captivity: the mass migrations of young flying termites. This happened in September 2008, when thousands of male termites left their mound each night, died, and covered the floors and water pits of the house holding their exhibit.

African tribes in several countries have termites as totems, and for this reason tribe members are forbidden to eat the reproductive alates. Termites are widely used in traditional popular medicine; they are used as treatments for diseases and other conditions such as asthma, bronchitis, hoarseness, influenza, sinusitis, tonsillitis and whooping cough. In Nigeria, Macrotermes nigeriensis is used for spiritual protection and to treat wounds and sick pregnant women. In Southeast Asia, termites are used in ritual practices. In Malaysia, Singapore and Thailand, termite mounds are commonly worshiped among the populace. Abandoned mounds are viewed as structures created by spirits, believing a local guardian dwells within the mound; this is known as Keramat and Datok Kong. In urban areas, local residents construct red-painted shrines over mounds that have been abandoned, where they pray for good health, protection and luck.

  1. It is unknown whether the termite was female or male. If it was a female, the body length would be far greater than 25 millimetres when mature.
  1. Behrensmeyer, A. K.; Turner, A. "Fossilworks, Gateway to the Paleobiology Database".
  2. Engel, M.S.; Grimaldi, D.A.; Krishna, K. (2009). "Termites (Isoptera): their phylogeny, classification, and rise to ecological dominance". American Museum Novitates (3650): 1–27. doi:10.1206/651.1. hdl:2246/5969. ISSN 0003-0082. S2CID 56166416.
  3. Evangelista, Dominic A.; Wipfler, Benjamin; Béthoux, Olivier; Donath, Alexander; Fujita, Mari; Kohli, Manpreet K.; Legendre, Frédéric; Liu, Shanlin; Machida, Ryuichiro; Misof, Bernhard; Peters, Ralph S. (2019-01-30). "An integrative phylogenomic approach illuminates the evolutionary history of cockroaches and termites (Blattodea)". Proceedings of the Royal Society B: Biological Sciences. 286 (1895): 20182076. doi:10.1098/rspb.2018.2076. ISSN 0962-8452. PMC6364590. PMID 30963947.
  4. "Termite". Merriam-Webster.com.
  5. Bignell, Roisin & Lo 2010, p. 2.
  6. Cranshaw, W. (2013). "11". Bugs Rule!: An Introduction to the World of Insects. Princeton, New Jersey: Princeton University Press. p. 188. ISBN 978-0-691-12495-7.
  7. Lobeck, A. Kohl (1939). Geomorphology; an Introduction to the Study of Landscapes (1st ed.). University of California: McGraw Hill Book Company, Incorporated. pp. 431–432. ASIN B002P5O9SC.
  8. Harper, Douglas. "Termite". Online Etymology Dictionary.
  9. "Termite". Merriam-Webster Online Dictionary. Retrieved5 January 2015.
  10. Cleveland, L.R.; Hall, S.K.; Sanders, E.P.; Collier, J. (1934). "The Wood-Feeding Roach Cryptocercus, its protozoa, and the symbiosis between protozoa and roach". Memoirs of the American Academy of Arts and Sciences. 17 (2): 185–382. doi:10.1093/aesa/28.2.216.
  11. McKittrick, F.A. (1965). "A contribution to the understanding of cockroach-termite affinities". Annals of the Entomological Society of America. 58 (1): 18–22. doi:10.1093/aesa/58.1.18. PMID 5834489.
  12. Ware, J.L.; Litman, J.; Klass, K.-D.; Spearman, L.A. (2008). "Relationships among the major lineages of Dictyoptera: the effect of outgroup selection on dictyopteran tree topology". Systematic Entomology. 33 (3): 429–450. doi:10.1111/j.1365-3113.2008.00424.x. S2CID 86777253.
  13. Inward, D.; Beccaloni, G.; Eggleton, P. (2007). "Death of an order: a comprehensive molecular phylogenetic study confirms that termites are eusocial cockroaches". Biology Letters. 3 (3): 331–5. doi:10.1098/rsbl.2007.0102. PMC2464702. PMID 17412673.
  14. Eggleton, P.; Beccaloni, G.; Inward, D. (2007). "Response to Lo et al.". Biology Letters. 3 (5): 564–565. doi:10.1098/rsbl.2007.0367. PMC2391203.
  15. Ohkuma, M.; Noda, S.; Hongoh, Y.; Nalepa, C.A.; Inoue, T. (2009). "Inheritance and diversification of symbiotic trichonymphid flagellates from a common ancestor of termites and the cockroach Cryptocercus". Proceedings of the Royal Society B: Biological Sciences. 276 (1655): 239–245. doi:10.1098/rspb.2008.1094. PMC2674353. PMID 18812290.
  16. Lo, N.; Tokuda, G.; Watanabe, H.; Rose, H.; Slaytor, M.; Maekawa, K.; Bandi, C.; Noda, H. (June 2000). "Evidence from multiple gene sequences indicates that termites evolved from wood-feeding cockroaches". Current Biology. 10 (13): 801–814. doi:10.1016/S0960-9822(00)00561-3. PMID 10898984. S2CID 14059547.
  17. Grimaldi, D.; Engel, M.S. (2005).Evolution of the insects (1st ed.). Cambridge: Cambridge University Press. p. 237. ISBN 978-0-521-82149-0.
  18. Klass, K.D.; Nalepa, C.; Lo, N. (2008). "Wood-feeding cockroaches as models for termite evolution (Insecta: Dictyoptera): Cryptocercus vs. Parasphaeria boleiriana". Molecular Phylogenetics & Evolution. 46 (3): 809–817. doi:10.1016/j.ympev.2007.11.028. PMID 18226554.
  19. Lo, N.; Engel, M.S.; Cameron, S.; Nalepa, C.A.; Tokuda, G.; Grimaldi, D.; Kitade, O..; Krishna, K.; Klass, K.-D.; Maekawa, K.; Miura, T.; Thompson, G.J. (2007). "Comment. Save Isoptera: a comment on Inward et al.". Biology Letters. 3 (5): 562–563. doi:10.1098/rsbl.2007.0264. PMC2391185. PMID 17698448.
  20. Costa, James (2006). The other insect societies. Harvard University Press. pp. 135–136. ISBN 978-0-674-02163-1.
  21. Capinera, J.L. (2008).Encyclopedia of Entomology (2nd ed.). Dordrecht: Springer. pp. 3033–3037, 3754. ISBN 978-1-4020-6242-1.
  22. Vrsanky, P.; Aristov, D. (2014). "Termites (Isoptera) from the Jurassic/Cretaceous boundary: Evidence for the longevity of their earliest genera". European Journal of Entomology. 111 (1): 137–141. doi:10.14411/eje.2014.014.
  23. Poinar, G.O. (2009). "Description of an early Cretaceous termite (Isoptera: Kalotermitidae) and its associated intestinal protozoa, with comments on their co-evolution". Parasites & Vectors. 2 (1–17): 12. doi:10.1186/1756-3305-2-12. PMC2669471. PMID 19226475.
  24. Legendre, F.; Nel, A.; Svenson, G.J.; Robillard, T.; Pellens, R.; Grandcolas, P.; Escriva, H. (2015). "Phylogeny of Dictyoptera: Dating the Origin of Cockroaches, Praying Mantises and Termites with Molecular Data and Controlled Fossil Evidence". PLOS ONE. 10 (7): e0130127. Bibcode:2015PLoSO..1030127L. doi:10.1371/journal.pone.0130127. PMC4511787. PMID 26200914.
  25. Luo, Z.X.; Wible, J.R. (2005). "A Late Jurassic digging mammal and early mammalian diversification". Science. 308 (5718): 103–107. Bibcode:2005Sci...308..103L. doi:10.1126/science.1108875. PMID 15802602. S2CID 7031381.
  26. Rohr, D.M.; Boucot, A. J.; Miller, J.; Abbott, M. (1986). "Oldest termite nest from the Upper Cretaceous of west Texas". Geology. 14 (1): 87. Bibcode:1986Geo....14...87R. doi:10.1130/0091-7613(1986)14<87:OTNFTU>2.0.CO;2.
  27. Weesner, F.M. (1960). "Evolution and Biology of the Termites". Annual Review of Entomology. 5 (1): 153–170. doi:10.1146/annurev.en.05.010160.001101.
  28. Tilyard, R.J. (1937). "Kansas Permian insects. Part XX the cockroaches, or order Blattaria". American Journal of Science. 34 (201): 169–202, 249–276. Bibcode:1937AmJS...34..169T. doi:10.2475/ajs.s5-34.201.169.
  29. Henry, M.S. (2013). Symbiosis: Associations of Invertebrates, Birds, Ruminants, and Other Biota. New York, New York: Elsevier. p. 59. ISBN 978-1-4832-7592-5.
  30. Krishna, K.; Grimaldi, D.A.; Krishna, V.; Engel, M.S. (2013). "Treatise on the Isoptera of the world"(PDF). Bulletin of the American Museum of Natural History. 1. 377 (7): 1–200. doi:10.1206/377.1. S2CID 87276148.
  31. Bell, W.J.; Roth, L.M.; Nalepa, C.A. (2007).Cockroaches: ecology, behavior, and natural history. Baltimore, Md.: Johns Hopkins University Press. p. 161. ISBN 978-0-8018-8616-4.
  32. Engel, M. (2011). "Family-group names for termites (Isoptera), redux". ZooKeys (148): 171–184. doi:10.3897/zookeys.148.1682. PMC3264418. PMID 22287896.
  33. Thorne, Barbara L (1997). "Evolution of eusociality in termites"(PDF). Annual Review of Ecology and Systematics. 28 (5): 27–53. doi:10.1146/annurev.ecolsys.28.1.27. PMC349550. Archived from the original(PDF) on 2010-05-30.
  34. Harrison, Mark C.; Jongepier, Evelien; Robertson, Hugh M.; Arning, Nicolas; Bitard-Feildel, Tristan; Chao, Hsu; Childers, Christopher P.; Dinh, Huyen; Doddapaneni, Harshavardhan; Dugan, Shannon; Gowin, Johannes; Greiner, Carolin; Han, Yi; Hu, Haofu; Hughes, Daniel S. T.; Huylmans, Ann-Kathrin; Kemena, Carsten; Kremer, Lukas P. M.; Lee, Sandra L.; Lopez-Ezquerra, Alberto; Mallet, Ludovic; Monroy-Kuhn, Jose M.; Moser, Annabell; Murali, Shwetha C.; Muzny, Donna M.; Otani, Saria; Piulachs, Maria-Dolors; Poelchau, Monica; Qu, Jiaxin; Schaub, Florentine; Wada-Katsumata, Ayako; Worley, Kim C.; Xie, Qiaolin; Ylla, Guillem; Poulsen, Michael; Gibbs, Richard A.; Schal, Coby; Richards, Stephen; Belles, Xavier; Korb, Judith; Bornberg-Bauer, Erich (2018). "Hemimetabolous genomes reveal molecular basis of termite eusociality". Nature Ecology & Evolution. 2 (3): 557–566. doi:10.1038/s41559-017-0459-1. PMC6482461. PMID 29403074.
  35. "Termites had first castes". Nature. 530 (7590): 256. 2016. Bibcode:2016Natur.530Q.256.. doi:10.1038/530256a. S2CID 49905391.
  36. Terrapon, Nicolas; Li, Cai; Robertson, Hugh M.; Ji, Lu; Meng, Xuehong; Booth, Warren; Chen, Zhensheng; Childers, Christopher P.; Glastad, Karl M.; Gokhale, Kaustubh; et al. (2014). "Molecular traces of alternative social organization in a termite genome". Nature Communications. 5: 3636. Bibcode:2014NatCo...5.3636T. doi:10.1038/ncomms4636. PMID 24845553.
  37. Poulsen, Michael; Hu, Haofu; Li, Cai; Chen, Zhensheng; Xu, Luohao; Otani, Saria; Nygaard, Sanne; Nobre, Tania; Klaubauf, Sylvia; Schindler, Philipp M .; et al. (2014). "Complementary symbiont contributions to plant decomposition in a fungus-farming termite". Proceedings of the National Academy of Sciences. 111 (40): 14500–14505. Bibcode:2014PNAS..11114500P. doi:10.1073/pnas.1319718111. PMC4209977. PMID 25246537.
  38. Kohler, T; Dietrich, C; Scheffrahn, RH; Brune, A (2012). "High‐resolution analysis of gut environment and bacterial microbiota reveals functional compartmentation of the gut in wood‐feeding higher termites (Nasutitermes spp.)". Applied and Environmental Microbiology. 78 (13): 4691–4701. Bibcode:2012ApEnM..78.4691K. doi:10.1128/aem.00683-12. PMC3370480. PMID 22544239.
  39. "Termite Biology and Ecology". Division of Technology, Industry and Economics Chemicals Branch. United Nations Environment Programme. Archived from the original on 10 November 2014. Retrieved12 January 2015.
  40. Meyer, V.W.; Braack, L.E.O.; Biggs, H.C.; Ebersohn, C. (1999). "Distribution and density of termite mounds in the northern Kruger National Park, with specific reference to those constructed by Macrotermes Holmgren (Isoptera: Termitidae)". African Entomology. 7 (1): 123–130.
  41. Eggleton, Paul (2020). "The State of the World's Insects". Annual Review of Environment and Resources. 45: 61–82. doi:10.1146/annurev-environ-012420-050035.
  42. Sanderson, M.G. (1996). "Biomass of termites and their emissions of methane and carbon dioxide: A global database". Global Biogeochemical Cycles. 10 (4): 543–557. Bibcode:1996GBioC..10..543S. doi:10.1029/96GB01893.
  43. Heather, N.W. (1971). "The exotic drywood termite Cryptotermes brevis (Walker) (Isoptera : Kalotermitidae) and endemic Australian drywood termites in Queensland". Australian Journal of Entomology. 10 (2): 134–141. doi:10.1111/j.1440-6055.1971.tb00022.x.
  44. Claybourne, Anna (2013). A colony of ants, and other insect groups. Chicago, Ill.: Heinemann Library. p. 38. ISBN 978-1-4329-6487-0.
  45. Engel, M.S.; Gross, M. (2008). "A giant termite from the Late Miocene of Styria, Austria (Isoptera)". Naturwissenschaften. 96 (2): 289–295. Bibcode:2009NW.....96..289E. doi:10.1007/s00114-008-0480-y. PMID 19052720. S2CID 21795900.
  46. Heidecker, J.L.; Leuthold, R.H. (1984). "The organisation of collective foraging in the harvester termite Hodotermes mossambicus (Isoptera)". Behavioral Ecology and Sociobiology. 14 (3): 195–202. doi:10.1007/BF00299619. S2CID 22158321.
  47. Costa-Leonardo, A.M.; Haifig, I. (2010). "Pheromones and exocrine glands in Isoptera". Vitamins and Hormones. 83: 521–549. doi:10.1016/S0083-6729(10)83021-3. ISBN 9780123815163. PMID 20831960.
  48. Bignell, Roisin & Lo 2010, p. 7.
  49. Bignell, Roisin & Lo 2010, pp. 7–9.
  50. Bignell, Roisin & Lo 2010, p. 11.
  51. Robinson, W.H. (2005).Urban Insects and Arachnids: A Handbook of Urban Entomology. Cambridge: Cambridge University Press. p. 291. ISBN 978-1-139-44347-0.
  52. Bignell, Roisin & Lo 2010, p. 12.
  53. Riparbelli, M.G; Dallai, R; Mercati, D; Bu, Y; Callaini, G (2009). "Centriole symmetry: a big tale from small organisms". Cell Motility and the Cytoskeleton. 66 (12): 1100–5. doi:10.1002/cm.20417. PMID 19746415.
  54. Nalepa, C.A.; Lenz, M. (2000). "The ootheca of Mastotermes darwiniensis Froggatt (Isoptera: Mastotermitidae): homology with cockroach oothecae". Proceedings of the Royal Society B: Biological Sciences. 267 (1454): 1809–1813. doi:10.1098/rspb.2000.1214. PMC1690738. PMID 12233781.
  55. Crosland, M.W.J.; Su, N.Y.; Scheffrahn, R.H. (2005). "Arolia in termites (Isoptera): functional significance and evolutionary loss". Insectes Sociaux. 52 (1): 63–66. doi:10.1007/s00040-004-0779-4. S2CID 26873138.
  56. Bignell, Roisin & Lo 2010, p. 9.
  57. Bignell, Roisin & Lo 2010, p. 10.
  58. Bignell, Roisin & Lo 2010, p. 13.
  59. "Termites". Australian Museum. Retrieved8 January 2015.
  60. Machida, M.; Kitade, O.; Miura, T.; Matsumoto, T. (2001). "Nitrogen recycling through proctodeal trophallaxis in the Japanese damp-wood termite Hodotermopsis japonica (Isoptera, Termopsidae)". Insectes Sociaux. 48 (1): 52–56. doi:10.1007/PL00001745. ISSN 1420-9098. S2CID 21310420.
  61. Higashi, Masahiko; Yamamura, Norio; Abe, Takuya; Burns, Thomas P. (1991-10-22). "Why don't all termite species have a sterile worker caste?". Proceedings of the Royal Society B: Biological Sciences. 246 (1315): 25–29. doi:10.1098/rspb.1991.0120. ISSN 0962-8452. PMID 1684665. S2CID 23067349.
  62. Bignell, Roisin & Lo 2010, p. 18.
  63. Krishna, K. "Termite". Encyclopædia Britannica. Retrieved11 September 2015.
  64. Busvine, J.R. (2013). Insects and Hygiene The biology and control of insect pests of medical and domestic importance (3rd ed.). Boston, MA: Springer US. p. 545. ISBN 978-1-4899-3198-6.
  65. Meek, S.P. (1934). Termite Control at an Ordnance Storage Depot. American Defense Preparedness Association. p. 159.
  66. Prestwich, G.D. (1982). "From tetracycles to macrocycles". Tetrahedron. 38 (13): 1911–1919. doi:10.1016/0040-4020(82)80040-9.
  67. Prestwich, G. D.; Bentley, B.L.; Carpenter, E.J. (1980). "Nitrogen sources for neotropical nasute termites: Fixation and selective foraging". Oecologia. 46 (3): 397–401. Bibcode:1980Oecol..46..397P. doi:10.1007/BF00346270. ISSN 1432-1939. PMID 28310050. S2CID 6134800.
  68. Thorne, B. L.; Breisch, N. L.; Muscedere, M. L. (2003-10-28). "Evolution of eusociality and the soldier caste in termites: Influence of intraspecific competition and accelerated inheritance". Proceedings of the National Academy of Sciences. 100 (22): 12808–12813. Bibcode:2003PNAS..10012808T. doi:10.1073/pnas.2133530100. ISSN 0027-8424. PMC240700. PMID 14555764.
  69. Horwood, M.A.; Eldridge, R.H. (2005). Termites in New South Wales Part 1. Termite biology(PDF) (Technical report). Forest Resources Research. ISSN 0155-7548. 96-38.
  70. Keller, L. (1998). "Queen lifespan and colony characteristics in ants and termites". Insectes Sociaux. 45 (3): 235–246. doi:10.1007/s000400050084. S2CID 24541087.
  71. Srinivasan, Amia (September 10, 2018). "What Termites Can Teach Us". The New Yorker. Archived from the original on March 7, 2020.
  72. Korb, J. (2008). "Termites, hemimetabolous diploid white ants?". Frontiers in Zoology. 5 (1): 15. doi:10.1186/1742-9994-5-15. PMC2564920. PMID 18822181.
  73. Davis, P. "Termite Identification". Entomology at Western Australian Department of Agriculture. Archived from the original on 2009-06-12.
  74. Neoh, K.B.; Lee, C.Y. (2011). "Developmental stages and caste composition of a mature and incipient colony of the drywood termite, Cryptotermes dudleyi (Isoptera: Kalotermitidae)". Journal of Economic Entomology. 104 (2): 622–8. doi:10.1603/ec10346. PMID 21510214. S2CID 23356632.
  75. "Native subterranean termites". University of Florida. Retrieved8 January 2015.
  76. Schneider, M.F. (1999). "Termite Life Cycle and Caste System". University of Freiburg. Retrieved8 January 2015.
  77. Simpson, S.J.; Sword, G.A.; Lo, N. (2011). "Polyphenism in Insects". Current Biology. 21 (18): 738–749. doi:10.1016/j.cub.2011.06.006. PMID 21959164. S2CID 656039.
  78. Tasaki E, Kobayashi K, Matsuura K, Iuchi Y (2017). "An Efficient Antioxidant System in a Long-Lived Termite Queen". PLOS ONE. 12 (1): e0167412. Bibcode:2017PLoSO..1267412T. doi:10.1371/journal.pone.0167412. PMC5226355. PMID 28076409.
  79. Miller, D.M. (5 March 2010). "Subterranean Termite Biology and Behavior". Virginia Tech (Virginia State University). Retrieved8 January 2015.
  80. Gouge, D.H.; Smith, K.A.; Olson, C.; Baker, P. (2001). "Drywood Termites". Cooperative Extension, College of Agriculture & Life Sciences. University of Arizona. Retrieved16 September 2015.
  81. Kaib, M.; Hacker, M.; Brandl, R. (2001). "Egg-laying in monogynous and polygynous colonies of the termite Macrotermes michaelseni (Isoptera, Macrotermitidae)". Insectes Sociaux. 48 (3): 231–237. doi:10.1007/PL00001771. S2CID 35656795.
  82. Gilbert, executive editors, G.A. Kerkut, L.I. (1985). Comprehensive insect physiology, biochemistry, and pharmacology (1st ed.). Oxford: Pergamon Press. p. 167. ISBN 978-0-08-026850-7.{{cite book}}:|first1= has generic name ()
  83. Wyatt, T.D. (2003).Pheromones and animal behaviour: communication by smell and taste (Repr. with corrections 2004. ed.). Cambridge: Cambridge University Press. p. 119. ISBN 978-0-521-48526-5.
  84. Morrow, E.H. (2004). "How the sperm lost its tail: the evolution of aflagellate sperm". Biological Reviews of the Cambridge Philosophical Society. 79 (4): 795–814. doi:10.1017/S1464793104006451. PMID 15682871. S2CID 25878093.
  85. "Supplementary reproductive". University of Hawaii. Archived from the original on 30 October 2014. Retrieved16 September 2015.
  86. Yashiro, T.; Matsuura, K. (2014). "Termite queens close the sperm gates of eggs to switch from sexual to asexual reproduction". Proceedings of the National Academy of Sciences. 111 (48): 17212–17217. Bibcode:2014PNAS..11117212Y. doi:10.1073/pnas.1412481111. PMC4260566. PMID 25404335.
  87. Matsuura, K.; Vargo, E.L.; Kawatsu, K.; Labadie, P. E.; Nakano, H.; Yashiro, T.; Tsuji, K. (2009). "Queen Succession Through Asexual Reproduction in Termites". Science. 323 (5922): 1687. Bibcode:2009Sci...323.1687M. doi:10.1126/science.1169702. PMID 19325106. S2CID 21785886.
  88. Fougeyrollas R, Dolejšová K, Sillam-Dussès D, Roy V, Poteaux C, Hanus R, Roisin Y (June 2015). "Asexual queen succession in the higher termite Embiratermes neotenicus". Proc. Biol. Sci. 282 (1809): 20150260. doi:10.1098/rspb.2015.0260. PMC4590441. PMID 26019158.
  89. Bignell, Roisin & Lo 2010, pp. 13–14.
  90. Freymann, B.P.; Buitenwerf, R.; Desouza, O.; Olff (2008). "The importance of termites (Isoptera) for the recycling of herbivore dung in tropical ecosystems: a review". European Journal of Entomology. 105 (2): 165–173. doi:10.14411/eje.2008.025.
  91. de Souza, O.F.; Brown, V.K. (2009). "Effects of habitat fragmentation on Amazonian termite communities". Journal of Tropical Ecology. 10 (2): 197–206. doi:10.1017/S0266467400007847.
  92. Tokuda, G.; Watanabe, H.; Matsumoto, T.; Noda, H. (1997). "Cellulose digestion in the wood-eating higher termite, Nasutitermes takasagoensis (Shiraki): distribution of cellulases and properties of endo-beta-1,4-glucanase". Zoological Science. 14 (1): 83–93. doi:10.2108/zsj.14.83. PMID 9200983. S2CID 2877588.
  93. Ritter, Michael (2006). The Physical Environment: an Introduction to Physical Geography. University of Wisconsin. p. 450. Archived from the original on 18 May 2007.
  94. Ikeda-Ohtsubo, W.; Brune, A. (2009). "Cospeciation of termite gut flagellates and their bacterial endosymbionts: Trichonympha species and Candidatus Endomicrobium trichonymphae". Molecular Ecology. 18 (2): 332–342. doi:10.1111/j.1365-294X.2008.04029.x. PMID 19192183. S2CID 28048145.
  95. Slaytor, M. (1992). "Cellulose digestion in termites and cockroaches: What role do symbionts play?". Comparative Biochemistry and Physiology B. 103 (4): 775–784. doi:10.1016/0305-0491(92)90194-V.
  96. "The Termite Gut and its Symbiotic Microbes". iBiology. Retrieved2020-05-16.
  97. Watanabe, H..; Noda, H.; Tokuda, G.; Lo, N. (1998). "A cellulase gene of termite origin". Nature. 394 (6691): 330–331. Bibcode:1998Natur.394..330W. doi:10.1038/28527. PMID 9690469. S2CID 4384555.
  98. Tokuda, G.; Watanabe, H. (2007). "Hidden cellulases in termites: revision of an old hypothesis". Biology Letters. 3 (3): 336–339. doi:10.1098/rsbl.2007.0073. PMC2464699. PMID 17374589.
  99. Li, Z.-Q.; Liu, B.-R.; Zeng, W.-H.; Xiao, W.-L.; Li, Q.-J.; Zhong, J.-H. (2013). "Character of Cellulase Activity in the Guts of Flagellate-Free Termites with Different Feeding Habits". Journal of Insect Science. 13 (37): 37. doi:10.1673/031.013.3701. PMC3738099. PMID 23895662.
  100. "The Termite Gut and its Symbiotic Microbes". iBiology. Retrieved2020-05-16.
  101. Geetha Iyer Scroll.in (Mar 09, 2017) Why Indians worship the mound of the much-hated termite "[The soldier termites] and the reproductive castes obtain their nutrients from the workers through oral or anal trophallaxis."
  102. Dietrich, C.; Kohler, T.; Brune, A. (2014). "The Cockroach origin of the termite gut microbiota: patterns in bacterial community structure reflect major evolutionary events". Applied and Environmental Microbiology. 80 (7): 2261–2269. Bibcode:2014ApEnM..80.2261D. doi:10.1128/AEM.04206-13. PMC3993134. PMID 24487532.
  103. Allen, C.T.; Foster, D.E.; Ueckert, D.N. (1980). "Seasonal Food Habits of a Desert Termite, Gnathamitermes tubiformans, in West Texas". Environmental Entomology. 9 (4): 461–466. doi:10.1093/ee/9.4.461.
  104. McMahan, E.A. (1966). "Studies of Termite Wood-feeding Preferences"(PDF). Hawaiian Entomological Society. 19 (2): 239–250. ISSN 0073-134X.
  105. Aanen, D.K.; Eggleton, P.; Rouland-Lefevre, C.; Guldberg-Froslev, T.; Rosendahl, S.; Boomsma, J.J. (2002). "The evolution of fungus-growing termites and their mutualistic fungal symbionts". Proceedings of the National Academy of Sciences. 99 (23): 14887–14892. Bibcode:2002PNAS...9914887A. doi:10.1073/pnas.222313099. JSTOR 3073687. PMC137514. PMID 12386341.
  106. Mueller, U.G.; Gerardo, N. (2002). "Fungus-farming insects: Multiple origins and diverse evolutionary histories". Proceedings of the National Academy of Sciences. 99 (24): 15247–15249. Bibcode:2002PNAS...9915247M. doi:10.1073/pnas.242594799. PMC137700. PMID 12438688.
  107. Roberts, E.M.; Todd, C.N.; Aanen, D.K.; Nobre, T.; Hilbert-Wolf, H.L.; O'Connor, P.M.; Tapanila, L.; Mtelela, C.; Stevens, N.J. (2016). "Oligocene termite nests with in situ fungus gardens from the Rukwa Rift Basin, Tanzania, support a paleogene African origin for insect agriculture". PLOS ONE. 11 (6): e0156847. Bibcode:2016PLoSO..1156847R. doi:10.1371/journal.pone.0156847. PMC4917219. PMID 27333288.
  108. Radek, R. (1999). "Flagellates, bacteria, and fungi associated with termites: diversity and function in nutrition – a review"(PDF). Ecotropica. 5: 183–196.
  109. Breznak, J.A.; Brune, A. (1993). "Role of microorganisms in the digestion of lignocellulose by termites". Annual Review of Entomology. 39 (1): 453–487. doi:10.1146/annurev.en.39.010194.002321.
  110. Kok, O.B.; Hewitt, P.H. (1990). "Bird and mammal predators of the harvester termite Hodotermes mossambicus (Hagen) in semi-arid regions of South Africa". South African Journal of Science. 86 (1): 34–37. ISSN 0038-2353.
  111. Hölldobler, B.; Wilson, E.O. (1990). The Ants. Cambridge, Massachusetts: Belknap Press of Harvard University Press. pp. 559–566. ISBN 978-0-674-04075-5.
  112. Culliney, T.W.; Grace, J.K. (2000). "Prospects for the biological control of subterranean termites (Isoptera: Rhinotermitidae), with special reference to Coptotermes formosanus". Bulletin of Entomological Research. 90 (1): 9–21. doi:10.1017/S0007485300000663. PMID 10948359.
  113. Dean, W.R.J.; Milton, S.J. (1995). "Plant and invertebrate assemblages on old fields in the arid southern Karoo, South Africa". African Journal of Ecology. 33 (1): 1–13. doi:10.1111/j.1365-2028.1995.tb00777.x.
  114. Wade, W.W. (2002). Ecology of Desert Systems. Burlington: Elsevier. p. 216. ISBN 978-0-08-050499-5.
  115. Reagan, D.P.; Waide, R.B. (1996). The food web of a tropical rain forest. Chicago: University of Chicago Press. p. 294. ISBN 978-0-226-70599-6.
  116. Bardgett, R.D.; Herrick, J.E.; Six, J.; Jones, T.H.; Strong, D.R.; van der Putten, W.H. (2013). Soil ecology and ecosystem services (1st ed.). Oxford: Oxford University Press. p. 178. ISBN 978-0-19-968816-6.
  117. Bignell, Roisin & Lo 2010, p. 509.
  118. Choe, J.C.; Crespi, B.J. (1997).The evolution of social behavior in insects and arachnids (1st ed.). Cambridge: Cambridge university press. p. 76. ISBN 978-0-521-58977-2.
  119. Abe, Y.; Bignell, D.E.; Higashi, T. (2014). Termites: Evolution, Sociality, Symbioses, Ecology. Springer. pp. 124–149. doi:10.1007/978-94-017-3223-9. ISBN 978-94-017-3223-9. S2CID 30804981.
  120. Wilson, D.S.; Clark, A.B. (1977). "Above ground defence in the harvester termite, Hodotermes mossambicus". Journal of the Entomological Society of South Africa. 40: 271–282.
  121. Lavelle, P.; Spain, A.V. (2001).Soil ecology (2nd ed.). Dordrecht: Kluwer Academic. p. 316. ISBN 978-0-306-48162-8.
  122. Richardson, P.K.R.; Bearder, S.K. (1984)."The Hyena Family". In MacDonald, D. (ed.). The Encyclopedia of Mammals. New York, NY: Facts on File Publication. pp. 158–159. ISBN 978-0-87196-871-5.
  123. Mills, G.; Harvey, M. (2001). African Predators. Washington, D.C.: Smithsonian Institution Press. p. 71. ISBN 978-1-56098-096-4.
  124. d'Errico, F.; Backwell, L. (2009). "Assessing the function of early hominin bone tools". Journal of Archaeological Science. 36 (8): 1764–1773. doi:10.1016/j.jas.2009.04.005.
  125. Lepage, M.G. (1981). "Étude de la prédation de Megaponera foetens (F.) sur les populations récoltantes de Macrotermitinae dans un ecosystème semi-aride (Kajiado-Kenya)". Insectes Sociaux (in Spanish). 28 (3): 247–262. doi:10.1007/BF02223627. S2CID 28763771.
  126. Levieux, J. (1966). "Note préliminaire sur les colonnes de chasse de Megaponera fœtens F. (Hyménoptère Formicidæ)". Insectes Sociaux (in French). 13 (2): 117–126. doi:10.1007/BF02223567. S2CID 2031222.
  127. Longhurst, C.; Baker, R.; Howse, P.E. (1979). "Chemical crypsis in predatory ants". Experientia. 35 (7): 870–872. doi:10.1007/BF01955119. S2CID 39854106.
  128. Wheeler, W.M. (1936). "Ecological relations of Ponerine and other ants to termites". Proceedings of the American Academy of Arts and Sciences. 71 (3): 159–171. doi:10.2307/20023221. JSTOR 20023221.
  129. Shattuck, S.O.; Heterick, B.E. (2011). "Revision of the ant genus Iridomyrmex (Hymenoptera : Formicidae)"(PDF). Zootaxa. 2845: 1–74. doi:10.11646/zootaxa.2743.1.1. ISBN 978-1-86977-676-3. ISSN 1175-5334.
  130. Traniello, J.F.A. (1981). "Enemy deterrence in the recruitment strategy of a termite: Soldier-organized foraging in Nasutitermes costalis". Proceedings of the National Academy of Sciences. 78 (3): 1976–1979. Bibcode:1981PNAS...78.1976T. doi:10.1073/pnas.78.3.1976. PMC319259. PMID 16592995.
  131. Schöning, C.; Moffett, M.W. (2007). "Driver Ants Invading a Termite Nest: why do the most catholic predators of all seldom take this abundant prey?"(PDF). Biotropica. 39 (5): 663–667. doi:10.1111/j.1744-7429.2007.00296.x. Archived from the original(PDF) on 2015-11-12. Retrieved2015-09-20.
  132. Mill, A.E. (1983). "Observations on Brazilian termite alate swarms and some structures used in the dispersal of reproductives (Isoptera: Termitidae)". Journal of Natural History. 17 (3): 309–320. doi:10.1080/00222938300770231.
  133. Schmid-Hempel 1998, p. 61.
  134. Schmid-Hempel 1998, p. 75.
  135. Wilson, E.O. (1971). The Insect Societies. Vol. 76 (5th ed.). Cambridge, Massachusetts: Belknap Press of Harvard University Press. p. 398. ISBN 978-0-674-45495-8.
  136. Schmid-Hempel 1998, p. 59.
  137. Schmid-Hempel 1998, pp. 301–302.
  138. Schmid-Hempel 1998, p. 19.
  139. Weiser, J.; Hrdy, I. (2009). "Pyemotes – mites as parasites of termites". Zeitschrift für Angewandte Entomologie. 51 (1–4): 94–97. doi:10.1111/j.1439-0418.1962.tb04062.x.
  140. Chouvenc, T.; Efstathion, C.A.; Elliott, M.L.; Su, N.Y. (2012). "Resource competition between two fungal parasites in subterranean termites". Die Naturwissenschaften. 99 (11): 949–58. Bibcode:2012NW.....99..949C. doi:10.1007/s00114-012-0977-2. PMID 23086391. S2CID 16393629.
  141. Schmid-Hempel 1998, pp. 38, 102.
  142. Wilson, Megan; Barden, Phillip; Ware, Jessica (2021-04-30). "A Review of Ectoparasitic Fungi Associated With Termites". Annals of the Entomological Society of America. 114 (4): 373–396. doi:10.1093/aesa/saab001. Retrieved7 May 2021.
  143. Chouvenc, T.; Mullins, A.J.; Efstathion, C.A.; Su, N.-Y. (2013). "Virus-like symptoms in a termite (Isoptera: Kalotermitidae) field colony". Florida Entomologist. 96 (4): 1612–1614. doi:10.1653/024.096.0450. S2CID 73570814.
  144. Al Fazairy, A.A.; Hassan, F.A. (2011). "Infection of Termites by Spodoptera littoralis Nuclear Polyhedrosis Virus". International Journal of Tropical Insect Science. 9 (1): 37–39. doi:10.1017/S1742758400009991. S2CID 84743428.
  145. Traniello, J.F.A.; Leuthold, R.H. (2000). Behavior and Ecology of Foraging in Termites. Springer Netherlands. pp. 141–168. doi:10.1007/978-94-017-3223-9_7. ISBN 978-94-017-3223-9.
  146. Reinhard, J.; Kaib, M. (2001). "Trail communication during foraging and recruitment in the subterranean termite Reticulitermes santonensis De Feytaud (Isoptera, Rhinotermitidae)". Journal of Insect Behavior. 14 (2): 157–171. doi:10.1023/A:1007881510237. S2CID 40887791.
  147. Costa-Leonardo, A.M.; Haifig, I. (2013). Termite communication during different behavioral activities in Biocommunication of Animals. Springer Netherlands. pp. 161–190. doi:10.1007/978-94-007-7414-8_10. ISBN 978-94-007-7413-1.
  148. Costa-Leonardo, A.M. (2006). "Morphology of the sternal gland in workers of Coptotermes gestroi (Isoptera, Rhinotermitidae)". Micron. 37 (6): 551–556. doi:10.1016/j.micron.2005.12.006. PMID 16458523.
  149. Traniello, J.F.; Busher, C. (1985). "Chemical regulation of polyethism during foraging in the neotropical termite Nasutitermes costalis". Journal of Chemical Ecology. 11 (3): 319–32. doi:10.1007/BF01411418. PMID 24309963. S2CID 27799126.
  150. Miramontes, O.; DeSouza, O.; Paiva, L.R.; Marins, A.; Orozco, S.; Aegerter, C.M. (2014). "Lévy flights and self-similar exploratory behaviour of termite workers: beyond model fitting". PLOS ONE. 9 (10): e111183. arXiv:1410.0930. Bibcode:2014PLoSO...9k1183M. doi:10.1371/journal.pone.0111183. PMC4213025. PMID 25353958.
  151. Jost, C.; Haifig, I.; de Camargo-Dietrich, C.R.R.; Costa-Leonardo, A.M. (2012). "A comparative tunnelling network approach to assess interspecific competition effects in termites". Insectes Sociaux. 59 (3): 369–379. doi:10.1007/s00040-012-0229-7. S2CID 14885485.
  152. Polizzi, J.M.; Forschler, B.T. (1998). "Intra- and interspecific agonism in Reticulitermes flavipes (Kollar) and R. virginicus (Banks) and effects of arena and group size in laboratory assays". Insectes Sociaux. 45 (1): 43–49. doi:10.1007/s000400050067. S2CID 36235510.
  153. Darlington, J.P.E.C. (1982). "The underground passages and storage pits used in foraging by a nest of the termite Macrotermes michaelseni in Kajiado, Kenya". Journal of Zoology. 198 (2): 237–247. doi:10.1111/j.1469-7998.1982.tb02073.x.
  154. Cornelius, M.L.; Osbrink, W.L. (2010). "Effect of soil type and moisture availability on the foraging behavior of the Formosan subterranean termite (Isoptera: Rhinotermitidae)". Journal of Economic Entomology. 103 (3): 799–807. doi:10.1603/EC09250. PMID 20568626. S2CID 23173060.
  155. Toledo Lima, J.; Costa-Leonardo, A.M. (2012). "Subterranean termites (Isoptera: Rhinotermitidae): Exploitation of equivalent food resources with different forms of placement". Insect Science. 19 (3): 412–418. doi:10.1111/j.1744-7917.2011.01453.x. S2CID 82046133.
  156. Jmhasly, P.; Leuthold, R.H. (1999). "Intraspecific colony recognition in the termites Macrotermes subhyalinus and Macrotermes bellicosus (Isoptera, Termitidae)". Insectes Sociaux. 46 (2): 164–170. doi:10.1007/s000400050128. S2CID 23037986.
  157. Messenger, M.T.; Su, N.Y. (2005). "Agonistic behavior between colonies of the Formosan subterranean termite (Isoptera: Rhinotermitidae) from Louis Armstrong Park, New Orleans, Louisiana". Sociobiology. 45 (2): 331–345.
  158. Korb, J.; Weil, T.; Hoffmann, K.; Foster, K.R.; Rehli, M. (2009). "A gene necessary for reproductive suppression in termites". Science. 324 (5928): 758. Bibcode:2009Sci...324..758K. doi:10.1126/science.1170660. PMID 19423819. S2CID 31608071.
  159. Mathew, T.T.G.; Reis, R.; DeSouza, O.; Ribeiro, S.P. (2005). "Predation and interference competition between ants (Hymenoptera: Formicidae) and arboreal termites (Isoptera: Termitidae)"(PDF). Sociobiology. 46 (2): 409–419.
  160. Evans, T.A.; Inta, R.; Lai, J.C.S.; Lenz, M. (2007). "Foraging vibration signals attract foragers and identify food size in the drywood termite, Cryptotermes secundus". Insectes Sociaux. 54 (4): 374–382. doi:10.1007/s00040-007-0958-1. S2CID 40214049.
  161. Costa-Leonardo, A.M.; Casarin, F.E.; Lima, J.T. (2009). "Chemical communication in isoptera". Neotropical Entomology. 38 (1): 747–52. doi:10.1590/S1519-566X2009000100001. PMID 19347093.
  162. Richard, F.-J.; Hunt, J.H. (2013). "Intracolony chemical communication in social insects"(PDF). Insectes Sociaux. 60 (3): 275–291. doi:10.1007/s00040-013-0306-6. S2CID 8108234. Archived from the original(PDF) on 2016-03-04. Retrieved2015-10-08.
  163. Dronnet, S.; Lohou, C.; Christides, J.P.; Bagnères, A.G. (2006). "Cuticular hydrocarbon composition reflects genetic relationship among colonies of the introduced termite Reticulitermes santonensis Feytaud". Journal of Chemical Ecology. 32 (5): 1027–1042. doi:10.1007/s10886-006-9043-x. PMID 16739021. S2CID 23956394.
  164. Rosengaus, R.B.; Traniello, J.F.A.; Chen, T.; Brown, J.J.; Karp, R.D. (1999). "Immunity in a social insect". Naturwissenschaften. 86 (12): 588–591. Bibcode:1999NW.....86..588R. doi:10.1007/s001140050679. S2CID 10769345.
  165. Wilson, D.S. (1977). "Above ground predator defense in the harvester termite, Hodotermes mossambicus (Hagen)". Journal of the Entomological Society of Southern Africa. 40: 271–282.
  166. Belbin, R.M. (2013). The Coming Shape of Organization. New York: Routledge. p. 27. ISBN 978-1-136-01553-3.
  167. Wilson, E.O. (2014). A window on eternity: a biologist's walk through Gorongosa National Park (First ed.). Simon & Schuster. pp. 85, 90. ISBN 978-1-4767-4741-5.
  168. Miura, T.; Matsumoto, T. (2000). "Soldier morphogenesis in a nasute termite: discovery of a disc-like structure forming a soldier nasus". Proceedings of the Royal Society B: Biological Sciences. 267 (1449): 1185–1189. doi:10.1098/rspb.2000.1127. PMC1690655. PMID 10902684.
  169. Prestwich, G.D.; Chen, D. (1981). "Soldier defense secretions of Trinervitermes bettonianus (Isoptera, Nasutitermitinae): Chemical variation in allopatric populations". Journal of Chemical Ecology. 7 (1): 147–157. doi:10.1007/BF00988642. PMID 24420434. S2CID 27654745.
  170. Chen, J.; Henderson, G.; Grimm, C. C.; Lloyd, S. W.; Laine, R. A. (1998-04-09). "Termites fumigate their nests with naphthalene". Nature. 392 (6676): 558–559. Bibcode:1998Natur.392..558C. doi:10.1038/33305. S2CID 4419882.
  171. Piper, Ross (2007), Extraordinary Animals: An Encyclopedia of Curious and Unusual Animals, Greenwood Press, p. 26, ISBN 978-0-313-33922-6
  172. Bordereau, C.; Robert, A.; Van Tuyen, V.; Peppuy, A. (1997). "Suicidal defensive behaviour by frontal gland dehiscence in Globitermes sulphureus Haviland soldiers (Isoptera)". Insectes Sociaux. 44 (3): 289–297. doi:10.1007/s000400050049. S2CID 19770804.
  173. Sobotnik, J.; Bourguignon, T.; Hanus, R.; Demianova, Z.; Pytelkova, J.; Mares, M.; Foltynova, P.; Preisler, J.; Cvacka, J.; Krasulova, J.; Roisin, Y. (2012). "Explosive backpacks in old termite workers". Science. 337 (6093): 436. Bibcode:2012Sci...337..436S. doi:10.1126/science.1219129. PMID 22837520. S2CID 206540025.
  174. ŠobotnÍk, J.; Bourguignon, T.; Hanus, R.; Weyda, F.; Roisin, Y. (2010). "Structure and function of defensive glands in soldiers of Glossotermes oculatus (Isoptera: Serritermitidae)". Biological Journal of the Linnean Society. 99 (4): 839–848. doi:10.1111/j.1095-8312.2010.01392.x.
  175. Ulyshen, M.D.; Shelton, T.G. (2011). "Evidence of cue synergism in termite corpse response behavior". Naturwissenschaften. 99 (2): 89–93. Bibcode:2012NW.....99...89U. doi:10.1007/s00114-011-0871-3. PMID 22167071. S2CID 2616753.
  176. Su, N.Y. (2005). "Response of the Formosan subterranean termites (Isoptera: Rhinotermitidae) to baits or nonrepellent termiticides in extended foraging arenas". Journal of Economic Entomology. 98 (6): 2143–2152. doi:10.1603/0022-0493-98.6.2143. PMID 16539144. S2CID 196618597.
  177. Sun, Q.; Haynes, K.F.; Zhou, X. (2013). "Differential undertaking response of a lower termite to congeneric and conspecific corpses". Scientific Reports. 3: 1650. Bibcode:2013NatSR...3E1650S. doi:10.1038/srep01650. PMC3629736. PMID 23598990.
  178. Neoh, K.-B.; Yeap, B.-K.; Tsunoda, K.; Yoshimura, T.; Lee, C.Y.; Korb, J. (2012). "Do termites avoid carcasses? behavioral responses depend on the nature of the carcasses". PLOS ONE. 7 (4): e36375. Bibcode:2012PLoSO...736375N. doi:10.1371/journal.pone.0036375. PMC3338677. PMID 22558452.
  179. Matsuura, K. (2006). "Termite-egg mimicry by a sclerotium-forming fungus". Proceedings of the Royal Society B: Biological Sciences. 273 (1591): 1203–1209. doi:10.1098/rspb.2005.3434. PMC1560272. PMID 16720392.
  180. Matsuura, K.; Yashiro, T.; Shimizu, K.; Tatsumi, S.; Tamura, T. (2009). "Cuckoo fungus mimics termite eggs by producing the cellulose-digesting enzyme β-glucosidase". Current Biology. 19 (1): 30–36. doi:10.1016/j.cub.2008.11.030. PMID 19110429. S2CID 18604426.
  181. Howard, R.W.; McDaniel, C.A.; Blomquist, G.J. (1980). "Chemical mimicry as an integrating mechanism: cuticular hydrocarbons of a termitophile and its host". Science. 210 (4468): 431–433. Bibcode:1980Sci...210..431H. doi:10.1126/science.210.4468.431. PMID 17837424. S2CID 33221252.
  182. Watson, J.A.L. (1973). "Austrospirachtha mimetes a new termitophilous corotocine from Northern Australia (Coleoptera: Staphylinidae)". Australian Journal of Entomology. 12 (4): 307–310. doi:10.1111/j.1440-6055.1973.tb01678.x.
  183. Forbes, H.O. (1878). "Termites Kept in Captivity by Ants". Nature. 19 (471): 4–5. Bibcode:1878Natur..19....4F. doi:10.1038/019004b0. S2CID 4125839.(subscription required)
  184. Darlington, J. (1985). "Attacks by doryline ants and termite nest defences (Hymenoptera; Formicidae; Isoptera; Termitidae)". Sociobiology. 11: 189–200.
  185. Quinet Y, Tekule N & de Biseau JC (2005). "Behavioural Interactions Between Crematogaster brevispinosa rochai Forel (Hymenoptera: Formicidae) and Two Nasutitermes Species (Isoptera: Termitidae)". Journal of Insect Behavior. 18 (1): 1–17. doi:10.1007/s10905-005-9343-y. S2CID 33487814.
  186. Coty, D.; Aria, C.; Garrouste, R.; Wils, P.; Legendre, F.; Nel, A.; Korb, J. (2014). "The First Ant-Termite Syninclusion in Amber with CT-Scan Analysis of Taphonomy". PLOS ONE. 9 (8): e104410. Bibcode:2014PLoSO...9j4410C. doi:10.1371/journal.pone.0104410. PMC4139309. PMID 25140873.
  187. Santos, P.P.; Vasconcellos, A.; Jahyny, B.; Delabie, J.H.C. (2010). "Ant fauna (Hymenoptera, Formicidae) associated to arboreal nests of Nasutitermes spp: (Isoptera, Termitidae) in a cacao plantation in southeastern Bahia, Brazil". Revista Brasileira de Entomologia. 54 (3): 450–454. doi:10.1590/S0085-56262010000300016.
  188. Jaffe, K.; Ramos, C.; Issa, S. (1995). "Trophic Interactions Between Ants and Termites that Share Common Nests". Annals of the Entomological Society of America. 88 (3): 328–333. doi:10.1093/aesa/88.3.328.
  189. Trager, J.C. (1991). "A Revision of the fire ants, Solenopsis geminata group (Hymenoptera: Formicidae: Myrmicinae)". Journal of the New York Entomological Society. 99 (2): 141–198. doi:10.5281/zenodo.24912. JSTOR 25009890.
  190. Cingel, N.A. van der (2001). An atlas of orchid pollination: America, Africa, Asia and Australia. Rotterdam: Balkema. p. 224. ISBN 978-90-5410-486-5.
  191. McHatton, R. (2011). "Orchid Pollination: exploring a fascinating world"(PDF). The American Orchid Society. p. 344. Retrieved5 September 2015.
  192. Cowie, R. (2014). Journey to a Waterfall a biologist in Africa. Raleigh, North Carolina: Lulu Press. p. 169. ISBN 978-1-304-66939-1.
  193. Tan, K.H. (2009). Environmental Soil Science (3rd ed.). Boca Raton, Florida: CRC Press. pp. 105–106. ISBN 978-1-4398-9501-6.
  194. Clark, Sarah (15 November 2005). "Plant extract stops termites dead". ABC. Archived from the original on 15 June 2009. Retrieved8 February 2014.
  195. Vasconcellos, Alexandre; Bandeira, Adelmar G.; Moura, Flávia Maria S.; Araújo, Virgínia Farias P.; Gusmão, Maria Avany B.; Reginaldo, Constantino (February 2010). "Termite assemblages in three habitats under different disturbance regimes in the semi-arid Caatinga of NE Brazil". Journal of Arid Environments. Elsevier. 74 (2): 298–302. Bibcode:2010JArEn..74..298V. doi:10.1016/j.jaridenv.2009.07.007. ISSN 0140-1963.
  196. Bignell, Roisin & Lo 2010, p. 3.
  197. Noirot, C.; Darlington, J.P.E.C. (2000). Termite Nests: Architecture, Regulation and Defence in Termites: Evolution, Sociality, Symbioses, Ecology. Springer. pp. 121–139. doi:10.1007/978-94-017-3223-9_6. ISBN 978-94-017-3223-9.
  198. Bignell, Roisin & Lo 2010, p. 20.
  199. Eggleton, P.; Bignell, D.E.; Sands, W.A.; Mawdsley, N. A.; Lawton, J. H.; Wood, T.G.; Bignell, N.C. (1996). "The Diversity, Abundance and Biomass of Termites under Differing Levels of Disturbance in the Mbalmayo Forest Reserve, Southern Cameroon". Philosophical Transactions of the Royal Society B: Biological Sciences. 351 (1335): 51–68. Bibcode:1996RSPTB.351...51E. doi:10.1098/rstb.1996.0004.
  200. Bignell, Roisin & Lo 2010, p. 21.
  201. De Visse, S.N.; Freymann, B.P.; Schnyder, H. (2008). "Trophic interactions among invertebrates in termitaria in the African savanna: a stable isotope approach". Ecological Entomology. 33 (6): 758–764. doi:10.1111/j.1365-2311.2008.01029.x. S2CID 33877331.
  202. Bignell, Roisin & Lo 2010, p. 22.
  203. Vane, C.H.; Kim, A.W.; Moss-Hayes, V.; Snape, C.E.; Diaz, M.C.; Khan, N.S.; Engelhart, S.E.; Horton, B.P. (2013). "Degradation of mangrove tissues by arboreal termites (Nasutitermes acajutlae) and their role in the mangrove C cycle (Puerto Rico): Chemical characterization and organic matter provenance using bulk δ13C, C/N, alkaline CuO oxidation-GC/MS, and solid-state"(PDF). Geochemistry, Geophysics, Geosystems. 14 (8): 3176–3191. Bibcode:2013GGG....14.3176V. doi:10.1002/ggge.20194.
  204. Roisin, Y.; Pasteels, J. M. (1986). "Reproductive mechanisms in termites: Polycalism and polygyny in Nasutitermes polygynus and N. costalis". Insectes Sociaux. 33 (2): 149–167. doi:10.1007/BF02224595. S2CID 41799894.
  205. Perna, A.; Jost, C.; Couturier, E.; Valverde, S.; Douady, S.; Theraulaz, G. (2008). "The structure of gallery networks in the nests of termite Cubitermes spp. revealed by X-ray tomography". Die Naturwissenschaften. 95 (9): 877–884. Bibcode:2008NW.....95..877P. doi:10.1007/s00114-008-0388-6. PMID 18493731. S2CID 15326313.
  206. Glenday, Craig (2014). Guinness World Records 2014. pp. 33. ISBN 978-1-908843-15-9.
  207. Jacklyn, P. (1991). "Evidence for Adaptive Variation in the Orientation of Amitermes (Isoptera, Termitinae) Mounds From Northern Australia". Australian Journal of Zoology. 39 (5): 569. doi:10.1071/ZO9910569.
  208. Jacklyn, P.M.; Munro, U. (2002). "Evidence for the use of magnetic cues in mound construction by the termite Amitermes meridionalis (Isoptera : Termitinae)". Australian Journal of Zoology. 50 (4): 357. doi:10.1071/ZO01061.
  209. Grigg, G.C. (1973). "Some Consequences of the Shape and Orientation of 'magnetic' Termite Mounds"(PDF). Australian Journal of Zoology. 21 (2): 231–237. doi:10.1071/ZO9730231.
  210. Hadlington, P. (1996). Australian Termites and Other Common Timber Pests (2nd ed.). Kensington, NSW, Australia: New South Wales University Press. pp. 28–30. ISBN 978-0-86840-399-1.
  211. Kahn, L.; Easton, B. (2010). Shelter II. Bolinas, California: Shelter Publications. p. 198. ISBN 978-0-936070-49-0.
  212. Su, N.Y.; Scheffrahn, R.H. (2000). Termites as Pests of Buildings in Termites: Evolution, Sociality, Symbioses, Ecology. Springer Netherlands. pp. 437–453. doi:10.1007/978-94-017-3223-9_20. ISBN 978-94-017-3223-9.
  213. Thorne, Ph.D, Barbara L. (1999). NPMA Research Report On Subterranean Termites. Dunn Loring, VA: NPMA. p. 22.
  214. "Termites". Victorian Building Authority. Government of Victoria. 2014. Archived from the original on 3 February 2018. Retrieved20 September 2015.
  215. Thorne, Ph.D, Barbara L. (1999). NPMA Research Report On Subterranean Termites. Dunn Loring, VA: NPMA. p. 2.
  216. Grace, J.K.; Cutten, G.M.; Scheffrahn, R.H.; McEkevan, D.K. (1991). "First infestation by Incisitermes minor of a Canadian building (Isoptera: Kalotermitidae)". Sociobiology. 18: 299–304.
  217. Sands, W.A. (1973). "Termites as Pests of Tropical Food Crops". Tropical Pest Management. 19 (2): 167–177. doi:10.1080/09670877309412751.
  218. Flores, A. (17 February 2010). "New Assay Helps Track Termites, Other Insects". Agricultural Research Service. United States Department of Agriculture. Retrieved15 January 2015.
  219. Su, N.Y.; Scheffrahn, R.H. (1990). "Economically important termites in the United States and their control"(PDF). Sociobiology. 17: 77–94. Archived from the original(PDF) on 2011-08-12.
  220. Thorne, Ph.D, Barbara L. (1999). NPMA Research Report On Subterranean Termites. Dunn Loring, VA: NPMA. p. 40.
  221. Elliott, Sara (26 May 2009). "How can copper keep termites at bay?". HowStuffWorks.
  222. "Questions and Answers About Termites"(PDF). Department of Consumer Affairs, Structural Pest Control Board of California. Retrieved19 April 2021.
  223. "EPA Registration and Label for Taurus SC Termiticide"(PDF). EPA.gov.
  224. "EPA Registration and Label for Termidor SC"(PDF). EPA.gov. Retrieved19 April 2021.
  225. Pidd, Helen (21 December 2021). "'A world first': Devon calls victory in 27-year war on termites". The Guardian. Archived from the original on 22 December 2021. Retrieved22 December 2021.
  226. Figueirêdo, R.E.C.R.; Vasconcellos, A.; Policarpo, I.S.; Alves, R.R.N. (2015). "Edible and medicinal termites: a global overview". Journal of Ethnobiology and Ethnomedicine. 11 (1): 29. doi:10.1186/s13002-015-0016-4. PMC4427943. PMID 25925503.
  227. Nyakupfuka, A. (2013). Global Delicacies: Discover Missing Links from Ancient Hawaiian Teachings to Clean the Plaque of your Soul and Reach Your Higher Self. Bloomington, Indiana: BalboaPress. pp. 40–41. ISBN 978-1-4525-6791-4.
  228. Bodenheimer, F.S. (1951). Insects as Human Food: A Chapter of the Ecology of Man. Netherlands: Springer. pp. 331–350. ISBN 978-94-017-6159-8.
  229. Geissler, P.W. (2011). "The significance of earth-eating: social and cultural aspects of geophagy among Luo children". Africa. 70 (4): 653–682. doi:10.3366/afr.2000.70.4.653. S2CID 145754470.
  230. Knudsen, J.W. (2002). "Akula udongo (earth eating habit): a social and cultural practice among Chagga women on the slopes of Mount Kilimanjaro". African Journal of Indigenous Knowledge Systems. 1 (1): 19–26. doi:10.4314/indilinga.v1i1.26322. ISSN 1683-0296. OCLC 145403765.
  231. Nchito, M.; Wenzel Geissler, P.; Mubila, L.; Friis, H.; Olsen, A. (2004). "Effects of iron and multimicronutrient supplementation on geophagy: a two-by-two factorial study among Zambian schoolchildren in Lusaka". Transactions of the Royal Society of Tropical Medicine and Hygiene. 98 (4): 218–227. doi:10.1016/S0035-9203(03)00045-2. PMID 15049460.
  232. Saathoff, E.; Olsen, A.; Kvalsvig, J.D.; Geissler, P.W. (2002). "Geophagy and its association with geohelminth infection in rural schoolchildren from northern KwaZulu-Natal, South Africa". Transactions of the Royal Society of Tropical Medicine and Hygiene. 96 (5): 485–490. doi:10.1016/S0035-9203(02)90413-X. PMID 12474473.
  233. Katayama, N.; Ishikawa, Y.; Takaoki, M.; Yamashita, M.; Nakayama, S.; Kiguchi, K.; Kok, R.; Wada, H.; Mitsuhashi, J. (2008). "Entomophagy: A key to space agriculture"(PDF). Advances in Space Research. 41 (5): 701–705. Bibcode:2008AdSpR..41..701S. doi:10.1016/j.asr.2007.01.027.
  234. Mitchell, J.D. (2002). "Termites as pests of crops, forestry, rangeland and structures in Southern Africa and their control". Sociobiology. 40 (1): 47–69. ISSN 0361-6525.
  235. Löffler, E.; Kubiniok, J. (1996). "Landform development and bioturbation on the Khorat plateau, Northeast Thailand"(PDF). Natural History Bulletin of the Siam Society. 44: 199–216.
  236. Evans, T.A.; Dawes, T.Z.; Ward, P.R.; Lo, N. (2011). "Ants and termites increase crop yield in a dry climate". Nature Communications. 2: 262. Bibcode:2011NatCo...2..262E. doi:10.1038/ncomms1257. PMC3072065. PMID 21448161.
  237. "Termite Power". DOE Joint Genome Institute. United States Department of Energy. 14 August 2006. Archived from the original on 22 September 2006. Retrieved11 September 2015.{{cite web}}: CS1 maint: unfit URL (link)
  238. Hirschler, B. (22 November 2007). "Termites' gut reaction set for biofuels". ABC News. Retrieved8 January 2015.
  239. Roach, J. (14 March 2006). "Termite Power: Can Pests' Guts Create New Fuel?". National Geographic News. Retrieved11 September 2015.
  240. Werfel, J.; Petersen, K.; Nagpal, R. (2014). "Designing Collective Behavior in a Termite-Inspired Robot Construction Team". Science. 343 (6172): 754–758. Bibcode:2014Sci...343..754W. doi:10.1126/science.1245842. PMID 24531967. S2CID 38776920.
  241. Gibney, E. (2014). "Termite-inspired robots build castles". Nature. doi:10.1038/nature.2014.14713. S2CID 112117767.
  242. "Termites Green Architecture in the Tropics". The Architect. Architectural Association of Kenya. Archived from the original on 22 March 2016. Retrieved17 October 2015.
  243. Tan, A.; Wong, N. (2013). "Parameterization Studies of Solar Chimneys in the Tropics". Energies. 6 (1): 145–163. doi:10.3390/en6010145.
  244. Tsoroti, S. (15 May 2014). "What's that building? Eastgate Mall". Harare News. Archived from the original on 11 April 2021. Retrieved8 January 2015.
  245. "Im Zoo Basel fliegen die Termiten aus". Neue Zürcher Zeitung (in German). 8 February 2014. Retrieved21 May 2011.
  246. Van-Huis, H. (2003). "Insects as food in Sub-Saharan Africa"(PDF). Insect Science and Its Application. 23 (3): 163–185. doi:10.1017/s1742758400023572.
  247. Neoh, K.B. (2013). "Termites and human society in Southeast Asia"(PDF). The Newsletter. 30 (66): 1–2.

Cited literature

Termite Article Talk Language Watch Edit This article is about social insects For other uses see Termite disambiguation Termites are eusocial insects that are classified at the taxonomic rank of infraorder Isoptera or alternatively as epifamily Termitoidae within the order Blattodea along with cockroaches Termites were once classified in a separate order from cockroaches but recent phylogenetic studies indicate that they evolved from cockroaches as they are deeply nested within the group and the sister group to wood eating cockroaches of the genus Cryptocercus Previous estimates suggested the divergence took place during the Jurassic or Triassic More recent estimates suggest that they have an origin during the Late Jurassic 3 with the first fossil records in the Early Cretaceous About 3 106 species are currently described with a few hundred more left to be described Although these insects are often called white ants 4 they are not ants and are not closely related to ants Termite Temporal range Early Cretaceous Recent PreꞒ Ꞓ O S D C P T J K Pg NFormosan subterranean termite Coptotermes formosanus Soldiers red coloured heads Workers pale coloured heads Scientific classificationKingdom AnimaliaPhylum ArthropodaClass InsectaCohort PolyneopteraSuperorder DictyopteraOrder BlattodeaInfraorder Isoptera Brulle 1832Families Cratomastotermitidae 1 Mastotermitidae Termopsidae 2 Archotermopsidae Hodotermitidae Stolotermitidae Kalotermitidae Archeorhinotermitidae Stylotermitidae Rhinotermitidae Serritermitidae Termitidae Like ants and some bees and wasps from the separate order Hymenoptera termites divide as workers and soldiers that are usually sterile All colonies have fertile males called kings and one or more fertile females called queens Termites mostly feed on dead plant material and cellulose generally in the form of wood leaf litter soil or animal dung Termites are major detritivores particularly in the subtropical and tropical regions and their recycling of wood and plant matter is of considerable ecological importance Termites are among the most successful groups of insects on Earth colonising most landmasses except Antarctica Their colonies range in size from a few hundred individuals to enormous societies with several million individuals Termite queens have the longest known lifespan of any insect with some queens reportedly living up to 30 to 50 years Unlike ants which undergo a complete metamorphosis each individual termite goes through an incomplete metamorphosis that proceeds through egg nymph and adult stages Colonies are described as superorganisms because the termites form part of a self regulating entity the colony itself 5 Termites are a delicacy in the diet of some human cultures and are used in many traditional medicines Several hundred species are economically significant as pests that can cause serious damage to buildings crops or plantation forests Some species such as the West Indian drywood termite Cryptotermes brevis are regarded as invasive species Contents 1 Etymology 2 Taxonomy and evolution 2 1 Basal termite families 2 2 Neoisoptera 3 Distribution and diversity 4 Description 4 1 Caste system 5 Life cycle 5 1 Reproduction 6 Behaviour and ecology 6 1 Diet 6 2 Predators 6 3 Parasites pathogens and viruses 6 4 Locomotion and foraging 6 5 Competition 6 6 Communication 6 7 Defence 6 8 Relationship with other organisms 6 9 Relationship with the environment 7 Nests 7 1 Mounds 7 2 Shelter tubes 8 Relationship with humans 8 1 As pests 8 2 As food 8 3 In agriculture 8 4 In science and technology 8 5 In culture 9 See also 10 Notes 11 References 11 1 Cited literature 12 External linksEtymology EditThe infraorder name Isoptera is derived from the Greek words iso equal and ptera winged which refers to the nearly equal size of the fore and hind wings 6 Termite derives from the Latin and Late Latin word termes woodworm white ant altered by the influence of Latin terere to rub wear erode from the earlier word tarmes A termite nest is also known as a termitary or termitarium plural termitaria or termitariums 7 In earlier English termites were known as wood ants or white ants 8 The modern term was first used in 1781 9 Taxonomy and evolution Edit The external appearance of the giant northern termite Mastotermes darwiniensis is suggestive of the close relationship between termites and cockroaches Termites were formerly placed in the order Isoptera As early as 1934 suggestions were made that they were closely related to wood eating cockroaches genus Cryptocercus the woodroach based on the similarity of their symbiotic gut flagellates 10 In the 1960s additional evidence supporting that hypothesis emerged when F A McKittrick noted similar morphological characteristics between some termites and Cryptocercus nymphs 11 In 2008 DNA analysis from 16S rRNA sequences 12 supported the position of termites being nested within the evolutionary tree containing the order Blattodea which included the cockroaches 13 14 The cockroach genus Cryptocercus shares the strongest phylogenetical similarity with termites and is considered to be a sister group to termites 15 16 Termites and Cryptocercus share similar morphological and social features for example most cockroaches do not exhibit social characteristics but Cryptocercus takes care of its young and exhibits other social behaviour such as trophallaxis and allogrooming 17 Termites are thought to be the descendants of the genus Cryptocercus 13 18 Some researchers have suggested a more conservative measure of retaining the termites as the Termitoidae an epifamily within the cockroach order which preserves the classification of termites at family level and below 19 Termites have long been accepted to be closely related to cockroaches and mantids and they are classified in the same superorder Dictyoptera 20 21 The oldest unambiguous termite fossils date to the early Cretaceous but given the diversity of Cretaceous termites and early fossil records showing mutualism between microorganisms and these insects they possibly originated earlier in the Jurassic or Triassic 22 23 24 Possible evidence of a Jurassic origin is the assumption that the extinct Fruitafossor consumed termites judging from its morphological similarity to modern termite eating mammals 25 The oldest termite nest discovered is believed to be from the Upper Cretaceous in West Texas where the oldest known faecal pellets were also discovered 26 Claims that termites emerged earlier have faced controversy For example F M Weesner indicated that the Mastotermitidae termites may go back to the Late Permian 251 million years ago 27 and fossil wings that have a close resemblance to the wings of Mastotermes of the Mastotermitidae the most primitive living termite have been discovered in the Permian layers in Kansas 28 It is even possible that the first termites emerged during the Carboniferous 29 The folded wings of the fossil wood roach Pycnoblattina arranged in a convex pattern between segments 1a and 2a resemble those seen in Mastotermes the only living insect with the same pattern 28 Krishna et al though consider that all of the Paleozoic and Triassic insects tentatively classified as termites are in fact unrelated to termites and should be excluded from the Isoptera 30 Other studies suggest that the origin of termites is more recent having diverged from Cryptocercus sometime during the Early Cretaceous 3 Macro image of a worker The primitive giant northern termite Mastotermes darwiniensis exhibits numerous cockroach like characteristics that are not shared with other termites such as laying its eggs in rafts and having anal lobes on the wings 31 It has been proposed that the Isoptera and Cryptocercidae be grouped in the clade Xylophagodea 32 Termites are sometimes called white ants but the only resemblance to the ants is due to their sociality which is due to convergent evolution 33 34 with termites being the first social insects to evolve a caste system more than 100 million years ago 35 Termite genomes are generally relatively large compared to those of other insects the first fully sequenced termite genome of Zootermopsis nevadensis which was published in the journal Nature Communications consists of roughly 500Mb 36 while two subsequently published genomes Macrotermes natalensis and Cryptotermes secundus are considerably larger at around 1 3Gb 37 34 External phylogeny Dictyoptera Manipulatoridae extinct Alienopteridae extinct Mantodea Mantises Blattodea Blaberoidea Solumblattodea Corydiodea Blattoidea Blattoidae Kittrickea Lamproblattidae Xylophagodea Cryptocercidae brown hooded cockroaches Termitoidae Termites Internal phylogeny 30 Termitoidae Cratomastotermitidae Mastotermitidae Euisoptera Carinatermes Termopsidae Mariconitermes Hodotermitidae Cratokalotermes Archotermopsidae Stolotermitidae Tanytermes Baissatermes Dharmatermes Kalotermitidae Neoisoptera Archeorhinotermitidae Stylotermitidae Rhinotermitidae Serritermitidae Termitidae As of 2013 about 3 106 living and fossil termite species are recognised classified in 12 families reproductive and or soldier castes are usually required for identification The infraorder Isoptera is divided into the following clade and family groups showing the subfamilies in their respective classification 30 Basal termite families Edit Infraorder Isoptera Epifamily Termitoidae Family Cratomastotermitidae Family Mastotermitidae dd Parvorder EuisopteraFamily Arceotermitidae Family Archotermopsidae Family Hodotermitidae Family Kalotermitidae Family Krishnatermitidae Family Melqartitermitidae Family Mylacrotermitidae Family Stolotermitidae Family Tanytermitidae Family Termopsidae dd Neoisoptera Edit The Neoisoptera literally meaning newer termites in an evolutionary sense are a recently coined nanorder that include families commonly referred to as higher termites although some authorities only apply this term to the largest family Termitidae The latter characteristically do not have Pseudergate nymphs many lower termite worker nymphs have the capacity to develop into reproductive castes see below Cellulose digestion in higher termites has co evolved with eukaryotic gut microbiota 38 and many genera have symbiotic relationships with fungi such as Termitomyces in contrast lower termites typically have flagellates and prokaryotes in their hindguts Five families are now included here Family Archeorhinotermitidae Family RhinotermitidaeCoptotermitinae Heterotermitinae Prorhinoterminae Psammotermitinae Rhinotermitinae Termitogetoninae dd Family Serritermitidae Family Stylotermitidae dd Family TermitidaeApicotermitinae Cubitermitinae Foraminitermitinae Macrotermitinae Nasutitermitinae Sphaerotermitinae Syntermitinae Termitinae dd dd Distribution and diversity EditTermites are found on all continents except Antarctica The diversity of termite species is low in North America and Europe 10 species known in Europe and 50 in North America but is high in South America where over 400 species are known 39 Of the 3 000 termite species currently classified 1 000 are found in Africa where mounds are extremely abundant in certain regions Approximately 1 1 million active termite mounds can be found in the northern Kruger National Park alone 40 In Asia there are 435 species of termites which are mainly distributed in China Within China termite species are restricted to mild tropical and subtropical habitats south of the Yangtze River 39 In Australia all ecological groups of termites dampwood drywood subterranean are endemic to the country with over 360 classified species 39 Because termites are highly social and abundant they represent a disproportionate amount of the world s insect biomass Termites and ants comprise about 1 of insect species but represent more than 50 of insect biomass 41 Due to their soft cuticles termites do not inhabit cool or cold habitats 42 There are three ecological groups of termites dampwood drywood and subterranean Dampwood termites are found only in coniferous forests and drywood termites are found in hardwood forests subterranean termites live in widely diverse areas 39 One species in the drywood group is the West Indian drywood termite Cryptotermes brevis which is an invasive species in Australia 43 Diversity of Isoptera by continent Asia Africa North America South America Europe AustraliaEstimated number of species 435 1 000 50 400 10 360Description Edit Close up view of a worker s head Termites are usually small measuring between 4 to 15 millimetres 3 16 to 9 16 in in length 39 The largest of all extant termites are the queens of the species Macrotermes bellicosus measuring up to over 10 centimetres 4 in in length 44 Another giant termite the extinct Gyatermes styriensis flourished in Austria during the Miocene and had a wingspan of 76 millimetres 3 in and a body length of 25 millimetres 1 in 45 note 1 Most worker and soldier termites are completely blind as they do not have a pair of eyes However some species such as Hodotermes mossambicus have compound eyes which they use for orientation and to distinguish sunlight from moonlight 46 The alates winged males and females have eyes along with lateral ocelli Lateral ocelli however are not found in all termites absent in the families Hodotermitidae Termopsidae and Archotermopsidae 47 48 Like other insects termites have a small tongue shaped labrum and a clypeus the clypeus is divided into a postclypeus and anteclypeus Termite antennae have a number of functions such as the sensing of touch taste odours including pheromones heat and vibration The three basic segments of a termite antenna include a scape a pedicel typically shorter than the scape and the flagellum all segments beyond the scape and pedicel 48 The mouth parts contain a maxillae a labium and a set of mandibles The maxillae and labium have palps that help termites sense food and handling 48 Consistent with all insects the anatomy of the termite thorax consists of three segments the prothorax the mesothorax and the metathorax 48 Each segment contains a pair of legs On alates the wings are located at the mesothorax and metathorax which is consistent with all four winged insects The mesothorax and metathorax have well developed exoskeletal plates the prothorax has smaller plates 49 Diagram showing a wing along with the clypeus and leg Termites have a ten segmented abdomen with two plates the tergites and the sternites 50 The tenth abdominal segment has a pair of short cerci 51 There are ten tergites of which nine are wide and one is elongated 52 The reproductive organs are similar to those in cockroaches but are more simplified For example the intromittent organ is not present in male alates and the sperm is either immotile or aflagellate However Mastotermitidae termites have multiflagellate sperm with limited motility 53 The genitals in females are also simplified Unlike in other termites Mastotermitidae females have an ovipositor a feature strikingly similar to that in female cockroaches 54 The non reproductive castes of termites are wingless and rely exclusively on their six legs for locomotion The alates fly only for a brief amount of time so they also rely on their legs 50 The appearance of the legs is similar in each caste but the soldiers have larger and heavier legs The structure of the legs is consistent with other insects the parts of a leg include a coxa trochanter femur tibia and the tarsus 50 The number of tibial spurs on an individual s leg varies Some species of termite have an arolium located between the claws which is present in species that climb on smooth surfaces but is absent in most termites 55 Unlike in ants the hind wings and fore wings are of equal length 6 Most of the time the alates are poor flyers their technique is to launch themselves in the air and fly in a random direction 56 Studies show that in comparison to larger termites smaller termites cannot fly long distances When a termite is in flight its wings remain at a right angle and when the termite is at rest its wings remain parallel to the body 57 Caste system Edit Caste system of termites A King B Queen C Secondary queen D Tertiary queen E Soldiers F Worker Worker termites undertake the most labour within the colony being responsible for foraging food storage and brood and nest maintenance 58 59 Workers are tasked with the digestion of cellulose in food and are thus the most likely caste to be found in infested wood The process of worker termites feeding other nestmates is known as trophallaxis Trophallaxis is an effective nutritional tactic to convert and recycle nitrogenous components 60 It frees the parents from feeding all but the first generation of offspring allowing for the group to grow much larger and ensuring that the necessary gut symbionts are transferred from one generation to another Some termite species may rely on nymphs to perform work without differentiating as a separate caste 59 Workers may be male or female and are usually sterile especially in termites that have a nest site that is separate from their foraging site Sterile workers are sometimes termed as true workers while those that are fertile as in the wood nesting Archotermopsidae are termed as false workers 61 The soldier caste has anatomical and behavioural specialisations and their sole purpose is to defend the colony 62 Many soldiers have large heads with highly modified powerful jaws so enlarged they cannot feed themselves Instead like juveniles they are fed by workers 62 63 Fontanelles simple holes in the forehead that exude defensive secretions are a feature of the family Rhinotermitidae 64 Many species are readily identified using the characteristics of the soldiers larger and darker head and large mandibles 59 62 Among certain termites soldiers may use their globular phragmotic heads to block their narrow tunnels 65 Different sorts of soldiers include minor and major soldiers and nasutes which have a horn like nozzle frontal projection a nasus 59 These unique soldiers are able to spray noxious sticky secretions containing diterpenes at their enemies 66 Nitrogen fixation plays an important role in nasute nutrition 67 Soldiers are usually sterile but some species of Archotermopsidae are known to have neotenic forms with soldier like heads while also having sexual organs 68 The reproductive caste of a mature colony includes a fertile female and male known as the queen and king 69 The queen of the colony is responsible for egg production for the colony Unlike in ants the king mates with her for life 70 In some species the abdomen of the queen swells up dramatically to increase fecundity a characteristic known as physogastrism 58 69 Depending on the species the queen starts producing reproductive alates at a certain time of the year and huge swarms emerge from the colony when nuptial flight begins These swarms attract a wide variety of predators 69 Life cycle Edit A young termite nymph Nymphs first moult into workers but others may further moult to become soldiers or alates Termite and shed wings from other termites on an interior window sill Shedding of wings is associated with reproductive swarming 71 Termites are often compared with the social Hymenoptera ants and various species of bees and wasps but their differing evolutionary origins result in major differences in life cycle In the eusocial Hymenoptera the workers are exclusively female Males drones are haploid and develop from unfertilised eggs while females both workers and the queen are diploid and develop from fertilised eggs In contrast worker termites which constitute the majority in a colony are diploid individuals of both sexes and develop from fertilised eggs Depending on species male and female workers may have different roles in a termite colony 72 The life cycle of a termite begins with an egg but is different from that of a bee or ant in that it goes through a developmental process called incomplete metamorphosis with egg nymph and adult stages 73 Nymphs resemble small adults and go through a series of moults as they grow In some species eggs go through four moulting stages and nymphs go through three 74 Nymphs first moult into workers and then some workers go through further moulting and become soldiers or alates workers become alates only by moulting into alate nymphs 75 The development of nymphs into adults can take months the time period depends on food availability temperature and the general population of the colony Since nymphs are unable to feed themselves workers must feed them but workers also take part in the social life of the colony and have certain other tasks to accomplish such as foraging building or maintaining the nest or tending to the queen 59 76 Pheromones regulate the caste system in termite colonies preventing all but a very few of the termites from becoming fertile queens 77 Queens of the eusocial termite Reticulitermes speratus are capable of a long lifespan without sacrificing fecundity These long lived queens have a significantly lower level of oxidative damage including oxidative DNA damage than workers soldiers and nymphs 78 The lower levels of damage appear to be due to increased catalase an enzyme that protects against oxidative stress 78 Reproduction Edit Alates swarming during nuptial flight after rain Termite alates only leave the colony when a nuptial flight takes place Alate males and females pair up together and then land in search of a suitable place for a colony 79 A termite king and queen do not mate until they find such a spot When they do they excavate a chamber big enough for both close up the entrance and proceed to mate 79 After mating the pair never go outside and spend the rest of their lives in the nest Nuptial flight time varies in each species For example alates in certain species emerge during the day in summer while others emerge during the winter 80 The nuptial flight may also begin at dusk when the alates swarm around areas with many lights The time when nuptial flight begins depends on the environmental conditions the time of day moisture wind speed and precipitation 80 The number of termites in a colony also varies with the larger species typically having 100 1 000 individuals However some termite colonies including those with many individuals can number in the millions 45 The queen only lays 10 20 eggs in the very early stages of the colony but lays as many as 1 000 a day when the colony is several years old 59 At maturity a primary queen has a great capacity to lay eggs In some species the mature queen has a greatly distended abdomen and may produce 40 000 eggs a day 81 The two mature ovaries may have some 2 000 ovarioles each 82 The abdomen increases the queen s body length to several times more than before mating and reduces her ability to move freely attendant workers provide assistance source source source source source source Egg grooming behaviour of Reticulitermes speratus workers in a nursery cell The king grows only slightly larger after initial mating and continues to mate with the queen for life a termite queen can live between 30 to 50 years this is very different from ant colonies in which a queen mates once with the males and stores the gametes for life as the male ants die shortly after mating 70 76 If a queen is absent a termite king produces pheromones which encourage the development of replacement termite queens 83 As the queen and king are monogamous sperm competition does not occur 84 Termites going through incomplete metamorphosis on the path to becoming alates form a subcaste in certain species of termite functioning as potential supplementary reproductives These supplementary reproductives only mature into primary reproductives upon the death of a king or queen or when the primary reproductives are separated from the colony 75 85 Supplementaries have the ability to replace a dead primary reproductive and there may also be more than a single supplementary within a colony 59 Some queens have the ability to switch from sexual reproduction to asexual reproduction Studies show that while termite queens mate with the king to produce colony workers the queens reproduce their replacements neotenic queens parthenogenetically 86 87 The neotropical termite Embiratermes neotenicus and several other related species produce colonies that contain a primary king accompanied by a primary queen or by up to 200 neotenic queens that had originated through thelytokous parthenogenesis of a founding primary queen 88 The form of parthenogenesis likely employed maintains heterozygosity in the passage of the genome from mother to daughter thus avoiding inbreeding depression Behaviour and ecology EditDiet Edit Termite faecal pellets Termites are detritivores consuming dead plants at any level of decomposition They also play a vital role in the ecosystem by recycling waste material such as dead wood faeces and plants 89 90 91 Many species eat cellulose having a specialised midgut that breaks down the fibre 92 Termites are considered to be a major source 11 of atmospheric methane one of the prime greenhouse gases produced from the breakdown of cellulose 93 Termites rely primarily upon symbiotic protozoa metamonads and other microbes such as flagellate protists in their guts to digest the cellulose for them allowing them to absorb the end products for their own use 94 95 The microbial ecosystem present in the termite gut contains many species found nowhere else on Earth Termites hatch without these symbionts present in their guts and develop them after fed a culture from other termites 96 Gut protozoa such as Trichonympha in turn rely on symbiotic bacteria embedded on their surfaces to produce some of the necessary digestive enzymes Most higher termites especially in the family Termitidae can produce their own cellulase enzymes but they rely primarily upon the bacteria The flagellates have been lost in Termitidae 97 98 99 Researches have found species of spirochetes living in termite guts capable of fixing atmospheric nitrogen to a form usable by the insect 100 Scientists understanding of the relationship between the termite digestive tract and the microbial endosymbionts is still rudimentary what is true in all termite species however is that the workers feed the other members of the colony with substances derived from the digestion of plant material either from the mouth or anus 60 101 Judging from closely related bacterial species it is strongly presumed that the termites and cockroach s gut microbiota derives from their dictyopteran ancestors 102 Certain species such as Gnathamitermes tubiformans have seasonal food habits For example they may preferentially consume Red three awn Aristida longiseta during the summer Buffalograss Buchloe dactyloides from May to August and blue grama Bouteloua gracilis during spring summer and autumn Colonies of G tubiformans consume less food in spring than they do during autumn when their feeding activity is high 103 Various woods differ in their susceptibility to termite attack the differences are attributed to such factors as moisture content hardness and resin and lignin content In one study the drywood termite Cryptotermes brevis strongly preferred poplar and maple woods to other woods that were generally rejected by the termite colony These preferences may in part have represented conditioned or learned behaviour 104 Some species of termite practice fungiculture They maintain a garden of specialised fungi of genus Termitomyces which are nourished by the excrement of the insects When the fungi are eaten their spores pass undamaged through the intestines of the termites to complete the cycle by germinating in the fresh faecal pellets 105 106 Molecular evidence suggests that the family Macrotermitinae developed agriculture about 31 million years ago It is assumed that more than 90 percent of dry wood in the semiarid savannah ecosystems of Africa and Asia are reprocessed by these termites Originally living in the rainforest fungus farming allowed them to colonise the African savannah and other new environments eventually expanding into Asia 107 Depending on their feeding habits termites are placed into two groups the lower termites and higher termites The lower termites predominately feed on wood As wood is difficult to digest termites prefer to consume fungus infected wood because it is easier to digest and the fungi are high in protein Meanwhile the higher termites consume a wide variety of materials including faeces humus grass leaves and roots 108 The gut of the lower termites contains many species of bacteria along with protozoa and Holomastigotoides while the higher termites only have a few species of bacteria with no protozoa 109 Predators Edit Crab spider with a captured alate Termites are consumed by a wide variety of predators One termite species alone Hodotermes mossambicus was found in the stomach contents of 65 birds and 19 mammals 110 Arthropods such as ants 111 112 centipedes cockroaches crickets dragonflies scorpions and spiders 113 reptiles such as lizards 114 and amphibians such as frogs 115 and toads consume termites with two spiders in the family Ammoxenidae being specialist termite predators 116 117 118 Other predators include aardvarks aardwolves anteaters bats bears bilbies many birds echidnas foxes galagos numbats mice and pangolins 116 119 120 121 The aardwolf is an insectivorous mammal that primarily feeds on termites it locates its food by sound and also by detecting the scent secreted by the soldiers a single aardwolf is capable of consuming thousands of termites in a single night by using its long sticky tongue 122 123 Sloth bears break open mounds to consume the nestmates while chimpanzees have developed tools to fish termites from their nest Wear pattern analysis of bone tools used by the early hominin Paranthropus robustus suggests that they used these tools to dig into termite mounds 124 A Matabele ant Megaponera analis kills a Macrotermes bellicosus termite soldier during a raid Among all predators ants are the greatest enemy to termites 111 112 Some ant genera are specialist predators of termites For example Megaponera is a strictly termite eating termitophagous genus that perform raiding activities some lasting several hours 125 126 Paltothyreus tarsatus is another termite raiding species with each individual stacking as many termites as possible in its mandibles before returning home all the while recruiting additional nestmates to the raiding site through chemical trails 111 The Malaysian basicerotine ants Eurhopalothrix heliscata uses a different strategy of termite hunting by pressing themselves into tight spaces as they hunt through rotting wood housing termite colonies Once inside the ants seize their prey by using their short but sharp mandibles 111 Tetramorium uelense is a specialised predator species that feeds on small termites A scout recruits 10 30 workers to an area where termites are present killing them by immobilising them with their stinger 127 Centromyrmex and Iridomyrmex colonies sometimes nest in termite mounds and so the termites are preyed on by these ants No evidence for any kind of relationship other than a predatory one is known 128 129 Other ants including Acanthostichus Camponotus Crematogaster Cylindromyrmex Leptogenys Odontomachus Ophthalmopone Pachycondyla Rhytidoponera Solenopsis and Wasmannia also prey on termites 119 111 130 In contrast to all these ant species and despite their enormous diversity of prey Dorylus ants rarely consume termites 131 Ants are not the only invertebrates that perform raids Many sphecoid wasps and several species including Polybia and Angiopolybia are known to raid termite mounds during the termites nuptial flight 132 Parasites pathogens and viruses Edit Termites are less likely to be attacked by parasites than bees wasps and ants as they are usually well protected in their mounds 133 134 Nevertheless termites are infected by a variety of parasites Some of these include dipteran flies 135 Pyemotes mites and a large number of nematode parasites Most nematode parasites are in the order Rhabditida 136 others are in the genus Mermis Diplogaster aerivora and Harteria gallinarum 137 Under imminent threat of an attack by parasites a colony may migrate to a new location 138 Certain fungal pathogens such as Aspergillus nomius and Metarhizium anisopliae are however major threats to a termite colony as they are not host specific and may infect large portions of the colony 139 140 transmission usually occurs via direct physical contact 141 M anisopliae is known to weaken the termite immune system Infection with A nomius only occurs when a colony is under great stress Over 34 fungal species are known to live as parasites on the exoskeleton of termites with many being host specific and only causing indirect harm to their host 142 Termites are infected by viruses including Entomopoxvirinae and the Nuclear Polyhedrosis Virus 143 144 Locomotion and foraging Edit Because the worker and soldier castes lack wings and thus never fly and the reproductives use their wings for just a brief amount of time termites predominantly rely upon their legs to move about 50 Foraging behaviour depends on the type of termite For example certain species feed on the wood structures they inhabit and others harvest food that is near the nest 145 Most workers are rarely found out in the open and do not forage unprotected they rely on sheeting and runways to protect them from predators 58 Subterranean termites construct tunnels and galleries to look for food and workers who manage to find food sources recruit additional nestmates by depositing a phagostimulant pheromone that attracts workers 146 Foraging workers use semiochemicals to communicate with each other 147 and workers who begin to forage outside of their nest release trail pheromones from their sternal glands 148 In one species Nasutitermes costalis there are three phases in a foraging expedition first soldiers scout an area When they find a food source they communicate to other soldiers and a small force of workers starts to emerge In the second phase workers appear in large numbers at the site The third phase is marked by a decrease in the number of soldiers present and an increase in the number of workers 149 Isolated termite workers may engage in Levy flight behaviour as an optimised strategy for finding their nestmates or foraging for food 150 Competition Edit Competition between two colonies always results in agonistic behaviour towards each other resulting in fights These fights can cause mortality on both sides and in some cases the gain or loss of territory 151 152 Cemetery pits may be present where the bodies of dead termites are buried 153 Studies show that when termites encounter each other in foraging areas some of the termites deliberately block passages to prevent other termites from entering 147 154 Dead termites from other colonies found in exploratory tunnels leads to the isolation of the area and thus the need to construct new tunnels 155 Conflict between two competitors does not always occur For example though they might block each other s passages colonies of Macrotermes bellicosus and Macrotermes subhyalinus are not always aggressive towards each other 156 Suicide cramming is known in Coptotermes formosanus Since C formosanus colonies may get into physical conflict some termites squeeze tightly into foraging tunnels and die successfully blocking the tunnel and ending all agonistic activities 157 Among the reproductive caste neotenic queens may compete with each other to become the dominant queen when there are no primary reproductives This struggle among the queens leads to the elimination of all but a single queen which with the king takes over the colony 158 Ants and termites may compete with each other for nesting space In particular ants that prey on termites usually have a negative impact on arboreal nesting species 159 Communication Edit Hordes of Nasutitermes on a march for food following and leaving trail pheromones Most termites are blind so communication primarily occurs through chemical mechanical and pheromonal cues 47 147 These methods of communication are used in a variety of activities including foraging locating reproductives construction of nests recognition of nestmates nuptial flight locating and fighting enemies and defending the nests 47 147 The most common way of communicating is through antennation 147 A number of pheromones are known including contact pheromones which are transmitted when workers are engaged in trophallaxis or grooming and alarm trail and sex pheromones The alarm pheromone and other defensive chemicals are secreted from the frontal gland Trail pheromones are secreted from the sternal gland and sex pheromones derive from two glandular sources the sternal and tergal glands 47 When termites go out to look for food they forage in columns along the ground through vegetation A trail can be identified by the faecal deposits or runways that are covered by objects Workers leave pheromones on these trails which are detected by other nestmates through olfactory receptors 63 Termites can also communicate through mechanical cues vibrations and physical contact 63 147 These signals are frequently used for alarm communication or for evaluating a food source 147 160 When termites construct their nests they use predominantly indirect communication No single termite would be in charge of any particular construction project Individual termites react rather than think but at a group level they exhibit a sort of collective cognition Specific structures or other objects such as pellets of soil or pillars cause termites to start building The termite adds these objects onto existing structures and such behaviour encourages building behaviour in other workers The result is a self organised process whereby the information that directs termite activity results from changes in the environment rather than from direct contact among individuals 147 Termites can distinguish nestmates and non nestmates through chemical communication and gut symbionts chemicals consisting of hydrocarbons released from the cuticle allow the recognition of alien termite species 161 162 Each colony has its own distinct odour This odour is a result of genetic and environmental factors such as the termites diet and the composition of the bacteria within the termites intestines 163 Defence Edit See also Insect defences Termites rush to a damaged area of the nest Termites rely on alarm communication to defend a colony 147 Alarm pheromones can be released when the nest has been breached or is being attacked by enemies or potential pathogens Termites always avoid nestmates infected with Metarhizium anisopliae spores through vibrational signals released by infected nestmates 164 Other methods of defence include intense jerking and secretion of fluids from the frontal gland and defecating faeces containing alarm pheromones 147 165 In some species some soldiers block tunnels to prevent their enemies from entering the nest and they may deliberately rupture themselves as an act of defence 166 In cases where the intrusion is coming from a breach that is larger than the soldier s head soldiers form a phalanx like formation around the breach and bite at intruders 167 If an invasion carried out by Megaponera analis is successful an entire colony may be destroyed although this scenario is rare 167 To termites any breach of their tunnels or nests is a cause for alarm When termites detect a potential breach the soldiers usually bang their heads apparently to attract other soldiers for defence and to recruit additional workers to repair any breach 63 Additionally an alarmed termite bumps into other termites which causes them to be alarmed and to leave pheromone trails to the disturbed area which is also a way to recruit extra workers 63 Nasute termite soldiers on rotten wood The pantropical subfamily Nasutitermitinae has a specialised caste of soldiers known as nasutes that have the ability to exude noxious liquids through a horn like frontal projection that they use for defence 168 Nasutes have lost their mandibles through the course of evolution and must be fed by workers 66 A wide variety of monoterpene hydrocarbon solvents have been identified in the liquids that nasutes secrete 169 Similarly Formosan subterranean termites have been known to secrete naphthalene to protect their nests 170 Soldiers of the species Globitermes sulphureus commit suicide by autothysis rupturing a large gland just beneath the surface of their cuticles The thick yellow fluid in the gland becomes very sticky on contact with the air entangling ants or other insects that are trying to invade the nest 171 172 Another termite Neocapriterme taracua also engages in suicidal defence Workers physically unable to use their mandibles while in a fight form a pouch full of chemicals then deliberately rupture themselves releasing toxic chemicals that paralyse and kill their enemies 173 The soldiers of the neotropical termite family Serritermitidae have a defence strategy which involves front gland autothysis with the body rupturing between the head and abdomen When soldiers guarding nest entrances are attacked by intruders they engage in autothysis creating a block that denies entry to any attacker 174 Workers use several different strategies to deal with their dead including burying cannibalism and avoiding a corpse altogether 175 176 177 To avoid pathogens termites occasionally engage in necrophoresis in which a nestmate carries away a corpse from the colony to dispose of it elsewhere 178 Which strategy is used depends on the nature of the corpse a worker is dealing with i e the age of the carcass 178 Relationship with other organisms Edit Rhizanthella gardneri is the only orchid known to be pollinated by termites A species of fungus is known to mimic termite eggs successfully avoiding its natural predators These small brown balls known as termite balls rarely kill the eggs and in some cases the workers tend to them 179 This fungus mimics these eggs by producing a cellulose digesting enzyme known as glucosidases 180 A unique mimicking behaviour exists between various species of Trichopsenius beetles and certain termite species within Reticulitermes The beetles share the same cuticle hydrocarbons as the termites and even biosynthesize them This chemical mimicry allows the beetles to integrate themselves within the termite colonies 181 The developed appendages on the physogastric abdomen of Austrospirachtha mimetes allows the beetle to mimic a termite worker 182 Some species of ant are known to capture termites to use as a fresh food source later on rather than killing them For example Formica nigra captures termites and those who try to escape are immediately seized and driven underground 183 Certain species of ants in the subfamily Ponerinae conduct these raids although other ant species go in alone to steal the eggs or nymphs 159 Ants such as Megaponera analis attack the outside of mounds and Dorylinae ants attack underground 159 184 Despite this some termites and ants can coexist peacefully Some species of termite including Nasutitermes corniger form associations with certain ant species to keep away predatory ant species 185 The earliest known association between Azteca ants and Nasutitermes termites date back to the Oligocene to Miocene period 186 An ant raiding party collecting Pseudocanthotermes militaris termites after a successful raid 54 species of ants are known to inhabit Nasutitermes mounds both occupied and abandoned ones 187 One reason many ants live in Nasutitermes mounds is due to the termites frequent occurrence in their geographical range another is to protect themselves from floods 187 188 Iridomyrmex also inhabits termite mounds although no evidence for any kind of relationship other than a predatory one is known 128 In rare cases certain species of termites live inside active ant colonies 189 Some invertebrate organisms such as beetles caterpillars flies and millipedes are termitophiles and dwell inside termite colonies they are unable to survive independently 63 As a result certain beetles and flies have evolved with their hosts They have developed a gland that secrete a substance that attracts the workers by licking them Mounds may also provide shelter and warmth to birds lizards snakes and scorpions 63 Termites are known to carry pollen and regularly visit flowers 190 so are regarded as potential pollinators for a number of flowering plants 191 One flower in particular Rhizanthella gardneri is regularly pollinated by foraging workers and it is perhaps the only Orchidaceae flower in the world to be pollinated by termites 190 Many plants have developed effective defences against termites However seedlings are vulnerable to termite attacks and need additional protection as their defence mechanisms only develop when they have passed the seedling stage 192 Defence is typically achieved by secreting antifeedant chemicals into the woody cell walls 193 This reduces the ability of termites to efficiently digest the cellulose A commercial product Blockaid has been developed in Australia that uses a range of plant extracts to create a paint on nontoxic termite barrier for buildings 193 An extract of a species of Australian figwort Eremophila has been shown to repel termites 194 tests have shown that termites are strongly repelled by the toxic material to the extent that they will starve rather than consume the food When kept close to the extract they become disoriented and eventually die 194 Relationship with the environment Edit Termite populations can be substantially impacted by environmental changes including those caused by human intervention A Brazilian study investigated the termite assemblages of three sites of Caatinga under different levels of anthropogenic disturbance in the semi arid region of northeastern Brazil were sampled using 65 x 2 m transects 195 A total of 26 species of termites were present in the three sites and 196 encounters were recorded in the transects The termite assemblages were considerably different among sites with a conspicuous reduction in both diversity and abundance with increased disturbance related to the reduction of tree density and soil cover and with the intensity of trampling by cattle and goats The wood feeders were the most severely affected feeding group Nests Edit source source source source source source source source source source source source Termite workers at work An arboreal termite nest in Mexico Termite nest in a Banksia Palm Beach Sydney A termite nest can be considered as being composed of two parts the inanimate and the animate The animate is all of the termites living inside the colony and the inanimate part is the structure itself which is constructed by the termites 196 Nests can be broadly separated into three main categories subterranean completely below ground epigeal protruding above the soil surface and arboreal built above ground but always connected to the ground via shelter tubes 197 Epigeal nests mounds protrude from the earth with ground contact and are made out of earth and mud 198 A nest has many functions such as providing a protected living space and providing shelter against predators Most termites construct underground colonies rather than multifunctional nests and mounds 199 Primitive termites of today nest in wooden structures such as logs stumps and the dead parts of trees as did termites millions of years ago 197 To build their nests termites primarily use faeces which have many desirable properties as a construction material 200 Other building materials include partly digested plant material used in carton nests arboreal nests built from faecal elements and wood and soil used in subterranean nest and mound construction Not all nests are visible as many nests in tropical forests are located underground 199 Species in the subfamily Apicotermitinae are good examples of subterranean nest builders as they only dwell inside tunnels 200 Other termites live in wood and tunnels are constructed as they feed on the wood Nests and mounds protect the termites soft bodies against desiccation light pathogens and parasites as well as providing a fortification against predators 201 Nests made out of carton are particularly weak and so the inhabitants use counter attack strategies against invading predators 202 Arboreal carton nests of mangrove swamp dwelling Nasutitermes are enriched in lignin and depleted in cellulose and xylans This change is caused by bacterial decay in the gut of the termites they use their faeces as a carton building material Arboreal termites nests can account for as much as 2 of above ground carbon storage in Puerto Rican mangrove swamps These Nasutitermes nests are mainly composed of partially biodegraded wood material from the stems and branches of mangrove trees namely Rhizophora mangle red mangrove Avicennia germinans black mangrove and Laguncularia racemosa white mangrove 203 Some species build complex nests called polycalic nests this habitat is called polycalism Polycalic species of termites form multiple nests or calies connected by subterranean chambers 119 The termite genera Apicotermes and Trinervitermes are known to have polycalic species 204 Polycalic nests appear to be less frequent in mound building species although polycalic arboreal nests have been observed in a few species of Nasutitermes 204 Mounds Edit Main article Mound building termites Wikimedia Commons has media related to Termite mounds Nests are considered mounds if they protrude from the earth s surface 200 A mound provides termites the same protection as a nest but is stronger 202 Mounds located in areas with torrential and continuous rainfall are at risk of mound erosion due to their clay rich construction Those made from carton can provide protection from the rain and in fact can withstand high precipitation 200 Certain areas in mounds are used as strong points in case of a breach For example Cubitermes colonies build narrow tunnels used as strong points as the diameter of the tunnels is small enough for soldiers to block 205 A highly protected chamber known as the queen s cell houses the queen and king and is used as a last line of defence 202 Species in the genus Macrotermes arguably build the most complex structures in the insect world constructing enormous mounds 200 These mounds are among the largest in the world reaching a height of 8 to 9 metres 26 to 29 feet and consist of chimneys pinnacles and ridges 63 Another termite species Amitermes meridionalis can build nests 3 to 4 metres 9 to 13 feet high and 2 5 metres 8 feet wide The tallest mound ever recorded was 12 8 metres 42 ft long found in the Democratic Republic of the Congo 206 The sculptured mounds sometimes have elaborate and distinctive forms such as those of the compass termite Amitermes meridionalis and A laurensis which builds tall wedge shaped mounds with the long axis oriented approximately north south which gives them their common name 207 208 This orientation has been experimentally shown to assist thermoregulation The north south orientation causes the internal temperature of a mound to increase rapidly during the morning while avoiding overheating from the midday sun The temperature then remains at a plateau for the rest of the day until the evening 209 Cathedral mounds in the Northern Territory Australia Mounds of compass or magnetic termites Amitermes oriented north south thereby avoiding mid day heat Termite mound in Queensland Australia Termites in a mound Analamazoatra Reserve Madagascar Termite mound in NamibiaShelter tubes Edit Nasutiterminae shelter tubes on a tree trunk provide cover for the trail from nest to forest floor Termites construct shelter tubes also known as earthen tubes or mud tubes that start from the ground These shelter tubes can be found on walls and other structures 210 Constructed by termites during the night a time of higher humidity these tubes provide protection to termites from potential predators especially ants 211 Shelter tubes also provide high humidity and darkness and allow workers to collect food sources that cannot be accessed in any other way 210 These passageways are made from soil and faeces and are normally brown in colour The size of these shelter tubes depends on the number of food sources that are available They range from less than 1 cm to several cm in width but may be dozens of metres in length 211 Relationship with humans EditAs pests Edit Termite mound as an obstacle on a runway at Khorixas Namibia Termite damage on external structure Owing to their wood eating habits many termite species can do significant damage to unprotected buildings and other wooden structures 212 Termites play an important role as decomposers of wood and vegetative material and the conflict with humans occurs where structures and landscapes containing structural wood components cellulose derived structural materials and ornamental vegetation provide termites with a reliable source of food and moisture 213 Their habit of remaining concealed often results in their presence being undetected until the timbers are severely damaged with only a thin exterior layer of wood remaining which protects them from the environment 214 Of the 3 106 species known only 183 species cause damage 83 species cause significant damage to wooden structures 212 In North America 18 subterranean species are pests 215 in Australia 16 species have an economic impact in the Indian subcontinent 26 species are considered pests and in tropical Africa 24 In Central America and the West Indies there are 17 pest species 212 Among the termite genera Coptotermes has the highest number of pest species of any genus with 28 species known to cause damage 212 Less than 10 of drywood termites are pests but they infect wooden structures and furniture in tropical subtropical and other regions Dampwood termites only attack lumber material exposed to rainfall or soil 212 Drywood termites thrive in warm climates and human activities can enable them to invade homes since they can be transported through contaminated goods containers and ships 212 Colonies of termites have been seen thriving in warm buildings located in cold regions 216 Some termites are considered invasive species Cryptotermes brevis the most widely introduced invasive termite species in the world has been introduced to all the islands in the West Indies and to Australia 43 212 Termite damage in wooden house stumps In addition to causing damage to buildings termites can also damage food crops 217 Termites may attack trees whose resistance to damage is low but generally ignore fast growing plants Most attacks occur at harvest time crops and trees are attacked during the dry season 217 The damage caused by termites costs the southwestern United States approximately 1 5 billion each year in wood structure damage but the true cost of damage worldwide cannot be determined 212 218 Drywood termites are responsible for a large proportion of the damage caused by termites 219 The goal of termite control is to keep structures and susceptible ornamental plants free from termites 220 Structures may be homes or business or elements such as wooden fence posts and telephone poles Regular and thorough inspections by a trained professional may be necessary to detect termite activity in the absence of more obvious signs like termite swarmers or alates inside or adjacent to a structure Termite monitors made of wood or cellulose adjacent to a structure may also provide indication of termite foraging activity where it will be in conflict with humans Termites can be controlled by application of Bordeaux mixture or other substances that contain copper such as chromated copper arsenate 221 In the United states application of a soil termiticide with the active ingredient Fipronil such as Termidor SC or Taurus SC by a licensed professional 222 is a common remedy approved by the Environmental Protection Agency for economically significant subterranean termites 223 224 A growing demand for alternative green and more natural extermination methods has increased demand for mechanical and biological control methods such as Orange Oil To better control the population of termites various methods have been developed to track termite movements 218 One early method involved distributing termite bait laced with immunoglobulin G IgG marker proteins from rabbits or chickens Termites collected from the field could be tested for the rabbit IgG markers using a rabbit IgG specific assay More recently developed less expensive alternatives include tracking the termites using egg white cow milk or soy milk proteins which can be sprayed on termites in the field Termites bearing these proteins can be traced using a protein specific ELISA test 218 In 1994 termites of the species Reticulitermes grassei were identified in two bungalows in Saunton Devon Anecdotal evidence suggests the infestation could date back 70 years before the official identification There are reports that gardeners had seen white ants and that a greenhouse had had to be replaced in the past The Saunton infestation was the first and only colony ever recorded in the UK In 1998 Termite Eradication Programme was set up with the intention of containing and eradicating the colony The TEP was managed by the Ministry of Housing Communities amp Local Government now the Department for Levelling Up Housing and Communities The TEP used insect growth regulators to prevent the termites from reaching maturity and reproducing In 2021 the UK s Termite Eradication Programme announced the eradication of the colony the first time a country has eradicated termites 225 As food Edit See also Entomophagy Mozambican boys from the Yawo tribe collecting flying termites These flying alates were collected as they came out of their nests in the ground during the early days of the rainy season 43 termite species are used as food by humans or are fed to livestock 226 These insects are particularly important in impoverished countries where malnutrition is common as the protein from termites can help improve the human diet Termites are consumed in many regions globally but this practice has only become popular in developed nations in recent years 226 Termites are consumed by people in many different cultures around the world In many parts of Africa the alates are an important factor in the diets of native populations 227 Groups have different ways of collecting or cultivating insects sometimes collecting soldiers from several species Though harder to acquire queens are regarded as a delicacy 228 Termite alates are high in nutrition with adequate levels of fat and protein They are regarded as pleasant in taste having a nut like flavour after they are cooked 227 Alates are collected when the rainy season begins During a nuptial flight they are typically seen around lights to which they are attracted and so nets are set up on lamps and captured alates are later collected The wings are removed through a technique that is similar to winnowing The best result comes when they are lightly roasted on a hot plate or fried until crisp Oil is not required as their bodies usually contain sufficient amounts of oil Termites are typically eaten when livestock is lean and tribal crops have not yet developed or produced any food or if food stocks from a previous growing season are limited 227 In addition to Africa termites are consumed in local or tribal areas in Asia and North and South America In Australia Indigenous Australians are aware that termites are edible but do not consume them even in times of scarcity there are few explanations as to why 227 228 Termite mounds are the main sources of soil consumption geophagy in many countries including Kenya Tanzania Zambia Zimbabwe and South Africa 229 230 231 232 Researchers have suggested that termites are suitable candidates for human consumption and space agriculture as they are high in protein and can be used to convert inedible waste to consumable products for humans 233 In agriculture Edit Scientists have developed a more affordable method of tracing the movement of termites using traceable proteins 218 Termites can be major agricultural pests particularly in East Africa and North Asia where crop losses can be severe 3 100 in crop loss in Africa 234 Counterbalancing this is the greatly improved water infiltration where termite tunnels in the soil allow rainwater to soak in deeply which helps reduce runoff and consequent soil erosion through bioturbation 235 In South America cultivated plants such as eucalyptus upland rice and sugarcane can be severely damaged by termite infestations with attacks on leaves roots and woody tissue Termites can also attack other plants including cassava coffee cotton fruit trees maize peanuts soybeans and vegetables 21 Mounds can disrupt farming activities making it difficult for farmers to operate farming machinery however despite farmers dislike of the mounds it is often the case that no net loss of production occurs 21 Termites can be beneficial to agriculture such as by boosting crop yields and enriching the soil Termites and ants can re colonise untilled land that contains crop stubble which colonies use for nourishment when they establish their nests The presence of nests in fields enables larger amounts of rainwater to soak into the ground and increases the amount of nitrogen in the soil both essential for the growth of crops 236 In science and technology Edit See also Renewable energy Termite inspired robots and Sustainable architecture The termite gut has inspired various research efforts aimed at replacing fossil fuels with cleaner renewable energy sources 237 Termites are efficient bioreactors capable of producing two litres of hydrogen from a single sheet of paper 238 Approximately 200 species of microbes live inside the termite hindgut releasing the hydrogen that was trapped inside wood and plants that they digest 237 239 Through the action of unidentified enzymes in the termite gut lignocellulose polymers are broken down into sugars and are transformed into hydrogen The bacteria within the gut turns the sugar and hydrogen into cellulose acetate an acetate ester of cellulose on which termites rely for energy 237 Community DNA sequencing of the microbes in the termite hindgut has been employed to provide a better understanding of the metabolic pathway 237 Genetic engineering may enable hydrogen to be generated in bioreactors from woody biomass 237 The development of autonomous robots capable of constructing intricate structures without human assistance has been inspired by the complex mounds that termites build 240 These robots work independently and can move by themselves on a tracked grid capable of climbing and lifting up bricks Such robots may be useful for future projects on Mars or for building levees to prevent flooding 241 Termites use sophisticated means to control the temperatures of their mounds As discussed above the shape and orientation of the mounds of the Australian compass termite stabilises their internal temperatures during the day As the towers heat up the solar chimney effect stack effect creates an updraft of air within the mound 242 Wind blowing across the tops of the towers enhances the circulation of air through the mounds which also include side vents in their construction The solar chimney effect has been in use for centuries in the Middle East and Near East for passive cooling as well as in Europe by the Romans 243 It is only relatively recently however that climate responsive construction techniques have become incorporated into modern architecture Especially in Africa the stack effect has become a popular means to achieve natural ventilation and passive cooling in modern buildings 242 In culture Edit The pink hued Eastgate Centre The Eastgate Centre is a shopping centre and office block in central Harare Zimbabwe whose architect Mick Pearce used passive cooling inspired by that used by the local termites 244 It was the first major building exploiting termite inspired cooling techniques to attract international attention Other such buildings include the Learning Resource Center at the Catholic University of Eastern Africa and the Council House 2 building in Melbourne Australia 242 Few zoos hold termites due to the difficulty in keeping them captive and to the reluctance of authorities to permit potential pests One of the few that do the Zoo Basel in Switzerland has two thriving Macrotermes bellicosus populations resulting in an event very rare in captivity the mass migrations of young flying termites This happened in September 2008 when thousands of male termites left their mound each night died and covered the floors and water pits of the house holding their exhibit 245 African tribes in several countries have termites as totems and for this reason tribe members are forbidden to eat the reproductive alates 246 Termites are widely used in traditional popular medicine they are used as treatments for diseases and other conditions such as asthma bronchitis hoarseness influenza sinusitis tonsillitis and whooping cough 226 In Nigeria Macrotermes nigeriensis is used for spiritual protection and to treat wounds and sick pregnant women In Southeast Asia termites are used in ritual practices In Malaysia Singapore and Thailand termite mounds are commonly worshiped among the populace 247 Abandoned mounds are viewed as structures created by spirits believing a local guardian dwells within the mound this is known as Keramat and Datok Kong In urban areas local residents construct red painted shrines over mounds that have been abandoned where they pray for good health protection and luck 247 See also EditStigmergy Termite shield XylophagyNotes Edit It is unknown whether the termite was female or male If it was a female the body length would be far greater than 25 millimetres when mature References Edit Behrensmeyer A K Turner A Fossilworks Gateway to the Paleobiology Database Engel M S Grimaldi D A Krishna K 2009 Termites Isoptera their phylogeny classification and rise to ecological dominance American Museum Novitates 3650 1 27 doi 10 1206 651 1 hdl 2246 5969 ISSN 0003 0082 S2CID 56166416 a b Evangelista Dominic A Wipfler Benjamin Bethoux Olivier Donath Alexander Fujita Mari Kohli Manpreet K Legendre Frederic Liu Shanlin Machida Ryuichiro Misof Bernhard Peters Ralph S 2019 01 30 An integrative phylogenomic approach illuminates the evolutionary history of cockroaches and termites Blattodea Proceedings of the Royal Society B Biological Sciences 286 1895 20182076 doi 10 1098 rspb 2018 2076 ISSN 0962 8452 PMC 6364590 PMID 30963947 Termite Merriam Webster com Bignell Roisin amp Lo 2010 p 2 a b Cranshaw W 2013 11 Bugs Rule An Introduction to the World of Insects Princeton New Jersey Princeton University Press p 188 ISBN 978 0 691 12495 7 Lobeck A Kohl 1939 Geomorphology an Introduction to the Study of Landscapes 1st ed University of California McGraw Hill Book Company Incorporated pp 431 432 ASIN B002P5O9SC Harper Douglas Termite Online Etymology Dictionary Termite Merriam Webster Online Dictionary Retrieved 5 January 2015 Cleveland L R Hall S K Sanders E P Collier J 1934 The Wood Feeding Roach Cryptocercus its protozoa and the symbiosis between protozoa and roach Memoirs of the American Academy of Arts and Sciences 17 2 185 382 doi 10 1093 aesa 28 2 216 McKittrick F A 1965 A contribution to the understanding of cockroach termite affinities Annals of the Entomological Society of America 58 1 18 22 doi 10 1093 aesa 58 1 18 PMID 5834489 Ware J L Litman J Klass K D Spearman L A 2008 Relationships among the major lineages of Dictyoptera the effect of outgroup selection on dictyopteran tree topology Systematic Entomology 33 3 429 450 doi 10 1111 j 1365 3113 2008 00424 x S2CID 86777253 a b Inward D Beccaloni G Eggleton P 2007 Death of an order a comprehensive molecular phylogenetic study confirms that termites are eusocial cockroaches Biology Letters 3 3 331 5 doi 10 1098 rsbl 2007 0102 PMC 2464702 PMID 17412673 Eggleton P Beccaloni G Inward D 2007 Response to Lo et al Biology Letters 3 5 564 565 doi 10 1098 rsbl 2007 0367 PMC 2391203 Ohkuma M Noda S Hongoh Y Nalepa C A Inoue T 2009 Inheritance and diversification of symbiotic trichonymphid flagellates from a common ancestor of termites and the cockroach Cryptocercus Proceedings of the Royal Society B Biological Sciences 276 1655 239 245 doi 10 1098 rspb 2008 1094 PMC 2674353 PMID 18812290 Lo N Tokuda G Watanabe H Rose H Slaytor M Maekawa K Bandi C Noda H June 2000 Evidence from multiple gene sequences indicates that termites evolved from wood feeding cockroaches Current Biology 10 13 801 814 doi 10 1016 S0960 9822 00 00561 3 PMID 10898984 S2CID 14059547 Grimaldi D Engel M S 2005 Evolution of the insects 1st ed Cambridge Cambridge University Press p 237 ISBN 978 0 521 82149 0 Klass K D Nalepa C Lo N 2008 Wood feeding cockroaches as models for termite evolution Insecta Dictyoptera Cryptocercus vs Parasphaeria boleiriana Molecular Phylogenetics amp Evolution 46 3 809 817 doi 10 1016 j ympev 2007 11 028 PMID 18226554 Lo N Engel M S Cameron S Nalepa C A Tokuda G Grimaldi D Kitade O Krishna K Klass K D Maekawa K Miura T Thompson G J 2007 Comment Save Isoptera a comment on Inward et al Biology Letters 3 5 562 563 doi 10 1098 rsbl 2007 0264 PMC 2391185 PMID 17698448 Costa James 2006 The other insect societies Harvard University Press pp 135 136 ISBN 978 0 674 02163 1 a b c Capinera J L 2008 Encyclopedia of Entomology 2nd ed Dordrecht Springer pp 3033 3037 3754 ISBN 978 1 4020 6242 1 Vrsanky P Aristov D 2014 Termites Isoptera from the Jurassic Cretaceous boundary Evidence for the longevity of their earliest genera European Journal of Entomology 111 1 137 141 doi 10 14411 eje 2014 014 Poinar G O 2009 Description of an early Cretaceous termite Isoptera Kalotermitidae and its associated intestinal protozoa with comments on their co evolution Parasites amp Vectors 2 1 17 12 doi 10 1186 1756 3305 2 12 PMC 2669471 PMID 19226475 Legendre F Nel A Svenson G J Robillard T Pellens R Grandcolas P Escriva H 2015 Phylogeny of Dictyoptera Dating the Origin of Cockroaches Praying Mantises and Termites with Molecular Data and Controlled Fossil Evidence PLOS ONE 10 7 e0130127 Bibcode 2015PLoSO 1030127L doi 10 1371 journal pone 0130127 PMC 4511787 PMID 26200914 Luo Z X Wible J R 2005 A Late Jurassic digging mammal and early mammalian diversification Science 308 5718 103 107 Bibcode 2005Sci 308 103L doi 10 1126 science 1108875 PMID 15802602 S2CID 7031381 Rohr D M Boucot A J Miller J Abbott M 1986 Oldest termite nest from the Upper Cretaceous of west Texas Geology 14 1 87 Bibcode 1986Geo 14 87R doi 10 1130 0091 7613 1986 14 lt 87 OTNFTU gt 2 0 CO 2 Weesner F M 1960 Evolution and Biology of the Termites Annual Review of Entomology 5 1 153 170 doi 10 1146 annurev en 05 010160 001101 a b Tilyard R J 1937 Kansas Permian insects Part XX the cockroaches or order Blattaria American Journal of Science 34 201 169 202 249 276 Bibcode 1937AmJS 34 169T doi 10 2475 ajs s5 34 201 169 Henry M S 2013 Symbiosis Associations of Invertebrates Birds Ruminants and Other Biota New York New York Elsevier p 59 ISBN 978 1 4832 7592 5 a b c Krishna K Grimaldi D A Krishna V Engel M S 2013 Treatise on the Isoptera of the world PDF Bulletin of the American Museum of Natural History 1 377 7 1 200 doi 10 1206 377 1 S2CID 87276148 Bell W J Roth L M Nalepa C A 2007 Cockroaches ecology behavior and natural history Baltimore Md Johns Hopkins University Press p 161 ISBN 978 0 8018 8616 4 Engel M 2011 Family group names for termites Isoptera redux ZooKeys 148 171 184 doi 10 3897 zookeys 148 1682 PMC 3264418 PMID 22287896 Thorne Barbara L 1997 Evolution of eusociality in termites PDF Annual Review of Ecology and Systematics 28 5 27 53 doi 10 1146 annurev ecolsys 28 1 27 PMC 349550 Archived from the original PDF on 2010 05 30 a b Harrison Mark C Jongepier Evelien Robertson Hugh M Arning Nicolas Bitard Feildel Tristan Chao Hsu Childers Christopher P Dinh Huyen Doddapaneni Harshavardhan Dugan Shannon Gowin Johannes Greiner Carolin Han Yi Hu Haofu Hughes Daniel S T Huylmans Ann Kathrin Kemena Carsten Kremer Lukas P M Lee Sandra L Lopez Ezquerra Alberto Mallet Ludovic Monroy Kuhn Jose M Moser Annabell Murali Shwetha C Muzny Donna M Otani Saria Piulachs Maria Dolors Poelchau Monica Qu Jiaxin Schaub Florentine Wada Katsumata Ayako Worley Kim C Xie Qiaolin Ylla Guillem Poulsen Michael Gibbs Richard A Schal Coby Richards Stephen Belles Xavier Korb Judith Bornberg Bauer Erich 2018 Hemimetabolous genomes reveal molecular basis of termite eusociality Nature Ecology amp Evolution 2 3 557 566 doi 10 1038 s41559 017 0459 1 PMC 6482461 PMID 29403074 Termites had first castes Nature 530 7590 256 2016 Bibcode 2016Natur 530Q 256 doi 10 1038 530256a S2CID 49905391 Terrapon Nicolas Li Cai Robertson Hugh M Ji Lu Meng Xuehong Booth Warren Chen Zhensheng Childers Christopher P Glastad Karl M Gokhale Kaustubh et al 2014 Molecular traces of alternative social organization in a termite genome Nature Communications 5 3636 Bibcode 2014NatCo 5 3636T doi 10 1038 ncomms4636 PMID 24845553 Poulsen Michael Hu Haofu Li Cai Chen Zhensheng Xu Luohao Otani Saria Nygaard Sanne Nobre Tania Klaubauf Sylvia Schindler Philipp M et al 2014 Complementary symbiont contributions to plant decomposition in a fungus farming termite Proceedings of the National Academy of Sciences 111 40 14500 14505 Bibcode 2014PNAS 11114500P doi 10 1073 pnas 1319718111 PMC 4209977 PMID 25246537 Kohler T Dietrich C Scheffrahn RH Brune A 2012 High resolution analysis of gut environment and bacterial microbiota reveals functional compartmentation of the gut in wood feeding higher termites Nasutitermes spp Applied and Environmental Microbiology 78 13 4691 4701 Bibcode 2012ApEnM 78 4691K doi 10 1128 aem 00683 12 PMC 3370480 PMID 22544239 a b c d e Termite Biology and Ecology Division of Technology Industry and Economics Chemicals Branch United Nations Environment Programme Archived from the original on 10 November 2014 Retrieved 12 January 2015 Meyer V W Braack L E O Biggs H C Ebersohn C 1999 Distribution and density of termite mounds in the northern Kruger National Park with specific reference to those constructed by Macrotermes Holmgren Isoptera Termitidae African Entomology 7 1 123 130 Eggleton Paul 2020 The State of the World s Insects Annual Review of Environment and Resources 45 61 82 doi 10 1146 annurev environ 012420 050035 Sanderson M G 1996 Biomass of termites and their emissions of methane and carbon dioxide A global database Global Biogeochemical Cycles 10 4 543 557 Bibcode 1996GBioC 10 543S doi 10 1029 96GB01893 a b Heather N W 1971 The exotic drywood termite Cryptotermes brevis Walker Isoptera Kalotermitidae and endemic Australian drywood termites in Queensland Australian Journal of Entomology 10 2 134 141 doi 10 1111 j 1440 6055 1971 tb00022 x Claybourne Anna 2013 A colony of ants and other insect groups Chicago Ill Heinemann Library p 38 ISBN 978 1 4329 6487 0 a b Engel M S Gross M 2008 A giant termite from the Late Miocene of Styria Austria Isoptera Naturwissenschaften 96 2 289 295 Bibcode 2009NW 96 289E doi 10 1007 s00114 008 0480 y PMID 19052720 S2CID 21795900 Heidecker J L Leuthold R H 1984 The organisation of collective foraging in the harvester termite Hodotermes mossambicus Isoptera Behavioral Ecology and Sociobiology 14 3 195 202 doi 10 1007 BF00299619 S2CID 22158321 a b c d Costa Leonardo A M Haifig I 2010 Pheromones and exocrine glands in Isoptera Vitamins and Hormones 83 521 549 doi 10 1016 S0083 6729 10 83021 3 ISBN 9780123815163 PMID 20831960 a b c d Bignell Roisin amp Lo 2010 p 7 Bignell Roisin amp Lo 2010 pp 7 9 a b c d Bignell Roisin amp Lo 2010 p 11 Robinson W H 2005 Urban Insects and Arachnids A Handbook of Urban Entomology Cambridge Cambridge University Press p 291 ISBN 978 1 139 44347 0 Bignell Roisin amp Lo 2010 p 12 Riparbelli M G Dallai R Mercati D Bu Y Callaini G 2009 Centriole symmetry a big tale from small organisms Cell Motility and the Cytoskeleton 66 12 1100 5 doi 10 1002 cm 20417 PMID 19746415 Nalepa C A Lenz M 2000 The ootheca of Mastotermes darwiniensis Froggatt Isoptera Mastotermitidae homology with cockroach oothecae Proceedings of the Royal Society B Biological Sciences 267 1454 1809 1813 doi 10 1098 rspb 2000 1214 PMC 1690738 PMID 12233781 Crosland M W J Su N Y Scheffrahn R H 2005 Arolia in termites Isoptera functional significance and evolutionary loss Insectes Sociaux 52 1 63 66 doi 10 1007 s00040 004 0779 4 S2CID 26873138 Bignell Roisin amp Lo 2010 p 9 Bignell Roisin amp Lo 2010 p 10 a b c Bignell Roisin amp Lo 2010 p 13 a b c d e f g Termites Australian Museum Retrieved 8 January 2015 a b Machida M Kitade O Miura T Matsumoto T 2001 Nitrogen recycling through proctodeal trophallaxis in the Japanese damp wood termite Hodotermopsis japonica Isoptera Termopsidae Insectes Sociaux 48 1 52 56 doi 10 1007 PL00001745 ISSN 1420 9098 S2CID 21310420 Higashi Masahiko Yamamura Norio Abe Takuya Burns Thomas P 1991 10 22 Why don t all termite species have a sterile worker caste Proceedings of the Royal Society B Biological Sciences 246 1315 25 29 doi 10 1098 rspb 1991 0120 ISSN 0962 8452 PMID 1684665 S2CID 23067349 a b c Bignell Roisin amp Lo 2010 p 18 a b c d e f g h Krishna K Termite Encyclopaedia Britannica Retrieved 11 September 2015 Busvine J R 2013 Insects and Hygiene The biology and control of insect pests of medical and domestic importance 3rd ed Boston MA Springer US p 545 ISBN 978 1 4899 3198 6 Meek S P 1934 Termite Control at an Ordnance Storage Depot American Defense Preparedness Association p 159 a b Prestwich G D 1982 From tetracycles to macrocycles Tetrahedron 38 13 1911 1919 doi 10 1016 0040 4020 82 80040 9 Prestwich G D Bentley B L Carpenter E J 1980 Nitrogen sources for neotropical nasute termites Fixation and selective foraging Oecologia 46 3 397 401 Bibcode 1980Oecol 46 397P doi 10 1007 BF00346270 ISSN 1432 1939 PMID 28310050 S2CID 6134800 Thorne B L Breisch N L Muscedere M L 2003 10 28 Evolution of eusociality and the soldier caste in termites Influence of intraspecific competition and accelerated inheritance Proceedings of the National Academy of Sciences 100 22 12808 12813 Bibcode 2003PNAS 10012808T doi 10 1073 pnas 2133530100 ISSN 0027 8424 PMC 240700 PMID 14555764 a b c Horwood M A Eldridge R H 2005 Termites in New South Wales Part 1 Termite biology PDF Technical report Forest Resources Research ISSN 0155 7548 96 38 a b Keller L 1998 Queen lifespan and colony characteristics in ants and termites Insectes Sociaux 45 3 235 246 doi 10 1007 s000400050084 S2CID 24541087 Srinivasan Amia September 10 2018 What Termites Can Teach Us The New Yorker Archived from the original on March 7 2020 Korb J 2008 Termites hemimetabolous diploid white ants Frontiers in Zoology 5 1 15 doi 10 1186 1742 9994 5 15 PMC 2564920 PMID 18822181 Davis P Termite Identification Entomology at Western Australian Department of Agriculture Archived from the original on 2009 06 12 Neoh K B Lee C Y 2011 Developmental stages and caste composition of a mature and incipient colony of the drywood termite Cryptotermes dudleyi Isoptera Kalotermitidae Journal of Economic Entomology 104 2 622 8 doi 10 1603 ec10346 PMID 21510214 S2CID 23356632 a b Native subterranean termites University of Florida Retrieved 8 January 2015 a b Schneider M F 1999 Termite Life Cycle and Caste System University of Freiburg Retrieved 8 January 2015 Simpson S J Sword G A Lo N 2011 Polyphenism in Insects Current Biology 21 18 738 749 doi 10 1016 j cub 2011 06 006 PMID 21959164 S2CID 656039 a b Tasaki E Kobayashi K Matsuura K Iuchi Y 2017 An Efficient Antioxidant System in a Long Lived Termite Queen PLOS ONE 12 1 e0167412 Bibcode 2017PLoSO 1267412T doi 10 1371 journal pone 0167412 PMC 5226355 PMID 28076409 a b Miller D M 5 March 2010 Subterranean Termite Biology and Behavior Virginia Tech Virginia State University Retrieved 8 January 2015 a b Gouge D H Smith K A Olson C Baker P 2001 Drywood Termites Cooperative Extension College of Agriculture amp Life Sciences University of Arizona Retrieved 16 September 2015 Kaib M Hacker M Brandl R 2001 Egg laying in monogynous and polygynous colonies of the termite Macrotermes michaelseni Isoptera Macrotermitidae Insectes Sociaux 48 3 231 237 doi 10 1007 PL00001771 S2CID 35656795 Gilbert executive editors G A Kerkut L I 1985 Comprehensive insect physiology biochemistry and pharmacology 1st ed Oxford Pergamon Press p 167 ISBN 978 0 08 026850 7 a href wiki Template Cite book title Template Cite book cite book a first1 has generic name help Wyatt T D 2003 Pheromones and animal behaviour communication by smell and taste Repr with corrections 2004 ed Cambridge Cambridge University Press p 119 ISBN 978 0 521 48526 5 Morrow E H 2004 How the sperm lost its tail the evolution of aflagellate sperm Biological Reviews of the Cambridge Philosophical Society 79 4 795 814 doi 10 1017 S1464793104006451 PMID 15682871 S2CID 25878093 Supplementary reproductive University of Hawaii Archived from the original on 30 October 2014 Retrieved 16 September 2015 Yashiro T Matsuura K 2014 Termite queens close the sperm gates of eggs to switch from sexual to asexual reproduction Proceedings of the National Academy of Sciences 111 48 17212 17217 Bibcode 2014PNAS 11117212Y doi 10 1073 pnas 1412481111 PMC 4260566 PMID 25404335 Matsuura K Vargo E L Kawatsu K Labadie P E Nakano H Yashiro T Tsuji K 2009 Queen Succession Through Asexual Reproduction in Termites Science 323 5922 1687 Bibcode 2009Sci 323 1687M doi 10 1126 science 1169702 PMID 19325106 S2CID 21785886 Fougeyrollas R Dolejsova K Sillam Dusses D Roy V Poteaux C Hanus R Roisin Y June 2015 Asexual queen succession in the higher termite Embiratermes neotenicus Proc Biol Sci 282 1809 20150260 doi 10 1098 rspb 2015 0260 PMC 4590441 PMID 26019158 Bignell Roisin amp Lo 2010 pp 13 14 Freymann B P Buitenwerf R Desouza O Olff 2008 The importance of termites Isoptera for the recycling of herbivore dung in tropical ecosystems a review European Journal of Entomology 105 2 165 173 doi 10 14411 eje 2008 025 de Souza O F Brown V K 2009 Effects of habitat fragmentation on Amazonian termite communities Journal of Tropical Ecology 10 2 197 206 doi 10 1017 S0266467400007847 Tokuda G Watanabe H Matsumoto T Noda H 1997 Cellulose digestion in the wood eating higher termite Nasutitermes takasagoensis Shiraki distribution of cellulases and properties of endo beta 1 4 glucanase Zoological Science 14 1 83 93 doi 10 2108 zsj 14 83 PMID 9200983 S2CID 2877588 Ritter Michael 2006 The Physical Environment an Introduction to Physical Geography University of Wisconsin p 450 Archived from the original on 18 May 2007 Ikeda Ohtsubo W Brune A 2009 Cospeciation of termite gut flagellates and their bacterial endosymbionts Trichonympha species and Candidatus Endomicrobium trichonymphae Molecular Ecology 18 2 332 342 doi 10 1111 j 1365 294X 2008 04029 x PMID 19192183 S2CID 28048145 Slaytor M 1992 Cellulose digestion in termites and cockroaches What role do symbionts play Comparative Biochemistry and Physiology B 103 4 775 784 doi 10 1016 0305 0491 92 90194 V The Termite Gut and its Symbiotic Microbes iBiology Retrieved 2020 05 16 Watanabe H Noda H Tokuda G Lo N 1998 A cellulase gene of termite origin Nature 394 6691 330 331 Bibcode 1998Natur 394 330W doi 10 1038 28527 PMID 9690469 S2CID 4384555 Tokuda G Watanabe H 2007 Hidden cellulases in termites revision of an old hypothesis Biology Letters 3 3 336 339 doi 10 1098 rsbl 2007 0073 PMC 2464699 PMID 17374589 Li Z Q Liu B R Zeng W H Xiao W L Li Q J Zhong J H 2013 Character of Cellulase Activity in the Guts of Flagellate Free Termites with Different Feeding Habits Journal of Insect Science 13 37 37 doi 10 1673 031 013 3701 PMC 3738099 PMID 23895662 The Termite Gut and its Symbiotic Microbes iBiology Retrieved 2020 05 16 Geetha Iyer Scroll in Mar 09 2017 Why Indians worship the mound of the much hated termite The soldier termites and the reproductive castes obtain their nutrients from the workers through oral or anal trophallaxis Dietrich C Kohler T Brune A 2014 The Cockroach origin of the termite gut microbiota patterns in bacterial community structure reflect major evolutionary events Applied and Environmental Microbiology 80 7 2261 2269 Bibcode 2014ApEnM 80 2261D doi 10 1128 AEM 04206 13 PMC 3993134 PMID 24487532 Allen C T Foster D E Ueckert D N 1980 Seasonal Food Habits of a Desert Termite Gnathamitermes tubiformans in West Texas Environmental Entomology 9 4 461 466 doi 10 1093 ee 9 4 461 McMahan E A 1966 Studies of Termite Wood feeding Preferences PDF Hawaiian Entomological Society 19 2 239 250 ISSN 0073 134X Aanen D K Eggleton P Rouland Lefevre C Guldberg Froslev T Rosendahl S Boomsma J J 2002 The evolution of fungus growing termites and their mutualistic fungal symbionts Proceedings of the National Academy of Sciences 99 23 14887 14892 Bibcode 2002PNAS 9914887A doi 10 1073 pnas 222313099 JSTOR 3073687 PMC 137514 PMID 12386341 Mueller U G Gerardo N 2002 Fungus farming insects Multiple origins and diverse evolutionary histories Proceedings of the National Academy of Sciences 99 24 15247 15249 Bibcode 2002PNAS 9915247M doi 10 1073 pnas 242594799 PMC 137700 PMID 12438688 Roberts E M Todd C N Aanen D K Nobre T Hilbert Wolf H L O Connor P M Tapanila L Mtelela C Stevens N J 2016 Oligocene termite nests with in situ fungus gardens from the Rukwa Rift Basin Tanzania support a paleogene African origin for insect agriculture PLOS ONE 11 6 e0156847 Bibcode 2016PLoSO 1156847R doi 10 1371 journal pone 0156847 PMC 4917219 PMID 27333288 Radek R 1999 Flagellates bacteria and fungi associated with termites diversity and function in nutrition a review PDF Ecotropica 5 183 196 Breznak J A Brune A 1993 Role of microorganisms in the digestion of lignocellulose by termites Annual Review of Entomology 39 1 453 487 doi 10 1146 annurev en 39 010194 002321 Kok O B Hewitt P H 1990 Bird and mammal predators of the harvester termite Hodotermes mossambicus Hagen in semi arid regions of South Africa South African Journal of Science 86 1 34 37 ISSN 0038 2353 a b c d e Holldobler B Wilson E O 1990 The Ants Cambridge Massachusetts Belknap Press of Harvard University Press pp 559 566 ISBN 978 0 674 04075 5 a b Culliney T W Grace J K 2000 Prospects for the biological control of subterranean termites Isoptera Rhinotermitidae with special reference to Coptotermes formosanus Bulletin of Entomological Research 90 1 9 21 doi 10 1017 S0007485300000663 PMID 10948359 Dean W R J Milton S J 1995 Plant and invertebrate assemblages on old fields in the arid southern Karoo South Africa African Journal of Ecology 33 1 1 13 doi 10 1111 j 1365 2028 1995 tb00777 x Wade W W 2002 Ecology of Desert Systems Burlington Elsevier p 216 ISBN 978 0 08 050499 5 Reagan D P Waide R B 1996 The food web of a tropical rain forest Chicago University of Chicago Press p 294 ISBN 978 0 226 70599 6 a b Bardgett R D Herrick J E Six J Jones T H Strong D R van der Putten W H 2013 Soil ecology and ecosystem services 1st ed Oxford Oxford University Press p 178 ISBN 978 0 19 968816 6 Bignell Roisin amp Lo 2010 p 509 Choe J C Crespi B J 1997 The evolution of social behavior in insects and arachnids 1st ed Cambridge Cambridge university press p 76 ISBN 978 0 521 58977 2 a b c Abe Y Bignell D E Higashi T 2014 Termites Evolution Sociality Symbioses Ecology Springer pp 124 149 doi 10 1007 978 94 017 3223 9 ISBN 978 94 017 3223 9 S2CID 30804981 Wilson D S Clark A B 1977 Above ground defence in the harvester termite Hodotermes mossambicus Journal of the Entomological Society of South Africa 40 271 282 Lavelle P Spain A V 2001 Soil ecology 2nd ed Dordrecht Kluwer Academic p 316 ISBN 978 0 306 48162 8 Richardson P K R Bearder S K 1984 The Hyena Family In MacDonald D ed The Encyclopedia of Mammals New York NY Facts on File Publication pp 158 159 ISBN 978 0 87196 871 5 Mills G Harvey M 2001 African Predators Washington D C Smithsonian Institution Press p 71 ISBN 978 1 56098 096 4 d Errico F Backwell L 2009 Assessing the function of early hominin bone tools Journal of Archaeological Science 36 8 1764 1773 doi 10 1016 j jas 2009 04 005 Lepage M G 1981 Etude de la predation de Megaponera foetens F sur les populations recoltantes de Macrotermitinae dans un ecosysteme semi aride Kajiado Kenya Insectes Sociaux in Spanish 28 3 247 262 doi 10 1007 BF02223627 S2CID 28763771 Levieux J 1966 Note preliminaire sur les colonnes de chasse de Megaponera fœtens F Hymenoptere Formicidae Insectes Sociaux in French 13 2 117 126 doi 10 1007 BF02223567 S2CID 2031222 Longhurst C Baker R Howse P E 1979 Chemical crypsis in predatory ants Experientia 35 7 870 872 doi 10 1007 BF01955119 S2CID 39854106 a b Wheeler W M 1936 Ecological relations of Ponerine and other ants to termites Proceedings of the American Academy of Arts and Sciences 71 3 159 171 doi 10 2307 20023221 JSTOR 20023221 Shattuck S O Heterick B E 2011 Revision of the ant genus Iridomyrmex Hymenoptera Formicidae PDF Zootaxa 2845 1 74 doi 10 11646 zootaxa 2743 1 1 ISBN 978 1 86977 676 3 ISSN 1175 5334 Traniello J F A 1981 Enemy deterrence in the recruitment strategy of a termite Soldier organized foraging in Nasutitermes costalis Proceedings of the National Academy of Sciences 78 3 1976 1979 Bibcode 1981PNAS 78 1976T doi 10 1073 pnas 78 3 1976 PMC 319259 PMID 16592995 Schoning C Moffett M W 2007 Driver Ants Invading a Termite Nest why do the most catholic predators of all seldom take this abundant prey PDF Biotropica 39 5 663 667 doi 10 1111 j 1744 7429 2007 00296 x Archived from the original PDF on 2015 11 12 Retrieved 2015 09 20 Mill A E 1983 Observations on Brazilian termite alate swarms and some structures used in the dispersal of reproductives Isoptera Termitidae Journal of Natural History 17 3 309 320 doi 10 1080 00222938300770231 Schmid Hempel 1998 p 61 Schmid Hempel 1998 p 75 Wilson E O 1971 The Insect Societies Vol 76 5th ed Cambridge Massachusetts Belknap Press of Harvard University Press p 398 ISBN 978 0 674 45495 8 Schmid Hempel 1998 p 59 Schmid Hempel 1998 pp 301 302 Schmid Hempel 1998 p 19 Weiser J Hrdy I 2009 Pyemotes mites as parasites of termites Zeitschrift fur Angewandte Entomologie 51 1 4 94 97 doi 10 1111 j 1439 0418 1962 tb04062 x Chouvenc T Efstathion C A Elliott M L Su N Y 2012 Resource competition between two fungal parasites in subterranean termites Die Naturwissenschaften 99 11 949 58 Bibcode 2012NW 99 949C doi 10 1007 s00114 012 0977 2 PMID 23086391 S2CID 16393629 Schmid Hempel 1998 pp 38 102 Wilson Megan Barden Phillip Ware Jessica 2021 04 30 A Review of Ectoparasitic Fungi Associated With Termites Annals of the Entomological Society of America 114 4 373 396 doi 10 1093 aesa saab001 Retrieved 7 May 2021 Chouvenc T Mullins A J Efstathion C A Su N Y 2013 Virus like symptoms in a termite Isoptera Kalotermitidae field colony Florida Entomologist 96 4 1612 1614 doi 10 1653 024 096 0450 S2CID 73570814 Al Fazairy A A Hassan F A 2011 Infection of Termites by Spodoptera littoralis Nuclear Polyhedrosis Virus International Journal of Tropical Insect Science 9 1 37 39 doi 10 1017 S1742758400009991 S2CID 84743428 Traniello J F A Leuthold R H 2000 Behavior and Ecology of Foraging in Termites Springer Netherlands pp 141 168 doi 10 1007 978 94 017 3223 9 7 ISBN 978 94 017 3223 9 Reinhard J Kaib M 2001 Trail communication during foraging and recruitment in the subterranean termite Reticulitermes santonensis De Feytaud Isoptera Rhinotermitidae Journal of Insect Behavior 14 2 157 171 doi 10 1023 A 1007881510237 S2CID 40887791 a b c d e f g h i j Costa Leonardo A M Haifig I 2013 Termite communication during different behavioral activitiesin Biocommunication of Animals Springer Netherlands pp 161 190 doi 10 1007 978 94 007 7414 8 10 ISBN 978 94 007 7413 1 Costa Leonardo A M 2006 Morphology of the sternal gland in workers of Coptotermes gestroi Isoptera Rhinotermitidae Micron 37 6 551 556 doi 10 1016 j micron 2005 12 006 PMID 16458523 Traniello J F Busher C 1985 Chemical regulation of polyethism during foraging in the neotropical termite Nasutitermes costalis Journal of Chemical Ecology 11 3 319 32 doi 10 1007 BF01411418 PMID 24309963 S2CID 27799126 Miramontes O DeSouza O Paiva L R Marins A Orozco S Aegerter C M 2014 Levy flights and self similar exploratory behaviour of termite workers beyond model fitting PLOS ONE 9 10 e111183 arXiv 1410 0930 Bibcode 2014PLoSO 9k1183M doi 10 1371 journal pone 0111183 PMC 4213025 PMID 25353958 Jost C Haifig I de Camargo Dietrich C R R Costa Leonardo A M 2012 A comparative tunnelling network approach to assess interspecific competition effects in termites Insectes Sociaux 59 3 369 379 doi 10 1007 s00040 012 0229 7 S2CID 14885485 Polizzi J M Forschler B T 1998 Intra and interspecific agonism in Reticulitermes flavipes Kollar and R virginicus Banks and effects of arena and group size in laboratory assays Insectes Sociaux 45 1 43 49 doi 10 1007 s000400050067 S2CID 36235510 Darlington J P E C 1982 The underground passages and storage pits used in foraging by a nest of the termite Macrotermes michaelseni in Kajiado Kenya Journal of Zoology 198 2 237 247 doi 10 1111 j 1469 7998 1982 tb02073 x Cornelius M L Osbrink W L 2010 Effect of soil type and moisture availability on the foraging behavior of the Formosan subterranean termite Isoptera Rhinotermitidae Journal of Economic Entomology 103 3 799 807 doi 10 1603 EC09250 PMID 20568626 S2CID 23173060 Toledo Lima J Costa Leonardo A M 2012 Subterranean termites Isoptera Rhinotermitidae Exploitation of equivalent food resources with different forms of placement Insect Science 19 3 412 418 doi 10 1111 j 1744 7917 2011 01453 x S2CID 82046133 Jmhasly P Leuthold R H 1999 Intraspecific colony recognition in the termites Macrotermes subhyalinus and Macrotermes bellicosus Isoptera Termitidae Insectes Sociaux 46 2 164 170 doi 10 1007 s000400050128 S2CID 23037986 Messenger M T Su N Y 2005 Agonistic behavior between colonies of the Formosan subterranean termite Isoptera Rhinotermitidae from Louis Armstrong Park New Orleans Louisiana Sociobiology 45 2 331 345 Korb J Weil T Hoffmann K Foster K R Rehli M 2009 A gene necessary for reproductive suppression in termites Science 324 5928 758 Bibcode 2009Sci 324 758K doi 10 1126 science 1170660 PMID 19423819 S2CID 31608071 a b c Mathew T T G Reis R DeSouza O Ribeiro S P 2005 Predation and interference competition between ants Hymenoptera Formicidae and arboreal termites Isoptera Termitidae PDF Sociobiology 46 2 409 419 Evans T A Inta R Lai J C S Lenz M 2007 Foraging vibration signals attract foragers and identify food size in the drywood termite Cryptotermes secundus Insectes Sociaux 54 4 374 382 doi 10 1007 s00040 007 0958 1 S2CID 40214049 Costa Leonardo A M Casarin F E Lima J T 2009 Chemical communication in isoptera Neotropical Entomology 38 1 747 52 doi 10 1590 S1519 566X2009000100001 PMID 19347093 Richard F J Hunt J H 2013 Intracolony chemical communication in social insects PDF Insectes Sociaux 60 3 275 291 doi 10 1007 s00040 013 0306 6 S2CID 8108234 Archived from the original PDF on 2016 03 04 Retrieved 2015 10 08 Dronnet S Lohou C Christides J P Bagneres A G 2006 Cuticular hydrocarbon composition reflects genetic relationship among colonies of the introduced termite Reticulitermes santonensis Feytaud Journal of Chemical Ecology 32 5 1027 1042 doi 10 1007 s10886 006 9043 x PMID 16739021 S2CID 23956394 Rosengaus R B Traniello J F A Chen T Brown J J Karp R D 1999 Immunity in a social insect Naturwissenschaften 86 12 588 591 Bibcode 1999NW 86 588R doi 10 1007 s001140050679 S2CID 10769345 Wilson D S 1977 Above ground predator defense in the harvester termite Hodotermes mossambicus Hagen Journal of the Entomological Society of Southern Africa 40 271 282 Belbin R M 2013 The Coming Shape of Organization New York Routledge p 27 ISBN 978 1 136 01553 3 a b Wilson E O 2014 A window on eternity a biologist s walk through Gorongosa National Park First ed Simon amp Schuster pp 85 90 ISBN 978 1 4767 4741 5 Miura T Matsumoto T 2000 Soldier morphogenesis in a nasute termite discovery of a disc like structure forming a soldier nasus Proceedings of the Royal Society B Biological Sciences 267 1449 1185 1189 doi 10 1098 rspb 2000 1127 PMC 1690655 PMID 10902684 Prestwich G D Chen D 1981 Soldier defense secretions of Trinervitermes bettonianus Isoptera Nasutitermitinae Chemical variation in allopatric populations Journal of Chemical Ecology 7 1 147 157 doi 10 1007 BF00988642 PMID 24420434 S2CID 27654745 Chen J Henderson G Grimm C C Lloyd S W Laine R A 1998 04 09 Termites fumigate their nests with naphthalene Nature 392 6676 558 559 Bibcode 1998Natur 392 558C doi 10 1038 33305 S2CID 4419882 Piper Ross 2007 Extraordinary Animals An Encyclopedia of Curious and Unusual Animals Greenwood Press p 26 ISBN 978 0 313 33922 6 Bordereau C Robert A Van Tuyen V Peppuy A 1997 Suicidal defensive behaviour by frontal gland dehiscence in Globitermes sulphureus Haviland soldiers Isoptera Insectes Sociaux 44 3 289 297 doi 10 1007 s000400050049 S2CID 19770804 Sobotnik J Bourguignon T Hanus R Demianova Z Pytelkova J Mares M Foltynova P Preisler J Cvacka J Krasulova J Roisin Y 2012 Explosive backpacks in old termite workers Science 337 6093 436 Bibcode 2012Sci 337 436S doi 10 1126 science 1219129 PMID 22837520 S2CID 206540025 SobotnIk J Bourguignon T Hanus R Weyda F Roisin Y 2010 Structure and function of defensive glands in soldiers of Glossotermes oculatus Isoptera Serritermitidae Biological Journal of the Linnean Society 99 4 839 848 doi 10 1111 j 1095 8312 2010 01392 x Ulyshen M D Shelton T G 2011 Evidence of cue synergism in termite corpse response behavior Naturwissenschaften 99 2 89 93 Bibcode 2012NW 99 89U doi 10 1007 s00114 011 0871 3 PMID 22167071 S2CID 2616753 Su N Y 2005 Response of the Formosan subterranean termites Isoptera Rhinotermitidae to baits or nonrepellent termiticides in extended foraging arenas Journal of Economic Entomology 98 6 2143 2152 doi 10 1603 0022 0493 98 6 2143 PMID 16539144 S2CID 196618597 Sun Q Haynes K F Zhou X 2013 Differential undertaking response of a lower termite to congeneric and conspecific corpses Scientific Reports 3 1650 Bibcode 2013NatSR 3E1650S doi 10 1038 srep01650 PMC 3629736 PMID 23598990 a b Neoh K B Yeap B K Tsunoda K Yoshimura T Lee C Y Korb J 2012 Do termites avoid carcasses behavioral responses depend on the nature of the carcasses PLOS ONE 7 4 e36375 Bibcode 2012PLoSO 736375N doi 10 1371 journal pone 0036375 PMC 3338677 PMID 22558452 Matsuura K 2006 Termite egg mimicry by a sclerotium forming fungus Proceedings of the Royal Society B Biological Sciences 273 1591 1203 1209 doi 10 1098 rspb 2005 3434 PMC 1560272 PMID 16720392 Matsuura K Yashiro T Shimizu K Tatsumi S Tamura T 2009 Cuckoo fungus mimics termite eggs by producing the cellulose digesting enzyme b glucosidase Current Biology 19 1 30 36 doi 10 1016 j cub 2008 11 030 PMID 19110429 S2CID 18604426 Howard R W McDaniel C A Blomquist G J 1980 Chemical mimicry as an integrating mechanism cuticular hydrocarbons of a termitophile and its host Science 210 4468 431 433 Bibcode 1980Sci 210 431H doi 10 1126 science 210 4468 431 PMID 17837424 S2CID 33221252 Watson J A L 1973 Austrospirachtha mimetes a new termitophilous corotocine from Northern Australia Coleoptera Staphylinidae Australian Journal of Entomology 12 4 307 310 doi 10 1111 j 1440 6055 1973 tb01678 x Forbes H O 1878 Termites Kept in Captivity by Ants Nature 19 471 4 5 Bibcode 1878Natur 19 4F doi 10 1038 019004b0 S2CID 4125839 subscription required Darlington J 1985 Attacks by doryline ants and termite nest defences Hymenoptera Formicidae Isoptera Termitidae Sociobiology 11 189 200 Quinet Y Tekule N amp de Biseau JC 2005 Behavioural Interactions Between Crematogaster brevispinosa rochai Forel Hymenoptera Formicidae and Two Nasutitermes Species Isoptera Termitidae Journal of Insect Behavior 18 1 1 17 doi 10 1007 s10905 005 9343 y S2CID 33487814 Coty D Aria C Garrouste R Wils P Legendre F Nel A Korb J 2014 The First Ant Termite Syninclusion in Amber with CT Scan Analysis of Taphonomy PLOS ONE 9 8 e104410 Bibcode 2014PLoSO 9j4410C doi 10 1371 journal pone 0104410 PMC 4139309 PMID 25140873 a b Santos P P Vasconcellos A Jahyny B Delabie J H C 2010 Ant fauna Hymenoptera Formicidae associated to arboreal nests of Nasutitermes spp Isoptera Termitidae in a cacao plantation in southeastern Bahia Brazil Revista Brasileira de Entomologia 54 3 450 454 doi 10 1590 S0085 56262010000300016 Jaffe K Ramos C Issa S 1995 Trophic Interactions Between Ants and Termites that Share Common Nests Annals of the Entomological Society of America 88 3 328 333 doi 10 1093 aesa 88 3 328 Trager J C 1991 A Revision of the fire ants Solenopsis geminata group Hymenoptera Formicidae Myrmicinae Journal of the New York Entomological Society 99 2 141 198 doi 10 5281 zenodo 24912 JSTOR 25009890 a b Cingel N A van der 2001 An atlas of orchid pollination America Africa Asia and Australia Rotterdam Balkema p 224 ISBN 978 90 5410 486 5 McHatton R 2011 Orchid Pollination exploring a fascinating world PDF The American Orchid Society p 344 Retrieved 5 September 2015 Cowie R 2014 Journey to a Waterfall a biologist in Africa Raleigh North Carolina Lulu Press p 169 ISBN 978 1 304 66939 1 a b Tan K H 2009 Environmental Soil Science 3rd ed Boca Raton Florida CRC Press pp 105 106 ISBN 978 1 4398 9501 6 a b Clark Sarah 15 November 2005 Plant extract stops termites dead ABC Archived from the original on 15 June 2009 Retrieved 8 February 2014 Vasconcellos Alexandre Bandeira Adelmar G Moura Flavia Maria S Araujo Virginia Farias P Gusmao Maria Avany B Reginaldo Constantino February 2010 Termite assemblages in three habitats under different disturbance regimes in the semi arid Caatinga of NE Brazil Journal of Arid Environments Elsevier 74 2 298 302 Bibcode 2010JArEn 74 298V doi 10 1016 j jaridenv 2009 07 007 ISSN 0140 1963 Bignell Roisin amp Lo 2010 p 3 a b Noirot C Darlington J P E C 2000 Termite Nests Architecture Regulation and Defencein Termites Evolution Sociality Symbioses Ecology Springer pp 121 139 doi 10 1007 978 94 017 3223 9 6 ISBN 978 94 017 3223 9 Bignell Roisin amp Lo 2010 p 20 a b Eggleton P Bignell D E Sands W A Mawdsley N A Lawton J H Wood T G Bignell N C 1996 The Diversity Abundance and Biomass of Termites under Differing Levels of Disturbance in the Mbalmayo Forest Reserve Southern Cameroon Philosophical Transactions of the Royal Society B Biological Sciences 351 1335 51 68 Bibcode 1996RSPTB 351 51E doi 10 1098 rstb 1996 0004 a b c d e Bignell Roisin amp Lo 2010 p 21 De Visse S N Freymann B P Schnyder H 2008 Trophic interactions among invertebrates in termitaria in the African savanna a stable isotope approach Ecological Entomology 33 6 758 764 doi 10 1111 j 1365 2311 2008 01029 x S2CID 33877331 a b c Bignell Roisin amp Lo 2010 p 22 Vane C H Kim A W Moss Hayes V Snape C E Diaz M C Khan N S Engelhart S E Horton B P 2013 Degradation of mangrove tissues by arboreal termites Nasutitermes acajutlae and their role in the mangrove C cycle Puerto Rico Chemical characterization and organic matter provenance using bulk d13C C N alkaline CuO oxidation GC MS and solid state PDF Geochemistry Geophysics Geosystems 14 8 3176 3191 Bibcode 2013GGG 14 3176V doi 10 1002 ggge 20194 a b Roisin Y Pasteels J M 1986 Reproductive mechanisms in termites Polycalism and polygyny in Nasutitermes polygynus and N costalis Insectes Sociaux 33 2 149 167 doi 10 1007 BF02224595 S2CID 41799894 Perna A Jost C Couturier E Valverde S Douady S Theraulaz G 2008 The structure of gallery networks in the nests of termite Cubitermes spp revealed by X ray tomography Die Naturwissenschaften 95 9 877 884 Bibcode 2008NW 95 877P doi 10 1007 s00114 008 0388 6 PMID 18493731 S2CID 15326313 Glenday Craig 2014 Guinness World Records 2014 pp 33 ISBN 978 1 908843 15 9 Jacklyn P 1991 Evidence for Adaptive Variation in the Orientation of Amitermes Isoptera Termitinae Mounds From Northern Australia Australian Journal of Zoology 39 5 569 doi 10 1071 ZO9910569 Jacklyn P M Munro U 2002 Evidence for the use of magnetic cues in mound construction by the termite Amitermes meridionalis Isoptera Termitinae Australian Journal of Zoology 50 4 357 doi 10 1071 ZO01061 Grigg G C 1973 Some Consequences of the Shape and Orientation of magnetic Termite Mounds PDF Australian Journal of Zoology 21 2 231 237 doi 10 1071 ZO9730231 a b Hadlington P 1996 Australian Termites and Other Common Timber Pests 2nd ed Kensington NSW Australia New South Wales University Press pp 28 30 ISBN 978 0 86840 399 1 a b Kahn L Easton B 2010 Shelter II Bolinas California Shelter Publications p 198 ISBN 978 0 936070 49 0 a b c d e f g h Su N Y Scheffrahn R H 2000 Termites as Pests of Buildingsin Termites Evolution Sociality Symbioses Ecology Springer Netherlands pp 437 453 doi 10 1007 978 94 017 3223 9 20 ISBN 978 94 017 3223 9 Thorne Ph D Barbara L 1999 NPMA Research Report On Subterranean Termites Dunn Loring VA NPMA p 22 Termites Victorian Building Authority Government of Victoria 2014 Archived from the original on 3 February 2018 Retrieved 20 September 2015 Thorne Ph D Barbara L 1999 NPMA Research Report On Subterranean Termites Dunn Loring VA NPMA p 2 Grace J K Cutten G M Scheffrahn R H McEkevan D K 1991 First infestation by Incisitermes minor of a Canadian building Isoptera Kalotermitidae Sociobiology 18 299 304 a b Sands W A 1973 Termites as Pests of Tropical Food Crops Tropical Pest Management 19 2 167 177 doi 10 1080 09670877309412751 a b c d Flores A 17 February 2010 New Assay Helps Track Termites Other Insects Agricultural Research Service United States Department of Agriculture Retrieved 15 January 2015 Su N Y Scheffrahn R H 1990 Economically important termites in the United States and their control PDF Sociobiology 17 77 94 Archived from the original PDF on 2011 08 12 Thorne Ph D Barbara L 1999 NPMA Research Report On Subterranean Termites Dunn Loring VA NPMA p 40 Elliott Sara 26 May 2009 How can copper keep termites at bay HowStuffWorks Questions and Answers About Termites PDF Department of Consumer Affairs Structural Pest Control Board of California Retrieved 19 April 2021 EPA Registration and Label for Taurus SC Termiticide PDF EPA gov EPA Registration and Label for Termidor SC PDF EPA gov Retrieved 19 April 2021 Pidd Helen 21 December 2021 A world first Devon calls victory in 27 year war on termites The Guardian Archived from the original on 22 December 2021 Retrieved 22 December 2021 a b c Figueiredo R E C R Vasconcellos A Policarpo I S Alves R R N 2015 Edible and medicinal termites a global overview Journal of Ethnobiology and Ethnomedicine 11 1 29 doi 10 1186 s13002 015 0016 4 PMC 4427943 PMID 25925503 a b c d Nyakupfuka A 2013 Global Delicacies Discover Missing Links from Ancient Hawaiian Teachings to Clean the Plaque of your Soul and Reach Your Higher Self Bloomington Indiana BalboaPress pp 40 41 ISBN 978 1 4525 6791 4 a b Bodenheimer F S 1951 Insects as Human Food A Chapter of the Ecology of Man Netherlands Springer pp 331 350 ISBN 978 94 017 6159 8 Geissler P W 2011 The significance of earth eating social and cultural aspects of geophagy among Luo children Africa 70 4 653 682 doi 10 3366 afr 2000 70 4 653 S2CID 145754470 Knudsen J W 2002 Akula udongo earth eating habit a social and cultural practice among Chagga women on the slopes of Mount Kilimanjaro African Journal of Indigenous Knowledge Systems 1 1 19 26 doi 10 4314 indilinga v1i1 26322 ISSN 1683 0296 OCLC 145403765 Nchito M Wenzel Geissler P Mubila L Friis H Olsen A 2004 Effects of iron and multimicronutrient supplementation on geophagy a two by two factorial study among Zambian schoolchildren in Lusaka Transactions of the Royal Society of Tropical Medicine and Hygiene 98 4 218 227 doi 10 1016 S0035 9203 03 00045 2 PMID 15049460 Saathoff E Olsen A Kvalsvig J D Geissler P W 2002 Geophagy and its association with geohelminth infection in rural schoolchildren from northern KwaZulu Natal South Africa Transactions of the Royal Society of Tropical Medicine and Hygiene 96 5 485 490 doi 10 1016 S0035 9203 02 90413 X PMID 12474473 Katayama N Ishikawa Y Takaoki M Yamashita M Nakayama S Kiguchi K Kok R Wada H Mitsuhashi J 2008 Entomophagy A key to space agriculture PDF Advances in Space Research 41 5 701 705 Bibcode 2008AdSpR 41 701S doi 10 1016 j asr 2007 01 027 Mitchell J D 2002 Termites as pests of crops forestry rangeland and structures in Southern Africa and their control Sociobiology 40 1 47 69 ISSN 0361 6525 Loffler E Kubiniok J 1996 Landform development and bioturbation on the Khorat plateau Northeast Thailand PDF Natural History Bulletin of the Siam Society 44 199 216 Evans T A Dawes T Z Ward P R Lo N 2011 Ants and termites increase crop yield in a dry climate Nature Communications 2 262 Bibcode 2011NatCo 2 262E doi 10 1038 ncomms1257 PMC 3072065 PMID 21448161 a b c d e Termite Power DOE Joint Genome Institute United States Department of Energy 14 August 2006 Archived from the original on 22 September 2006 Retrieved 11 September 2015 a href wiki Template Cite web title Template Cite web cite web a CS1 maint unfit URL link Hirschler B 22 November 2007 Termites gut reaction set for biofuels ABC News Retrieved 8 January 2015 Roach J 14 March 2006 Termite Power Can Pests Guts Create New Fuel National Geographic News Retrieved 11 September 2015 Werfel J Petersen K Nagpal R 2014 Designing Collective Behavior in a Termite Inspired Robot Construction Team Science 343 6172 754 758 Bibcode 2014Sci 343 754W doi 10 1126 science 1245842 PMID 24531967 S2CID 38776920 Gibney E 2014 Termite inspired robots build castles Nature doi 10 1038 nature 2014 14713 S2CID 112117767 a b c Termites Green Architecture in the Tropics The Architect Architectural Association of Kenya Archived from the original on 22 March 2016 Retrieved 17 October 2015 Tan A Wong N 2013 Parameterization Studies of Solar Chimneys in the Tropics Energies 6 1 145 163 doi 10 3390 en6010145 Tsoroti S 15 May 2014 What s that building Eastgate Mall Harare News Archived from the original on 11 April 2021 Retrieved 8 January 2015 Im Zoo Basel fliegen die Termiten aus Neue Zurcher Zeitung in German 8 February 2014 Retrieved 21 May 2011 Van Huis H 2003 Insects as food in Sub Saharan Africa PDF Insect Science and Its Application 23 3 163 185 doi 10 1017 s1742758400023572 a b Neoh K B 2013 Termites and human society in Southeast Asia PDF The Newsletter 30 66 1 2 Cited literature a, wikipedia, wiki, book,

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