Termite

Termite

Overview
Termites are a group of eusocial
Eusociality
Eusociality is a term used for the highest level of social organization in a hierarchical classification....

 insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...

s that, until recently, were classified at the taxonomic rank
Taxonomy
Taxonomy is the science of identifying and naming species, and arranging them into a classification. The field of taxonomy, sometimes referred to as "biological taxonomy", revolves around the description and use of taxonomic units, known as taxa...

 of order
Order (biology)
In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family...

 Isoptera (see taxonomy below), but are now accepted as the epifamily Termitoidae, of the cockroach order Blattodea. While termites are commonly known, especially in Australia
Australia
Australia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...

, as "white ants," they are only distantly related to the ants.

Like ant
Ant
Ants are social insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. More than...

s, some bee
Bee
Bees are flying insects closely related to wasps and ants, and are known for their role in pollination and for producing honey and beeswax. Bees are a monophyletic lineage within the superfamily Apoidea, presently classified by the unranked taxon name Anthophila...

s, and wasp
Wasp
The term wasp is typically defined as any insect of the order Hymenoptera and suborder Apocrita that is neither a bee nor an ant. Almost every pest insect species has at least one wasp species that preys upon it or parasitizes it, making wasps critically important in natural control of their...

s—which are all placed in the separate order Hymenoptera
Hymenoptera
Hymenoptera is one of the largest orders of insects, comprising the sawflies, wasps, bees and ants. There are over 130,000 recognized species, with many more remaining to be described. The name refers to the heavy wings of the insects, and is derived from the Ancient Greek ὑμήν : membrane and...

—termites divide labour among castes, produce overlapping generations and take care of young collectively.
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Encyclopedia
Termites are a group of eusocial
Eusociality
Eusociality is a term used for the highest level of social organization in a hierarchical classification....

 insect
Insect
Insects are a class of living creatures within the arthropods that have a chitinous exoskeleton, a three-part body , three pairs of jointed legs, compound eyes, and two antennae...

s that, until recently, were classified at the taxonomic rank
Taxonomy
Taxonomy is the science of identifying and naming species, and arranging them into a classification. The field of taxonomy, sometimes referred to as "biological taxonomy", revolves around the description and use of taxonomic units, known as taxa...

 of order
Order (biology)
In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family...

 Isoptera (see taxonomy below), but are now accepted as the epifamily Termitoidae, of the cockroach order Blattodea. While termites are commonly known, especially in Australia
Australia
Australia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...

, as "white ants," they are only distantly related to the ants.

Like ant
Ant
Ants are social insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. More than...

s, some bee
Bee
Bees are flying insects closely related to wasps and ants, and are known for their role in pollination and for producing honey and beeswax. Bees are a monophyletic lineage within the superfamily Apoidea, presently classified by the unranked taxon name Anthophila...

s, and wasp
Wasp
The term wasp is typically defined as any insect of the order Hymenoptera and suborder Apocrita that is neither a bee nor an ant. Almost every pest insect species has at least one wasp species that preys upon it or parasitizes it, making wasps critically important in natural control of their...

s—which are all placed in the separate order Hymenoptera
Hymenoptera
Hymenoptera is one of the largest orders of insects, comprising the sawflies, wasps, bees and ants. There are over 130,000 recognized species, with many more remaining to be described. The name refers to the heavy wings of the insects, and is derived from the Ancient Greek ὑμήν : membrane and...

—termites divide labour among castes, produce overlapping generations and take care of young collectively. Termites mostly feed on dead plant material
Detritus
Detritus is a biological term used to describe dead or waste organic material.Detritus may also refer to:* Detritus , a geological term used to describe the particles of rock produced by weathering...

, generally in the form of wood, leaf litter
Plant litter
Plant litter, leaf litter or tree litter is dead plant material, such as leaves, bark, needles, and twigs, that has fallen to the ground. Litter provides habitat for small animals, fungi, and plants, and the material may be used to construct nests. As litter decomposes, nutrients are released to...

, soil, or animal dung, and about 10 percent of the estimated 4,000 species (about 2,600 taxonomically known) are economically significant as pests that can cause serious structural damage to buildings, crops or plantation forests. Termites are major detritivore
Detritivore
Detritivores, also known as detritophages or detritus feeders or detritus eaters or saprophages, are heterotrophs that obtain nutrients by consuming detritus . By doing so, they contribute to decomposition and the nutrient cycles...

s, particularly in the subtropical
Subtropics
The subtropics are the geographical and climatical zone of the Earth immediately north and south of the tropical zone, which is bounded by the Tropic of Cancer and the Tropic of Capricorn, at latitudes 23.5°N and 23.5°S...

 and tropical
Tropics
The tropics is a region of the Earth surrounding the Equator. It is limited in latitude by the Tropic of Cancer in the northern hemisphere at approximately  N and the Tropic of Capricorn in the southern hemisphere at  S; these latitudes correspond to the axial tilt of the Earth...

 regions, and their recycling of wood and other plant matter is of considerable ecological importance.

As eusocial
Eusociality
Eusociality is a term used for the highest level of social organization in a hierarchical classification....

 insects, termites live in colonies that, at maturity, number from several hundred to several million individuals. Colonies use decentralised, self-organised
Self-organization
Self-organization is the process where a structure or pattern appears in a system without a central authority or external element imposing it through planning...

 systems of activity guided by swarm intelligence
Swarm intelligence
Swarm intelligence is the collective behaviour of decentralized, self-organized systems, natural or artificial. The concept is employed in work on artificial intelligence...

 which exploit food sources and environments unavailable to any single insect acting alone. A typical colony contains nymphs (semi-mature young), workers, soldiers, and reproductive individuals of both genders, sometimes containing several egg-laying queens.

Reproductives



A female that has flown, mated, and is producing eggs is called a "queen." Similarly, a male that has flown, mated, and is in proximity to a queen is termed a "king." Research using genetic techniques to determine relatedness of colony members has shown that the original idea that colonies are only ever headed by a monogamous royal pair is wrong. Multiple pairs of reproductives within a colony are commonly encountered. In the families Rhinotermitidae and Termitidae, and possibly others, sperm competition
Sperm competition
Sperm competition is a term used to refer to the competitive process between spermatozoa of two different males to fertilize an egg of a lone female. Competition occurs whenever females engage in promiscuous mating to increase their chances in producing more viable offspring...

 does not seem to occur (male genitalia are very simple and the sperm are anucleate), suggesting that only one male (king) generally mates within the colony.

At maturity, a primary queen has a great capacity to lay eggs. In physogastric species, the queen adds an extra set of ovaries with each molt
Ecdysis
Ecdysis is the moulting of the cuticula in many invertebrates. This process of moulting is the defining feature of the clade Ecdysozoa, comprising the arthropods, nematodes, velvet worms, horsehair worms, rotifers, tardigrades and Cephalorhyncha...

, resulting in a greatly distended abdomen
Abdomen
In vertebrates such as mammals the abdomen constitutes the part of the body between the thorax and pelvis. The region enclosed by the abdomen is termed the abdominal cavity...

 and increased fecundity
Fecundity
Fecundity, derived from the word fecund, generally refers to the ability to reproduce. In demography, fecundity is the potential reproductive capacity of an individual or population. In biology, the definition is more equivalent to fertility, or the actual reproductive rate of an organism or...

, often reported to reach a production of more than 2,000 eggs a day. The distended abdomen increases the queen's body length to several times more than before mating and reduces her ability to move freely, though attendant workers provide assistance. The queen is widely believed to be a primary source of pheromone
Pheromone
A pheromone is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting outside the body of the secreting individual to impact the behavior of the receiving individual...

s useful in colony integration, and these are thought to be spread through shared feeding (trophallaxis
Trophallaxis
Trophallaxis is the transfer of food or other fluids among members of a community through mouth-to-mouth or anus-to-mouth feeding. It is most highly developed in social insects such as ants, termites, wasps and bees. The word was introduced by the entomologist William Morton Wheeler in 1918...

).

The king grows only slightly larger after initial mating and continues to mate with the queen for life. This is very different from ant colonies, in which a queen mates once with the male(s) and stores the gametes for life, as the male ants die shortly after mating.

The winged (or "alate'") caste, also referred to as the reproductive caste, are generally the only termites with well-developed eyes, although workers of some harvesting species do have well-developed compound eyes, and, in other species, soldiers with eyes occasionally appear. Termites on the path to becoming alates (going through incomplete metamorphosis) form a subcaste in certain species of termites, functioning as workers ("pseudergates") and also as potential supplementary reproductives. Supplementaries have the ability to replace a dead primary reproductive and, at least in some species, several are recruited once a primary queen is lost.

In areas with a distinct dry season
Dry season
The dry season is a term commonly used when describing the weather in the tropics. The weather in the tropics is dominated by the tropical rain belt, which oscillates from the northern to the southern tropics over the course of the year...

, the alates leave the nest in large swarms after the first good soaking rain of the rainy season. In other regions, flights may occur throughout the year, or more commonly, in the spring and autumn. Termites are relatively poor fliers and are readily blown downwind in wind speeds of less than 2 km/h, shedding their wings soon after landing at an acceptable site, where they mate and attempt to form a nest in damp timber or earth.

Workers



Worker termites undertake the labors of foraging, food storage, brood and nest maintenance, and some defense duties in certain species. Workers are the main caste in the colony for the digestion of cellulose
Cellulose
Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to over ten thousand β linked D-glucose units....

 in food and are the most likely to be found in infested wood. This is achieved in one of two ways. In all termite families except the Termitidae, there are flagellate
Flagellate
Flagellates are organisms with one or more whip-like organelles called flagella. Some cells in animals may be flagellate, for instance the spermatozoa of most phyla. Flowering plants do not produce flagellate cells, but ferns, mosses, green algae, some gymnosperms and other closely related plants...

 protist
Protist
Protists are a diverse group of eukaryotic microorganisms. Historically, protists were treated as the kingdom Protista, which includes mostly unicellular organisms that do not fit into the other kingdoms, but this group is contested in modern taxonomy...

s in the gut that assist in cellulose digestion. However, in the Termitidae, which account for approximately 60 percent of all termite species, the flagellates have been lost and this digestive role is taken up, in part, by a consortium of prokaryotic organisms. This simple story, which has been in entomology
Entomology
Entomology is the scientific study of insects, a branch of arthropodology...

 textbooks for decades, is complicated by the finding that all studied termites can produce their own cellulase
Cellulase
400px|thumb|right|alt = Colored dice with checkered background|Ribbon representation of the Streptomyces lividans beta-1,4-endoglucanase catalytic domain - an example from the family 12 glycoside hydrolases...

 enzyme
Enzyme
Enzymes are proteins that catalyze chemical reactions. In enzymatic reactions, the molecules at the beginning of the process, called substrates, are converted into different molecules, called products. Almost all chemical reactions in a biological cell need enzymes in order to occur at rates...

s, and therefore might digest wood in the absence of their symbiotic microbes although there is now evidence suggesting that these gut microbes make use of termite-produced cellulase enzymes. Our knowledge of the relationships between the microbial and termite parts of their digestion 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
Mouth
The mouth is the first portion of the alimentary canal that receives food andsaliva. The oral mucosa is the mucous membrane epithelium lining the inside of the mouth....

 or anus
Anus
The anus is an opening at the opposite end of an animal's digestive tract from the mouth. Its function is to control the expulsion of feces, unwanted semi-solid matter produced during digestion, which, depending on the type of animal, may be one or more of: matter which the animal cannot digest,...

. This process of feeding of one colony member by another is known as trophallaxis
Trophallaxis
Trophallaxis is the transfer of food or other fluids among members of a community through mouth-to-mouth or anus-to-mouth feeding. It is most highly developed in social insects such as ants, termites, wasps and bees. The word was introduced by the entomologist William Morton Wheeler in 1918...

 and is one of the keys to the success of the group. 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 do not have a true worker caste, instead relying on nymphs that perform the same work without differentiating as a separate caste.

Soldiers



The soldier caste has anatomical and behavioural specializations, providing strength and armour which are primarily useful against ant attack. The proportion of soldiers within a colony varies both within and among species. Many soldiers have jaws so enlarged that they cannot feed themselves, but instead, like juveniles, are fed by workers. The pantropical subfamily Nasutitermitinae have soldiers with the ability to exude noxious liquids through either a horn-like nozzle (nasus). Simple holes in the forehead called "fontanelle
Fontanelle
A fontanelle is an anatomical feature on an infant's skull.-Anatomy:Fontanelles are soft spots on a baby's head which, during birth, enable the bony plates of the skull to flex, allowing the child's head to pass through the birth canal. The ossification of the bones of the skull causes the...

s" and which exude defensive secretions are a feature of the family Rhinotermitidae. Many species are readily identified using the characteristics of the soldiers' heads, mandibles, or nasus. Among the drywood termites, a soldier's globular ("phragmotic") head can be used to block their narrow tunnels. Termite soldiers are usually blind, but in some families, particularly among the dampwood termites, soldiers developing from the reproductive line may have at least partly functional eyes.

The specialization of the soldier caste is principally a defence against predation by ants. The wide range of jaw types and phragmotic heads provides methods that effectively block narrow termite tunnels against ant entry. A tunnel-blocking soldier can rebuff attacks from many ants. Usually more soldiers stand by behind the initial soldier so once the first one falls another soldier will take the place. In cases where the intrusion is coming from a breach that is larger than the soldier's head, defense requires special formations where soldiers form a phalanx
Phalanx formation
The phalanx is a rectangular mass military formation, usually composed entirely of heavy infantry armed with spears, pikes, sarissas, or similar weapons...

-like formation around the breach and bite at intruders or exude toxins from the nasus or fontanelle. This formation involves self-sacrifice because once the workers have repaired the breach during fighting, no return is provided, thus leading to the death of all defenders. Another form of self-sacrifice is performed by Southeast Asian tar-baby termites (Globitermes sulphureus
Globitermes sulphureus
Globitermes sulphureus is a species of termite that is very common in Central and Southern Vietnam and also present in other areas of South East Asia, including Malaysia. They live in nests made of earth that can be up to 1.5m tall and can contain tens of thousands of individuals...

). The soldiers of this species commit suicide by autothysis
Autothysis
Autothysis is the process where an animal destroys itself via an internal rupturing or explosion of an organ which ruptures the skin. The term was proposed by Maschwitz and Maschwitz in 1974 to describe the defensive mechanism of the carpenter ant . It is caused by a contraction of muscles around a...

—rupturing a large gland just beneath the surface of their cuticle. The thick yellow fluid in the gland becomes very sticky on contact with the air, entangling ants or other insects who are trying to invade the nest.

Termites undergo incomplete metamorphosis. Freshly hatched young appear as tiny termites that grow without significant morphological changes (other than wings and soldier specializations). Some species of termite have dimorphic
Polymorphism (biology)
Polymorphism in biology occurs when two or more clearly different phenotypes exist in the same population of a species — in other words, the occurrence of more than one form or morph...

 soldiers (up to three times the size of smaller soldiers). Though their value is unknown, speculation is that they may function as an elite class that defends only the inner tunnels of the mound. Evidence for this is that, even when provoked, these large soldiers do not defend themselves but retreat deeper into the mound. On the other hand, dimorphic soldiers are common in some Australian species of Schedorhinotermes that neither build mounds nor appear to maintain complex nest structures. Some termite taxa are without soldiers; perhaps the best known of these are in the Apicotermitinae.

Diet


Termites are generally grouped according to their feeding behaviour. Thus, the commonly used general groupings are subterranean, soil-feeding, drywood, dampwood, and grass-eating. Of these, subterraneans and drywoods are primarily responsible for damage to human-made structures.

All termites eat cellulose
Cellulose
Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to over ten thousand β linked D-glucose units....

 in its various forms as plant fibre. Cellulose is a rich energy source (as demonstrated by the amount of energy released when wood is burned), but remains difficult to digest. Termites rely primarily upon symbiotic protozoa (metamonad
Metamonad
The metamonads are a large group of flagellate protozoa. Their composition is not entirely settled, but they include the retortamonads, diplomonads, and possibly the parabasalids and oxymonads as well...

s) such as Trichonympha
Trichonympha
Trichonympha is a genus of parabasalian protists that live in the intestines of many, if not most, termite species. They are symbiotes, in that they break down the cellulose in the wood and plant fibers their hosts eat....

, and other microbes in their gut to digest the cellulose for them and absorb the end products for their own use. Gut protozoa, such as Trichonympha
Trichonympha
Trichonympha is a genus of parabasalian protists that live in the intestines of many, if not most, termite species. They are symbiotes, in that they break down the cellulose in the wood and plant fibers their hosts eat....

, in turn rely on symbiotic bacteria
Bacteria
Bacteria are a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria have a wide range of shapes, ranging from spheres to rods and spirals...

 embedded on their surfaces to produce some of the necessary digestive enzyme
Digestive enzyme
'Digestive enzymes' are enzymes that break down polymeric macromolecules into their smaller building blocks, in order to facilitate their absorption by the body. Digestive enzymes are found in the digestive tract of animals where they aid in the digestion of food as well as inside the cells,...

s. This relationship is one of the finest examples of mutualism among animals. Most so-called higher termites, especially in the Family
Family
In human context, a family is a group of people affiliated by consanguinity, affinity, or co-residence. In most societies it is the principal institution for the socialization of children...

 Termitidae, can produce their own cellulase
Cellulase
400px|thumb|right|alt = Colored dice with checkered background|Ribbon representation of the Streptomyces lividans beta-1,4-endoglucanase catalytic domain - an example from the family 12 glycoside hydrolases...

 enzymes. However, they still retain a rich gut fauna and primarily rely upon the bacteria. Owing to closely related bacterial species, it is strongly presumed that the termites' gut flora
Gut flora
Gut flora consists of microorganisms that live in the digestive tracts of animals and is the largest reservoir of human flora. In this context, gut is synonymous with intestinal, and flora with microbiota and microflora....

 are descended from the gut flora of the ancestral wood-eating cockroach
Cockroach
Cockroaches are insects of the order Blattaria or Blattodea, of which about 30 species out of 4,500 total are associated with human habitations...

es, like those of the genus Cryptocercus
Cryptocercus
Cryptocercus is a genus of Dictyoptera in the family Polyphagidae, of which this genus is the only member. Species are known as wood roaches or brown-hooded cockroaches....

.

Some species of termite practice fungiculture
Fungiculture
Fungiculture is the process of producing food, medicine, and other products by the cultivation of mushrooms and other fungi.The word is also commonly used to refer to the practice of cultivating fungi by leafcutter ants, termites, ambrosia beetles, and marsh periwinkles.- Introduction :Mushrooms...

. They maintain a “garden” of specialized fungi of genus Termitomyces
Termitomyces
Termitomyces is a genus of basidiomycete fungi belonging to the family Lyophyllaceae. There are about 30 species in the genus. They are the food source for a subfamily of termites, the Macrotermitinae, who enjoy an obligate symbiosis with the genus similar to that between Atta and Attamyces...

, 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. They are also well known for eating smaller insects in a last resort environment.

Nests



Termite workers build and maintain nests which house the colony. These are elaborate structures made using a combination of soil, mud, chewed wood/cellulose, saliva, and faeces. A nest has many functions such as providing a protected living space and water conservation (through controlled condensation). There are nursery chambers deep within the nest where eggs and first instar larvae are tended. Some species maintain fungal gardens that are fed on collected plant matter, providing a nutritious mycelium
Mycelium
thumb|right|Fungal myceliaMycelium is the vegetative part of a fungus, consisting of a mass of branching, thread-like hyphae. The mass of hyphae is sometimes called shiro, especially within the fairy ring fungi. Fungal colonies composed of mycelia are found in soil and on or within many other...

 on which the colony then feeds (see "Diet," above). Nests are punctuated by a maze of tunnel-like galleries that provide air conditioning and control the CO2/O2 balance, as well as allow the termites to move through the nest.

Nests are commonly built underground, in large pieces of timber, inside fallen trees or atop living trees. Some species build nests aboveground, and they can develop into mounds. Homeowners need to be careful of tree stumps that have not been dug up. These are prime candidates for termite nests and being close to homes, termites usually end up destroying the siding and sometimes even wooden beams.

Mounds


Mounds (also known as "termitaria") occur when an aboveground nest grows beyond its initially concealing surface. They are commonly called “ant hills” in Africa and Australia, despite the technical incorrectness of that name.

In tropical savannas the mounds may be very large, with an extreme of 9 metres (30 ft) high in the case of large conical mounds constructed by some Macrotermes species in well-wooded areas in Africa. Two to three metres, however, would be typical for the largest mounds in most savanna
Savanna
A savanna, or savannah, is a grassland ecosystem characterized by the trees being sufficiently small or widely spaced so that the canopy does not close. The open canopy allows sufficient light to reach the ground to support an unbroken herbaceous layer consisting primarily of C4 grasses.Some...

s. The shape ranges from somewhat amorphous domes or cones usually covered in grass and/or woody shrubs, to sculptured hard earth mounds, or a mixture of the two. Despite the irregular mound shapes, the different species in an area can usually be identified by simply looking at the mounds.

The sculptured mounds sometimes have elaborate and distinctive forms, such as those of the compass termite (Amitermes meridionalis & A. laurensis) which build tall wedge-shaped mounds with the long axis oriented approximately north–south which gives them their alternative name of compass termites. This orientation has been experimentally shown to assist thermoregulation
Thermoregulation
Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different...

. The thin end of the nest faces towards the sun at its peak intensity hence taking up the least possible heat, this allows these termites to stay above ground where other species are forced to move into deeper below ground areas. This allows the compass termites to live in poorly drained areas where other species would be caught between a choice of baking or drowning
The column of hot air rising in the aboveground mounds helps drive air circulation currents inside the subterranean network. The structure of these mounds can be quite complex. The temperature control
Temperature control
Temperature control is a process in which change of temperature of a space is measured or otherwise detected, and the passage of heat energy into or out of the space is adjusted to achieve a desired average temperature....

 is essential for those species that cultivate fungal gardens and even for those that don't, much effort and energy is spent maintaining the brood within a narrow temperature range, often only plus or minus 1 degree C over a day.

In some parts of the African savanna, a high density of aboveground mounds dominates the landscape. For instance, in some parts of the Busanga Plain area of Zambia, small mounds of about 1 m diameter with a density of about 100 per hectare can be seen on grassland between larger tree- and bush-covered mounds about 25 m in diameter with a density around 1 per hectare, and both show up well on high-resolution satellite images taken in the wet season.




Shelter tubes



Termites are weak and relatively fragile insects that need to stay moist to survive. They can be overpowered by ants and other predators when exposed. They avoid these perils by covering their trails with tubing made of faeces, plant matter, saliva and soil. Thus the termites can remain hidden and wall out unfavourable environmental conditions. Sometimes these shelter tubes will extend for many metres, such as up the outside of a tree reaching from the soil to dead branches.

To a subterranean termite any breach of their tunnels or nest is a cause for alarm. When the Formosan subterranean termite
Formosan subterranean termite
The Formosan subterranean termite is an invasive species of termite. It has been transported worldwide from its native range in southern China to Formosa and Japan...

 (Coptotermes formosanus) and the Eastern subterranean termite
Eastern subterranean termite
Reticulitermes flavipes, the eastern subterranean termite is the most common termite found in North America. These termites are the most economically important wood destroying insects in the United States and are classified as pests. They feed on cellulose material such as the structural wood in...

 (Reticulitermes flavipes) detect a potential breach, the soldiers will usually bang their heads apparently to attract other soldiers for defence and recruit additional workers to repair any breach. This head-banging response to vibration is also useful when attempting to locate termites in house frames.

Timber damage





Owing to their wood-eating habits, many termite species can do great damage to unprotected buildings and other wooden structures. Their habit of remaining concealed often results in their presence being undetected until the timbers are severely damaged and exhibit surface changes. Once termites have entered a building, they do not limit themselves to wood; they also damage paper
Paper
Paper is a thin material mainly used for writing upon, printing upon, drawing or for packaging. It is produced by pressing together moist fibers, typically cellulose pulp derived from wood, rags or grasses, and drying them into flexible sheets....

, cloth, carpet
Carpet
A carpet is a textile floor covering consisting of an upper layer of "pile" attached to a backing. The pile is generally either made from wool or a manmade fibre such as polypropylene,nylon or polyester and usually consists of twisted tufts which are often heat-treated to maintain their...

s, and other cellulosic materials. Particles taken from soft plastics, plaster, rubber, and sealants such as silicone rubber
Silicone rubber
Silicone rubber is an elastomer composed of silicone—itself a polymer—containing silicon together with carbon, hydrogen, and oxygen. Silicone rubbers are widely used in industry, and there are multiple formulations...

 and acrylics are often employed in construction.

Humans have moved many wood-eating species between continents, but have also caused drastic population decline in others through habitat loss
Habitat destruction
Habitat destruction is the process in which natural habitat is rendered functionally unable to support the species present. In this process, the organisms that previously used the site are displaced or destroyed, reducing biodiversity. Habitat destruction by human activity mainly for the purpose of...

 and pesticide application.

Termites are commonly viewed as pests in many countries, because of the damage they can cause to structures and similar nuisances. In April 2011 wood-eating termites were blamed for reportedly consuming more than $220,000 worth of Indian rupee notes.

Precautions:
  • Avoid contact of susceptible timber with ground by using termite-resistant concrete
    Concrete
    Concrete is a composite construction material, composed of cement and other cementitious materials such as fly ash and slag cement, aggregate , water and chemical admixtures.The word concrete comes from the Latin word...

    , steel
    Steel
    Steel is an alloy that consists mostly of iron and has a carbon content between 0.2% and 2.1% by weight, depending on the grade. Carbon is the most common alloying material for iron, but various other alloying elements are used, such as manganese, chromium, vanadium, and tungsten...

    , or masonry
    Masonry
    Masonry is the building of structures from individual units laid in and bound together by mortar; the term masonry can also refer to the units themselves. The common materials of masonry construction are brick, stone, marble, granite, travertine, limestone; concrete block, glass block, stucco, and...

     foundation with appropriate barriers. Even so, termites are able to bridge these with shelter tubes, and it has been known for termites to chew through piping made of soft plastics and even some metals, such as lead
    Lead
    Lead is a main-group element in the carbon group with the symbol Pb and atomic number 82. Lead is a soft, malleable poor metal. It is also counted as one of the heavy metals. Metallic lead has a bluish-white color after being freshly cut, but it soon tarnishes to a dull grayish color when exposed...

    , to exploit moisture. In general, new buildings should be constructed with embedded physical termite barriers so that there are no easy means for termites to gain concealed entry. While barriers of poisoned soil, so-called termite pre-treatment
    Termite pre-treatment
    - What are Termite Pre-Treatments? :"Pre-treatment" is a term for termiticide that is applied to the soil. This page relates principally to the continental United States as other parts of the world often have less toxic approaches to termite management...

    , have been in general use since the 1970s, it is preferable that these be used only for existing buildings without effective physical barriers.
  • The intent of termite barriers (whether physical, poisoned soil, or some of the new poisoned plastics) is to prevent the termites from gaining unseen access to structures. In most instances, termites attempting to enter a barriered building will be forced into the less favourable approach of building shelter tubes up the outside walls; thus, they can be clearly visible both to the building occupants and a range of predators.
  • Timber treatment.
  • Termite pre-treatment
    Termite pre-treatment
    - What are Termite Pre-Treatments? :"Pre-treatment" is a term for termiticide that is applied to the soil. This page relates principally to the continental United States as other parts of the world often have less toxic approaches to termite management...

    .
  • Use of timber that is naturally resistant to termites such as Syncarpia glomulifera
    Syncarpia glomulifera
    Syncarpia glomulifera is a tree native to New South Wales and Queensland in Australia, which can grow to more than 30 metres in height...

    (Turpentine Tree), Tectona grandis (Teak), Callitris glaucophylla (White Cypress), or one of the sequoias
    Sequoioideae
    Sequoioideae is a subfamily in the Cupressaceae family, with three genera.-Genera:The three redwood subfamily genera are: Sequoia and Sequoiadendron of California and Oregon, USA; and Metasequoia in China. The redwood species contains the largest and tallest trees in the world. These trees can live...

    . Note that there is no tree species whose every individual tree yields only timbers that are immune to termite damage, so that even with well-known termite-resistant timber types, there will occasionally be pieces that are attacked.


When termites have already penetrated a building, the first action is usually to destroy the colony with insecticide
Insecticide
An insecticide is a pesticide used against insects. They include ovicides and larvicides used against the eggs and larvae of insects respectively. Insecticides are used in agriculture, medicine, industry and the household. The use of insecticides is believed to be one of the major factors behind...

s before removing the termites' means of access and fixing the problems that encouraged them in the first place. Baits (feeder stations) with small quantities of disruptive insect hormones or other very slow acting toxins have become the preferred least-toxic management tool in most western countries. This has replaced the dusting of toxins direct into termite tunnels that had been widely done since the early 1930s (originating in Australia). The main dust toxicants have been the inorganic metallic poison arsenic trioxide
Arsenic trioxide
Arsenic trioxide is the inorganic compound with the formula As2O3. This commercially important oxide of arsenic is the main precursor to other arsenic compounds, including organoarsenic compounds. Approximately 50,000 tonnes are produced annually...

, insect growth regulators (hormone
Hormone
A hormone is a chemical released by a cell or a gland in one part of the body that sends out messages that affect cells in other parts of the organism. Only a small amount of hormone is required to alter cell metabolism. In essence, it is a chemical messenger that transports a signal from one...

s) such as triflumuron and, more recently fipronil
Fipronil
Fipronil is a broad spectrum insecticide that disrupts the insect central nervous system by blocking the passage of chloride ions through the GABA receptor and glutamate-gated chloride channels, components of the central nervous system. This causes hyperexcitation of contaminated insects' nerves...

, a phenyl-pyrazole. Blowing dusts into termite workings is a highly skilled process. All these slow-acting poison
Poison
In the context of biology, poisons are substances that can cause disturbances to organisms, usually by chemical reaction or other activity on the molecular scale, when a sufficient quantity is absorbed by an organism....

s can be distributed by the workers for hours or weeks before any symptoms occur and are capable of destroying the entire colony. More modern variations include chlorfluazuron, diflubenzuron
Diflubenzuron
Diflubenzuron is an insecticide of the benzamide class. It is used in forest management and on field crops to selectively control insect pests, particularly forest tent caterpillar moths, boll weevils, gypsy moths, and other types of moths. The mechanism of action of diflubenzuron involves...

, hexaflumuron, and novaflumuron as bait toxicants and fipronil
Fipronil
Fipronil is a broad spectrum insecticide that disrupts the insect central nervous system by blocking the passage of chloride ions through the GABA receptor and glutamate-gated chloride channels, components of the central nervous system. This causes hyperexcitation of contaminated insects' nerves...

, imidacloprid
Imidacloprid
Imidacloprid is a nicotine-based, systemic insecticide, which acts as a neurotoxin and belongs to a class of chemicals called the neonicotinoids. Although it is now off patent, the primary manufacturer of this chemical is Bayer CropScience,...

 and chlorantraniprole as soil poisons. Soil poisons are the least-preferred method of control as this requires large doses of toxin and results in uncontrollable release to the environment.

The termite’s effects are damaging, costing the southwestern United States approximately $1.5 billion each year in wood structure damage. In order to better control the population of termites, researchers at the Agricultural Research Service
Agricultural Research Service
The Agricultural Research Service is the principal in-house research agency of the United States Department of Agriculture . ARS is one of four agencies in USDA's Research, Education and Economics mission area...

 have found a way to track the movement of the destructive pests. In 1990, researchers found a way to safely and reliably track termites using immunoglobulin G
Immunoglobulin G
Immunoglobulin G are antibody molecules. Each IgG is composed of four peptide chains — two heavy chains γ and two light chains. Each IgG has two antigen binding sites. Other immunoglobulins may be described in terms of polymers with the IgG structure considered the monomer.IgG constitutes 75%...

 (IgG) marker proteins from rabbits or chickens. In field tests, termite bait was laced with the rabbit IgG and the termites were randomly exposed to feeding on this bait. Termites were later collected from the field and tested for the rabbit-IgG markers using a rabbit-IgG-specific assay. However, this method of testing for the tracking proteins is expensive. Recently, researchers have developed a new way of tracking the termites using egg white, cow milk, or soy milk proteins, which can be sprayed on the termites in the field. This new method is less expensive because the proteins can be traced using a protein-specific ELISA test. The ELISA test is more affordable, because it is designed for mass production. Researchers hope to use this method of tracking termites to find a more cost-effective way to control the damaging pests. http://www.ars.usda.gov/is/pr/2010/100217.htm
]

Termites in the human diet


In many cultures, termites (particularly the winged ones known as alates) are used for food. The alates are nutritious, having a good store of fat and protein, and are palatable in most species with a nutty flavour when cooked. They are easily gathered at the beginning of the rainy season in West
West Africa
West Africa or Western Africa is the westernmost region of the African continent. Geopolitically, the UN definition of Western Africa includes the following 16 countries and an area of approximately 5 million square km:-Flags of West Africa:...

, Central
Central Africa
Central Africa is a core region of the African continent which includes Burundi, the Central African Republic, Chad, the Democratic Republic of the Congo, and Rwanda....

 and Southern Africa
Southern Africa
Southern Africa is the southernmost region of the African continent, variably defined by geography or geopolitics. Within the region are numerous territories, including the Republic of South Africa ; nowadays, the simpler term South Africa is generally reserved for the country in English.-UN...

 when they swarm, as they are attracted to lights and can be gathered up when they land on nets put up around a lamp. The wings are shed and can be removed by a technique similar to winnowing. They are best gently roasted on a hot plate or lightly fried until slightly crisp; oil
Cooking oil
Cooking oil is purified fat of plant origin, which is usually liquid at room temperature ....

 is not usually needed since their bodies are naturally high in oil. Traditionally they make a welcome treat at the beginning of the rainy season when livestock is lean, new crops have not yet produced food, and stored produce from the previous growing season is running low.

They are also eaten in Indonesia
Indonesia
Indonesia , officially the Republic of Indonesia , is a country in Southeast Asia and Oceania. Indonesia is an archipelago comprising approximately 13,000 islands. It has 33 provinces with over 238 million people, and is the world's fourth most populous country. Indonesia is a republic, with an...

, including Central Java
Central Java
Central Java is a province of Indonesia. The administrative capital is Semarang. It is one of six provinces on the island of Java.This province is the province of high Human Development in Indonesia and its Points Development Index countries is equivalent to Lebanon. The province of Central Java...

, where they are roasted or fried.

Agriculture


Termites can be major agricultural pests, particularly in Africa and Asia, where crop losses can be severe. Counterbalancing this is the greatly improved water infiltration where termite tunnels in the soil allow rainwater to soak in deeply and help reduce runoff and consequent soil erosion.

Termites as a source of energy


The U.S. Department of Energy is researching ways to replace fossil fuels with renewable sources of cleaner energy, and termites are considered a possible way to reach this goal through metagenomics
Metagenomics
Metagenomics is the study of metagenomes, genetic material recovered directly from environmental samples. The broad field may also be referred to as environmental genomics, ecogenomics or community genomics. Traditional microbiology and microbial genome sequencing rely upon cultivated clonal cultures...

.

Termites may produce up to two litres of hydrogen
Hydrogen
Hydrogen is the chemical element with atomic number 1. It is represented by the symbol H. With an average atomic weight of , hydrogen is the lightest and most abundant chemical element, constituting roughly 75% of the Universe's chemical elemental mass. Stars in the main sequence are mainly...

 from digesting a single sheet of paper, making them one of the planet’s most efficient bioreactor
Bioreactor
A bioreactor may refer to any manufactured or engineered device or system that supports a biologically active environment. In one case, a bioreactor is a vessel in which a chemical process is carried out which involves organisms or biochemically active substances derived from such organisms. This...

s. Termites achieve this high degree of efficiency by exploiting the metabolic capabilities of about 200 different species of microbes that inhabit their hindguts. The microbial community in the termite gut efficiently manufactures large quantities of hydrogen; the complex lignocellulose polymer
Polymer
A polymer is a large molecule composed of repeating structural units. These subunits are typically connected by covalent chemical bonds...

s within wood are broken down into simple sugars by fermenting bacteria in the termite’s gut, using enzymes that produce hydrogen as a byproduct. A second wave of bacteria uses the simple sugars and hydrogen to make the acetate the termite requires for energy. By sequencing the termite's microbial community, the DOE hopes to get a better understanding of these biochemical pathways. If it can be determined which enzymes are used to create hydrogen, and which genes produce them, this process could potentially be scaled up with bioreactors to generate hydrogen from woody biomass, such as poplar
Poplar
Populus is a genus of 25–35 species of deciduous flowering plants in the family Salicaceae, native to most of the Northern Hemisphere. English names variously applied to different species include poplar , aspen, and cottonwood....

, in commercial quantities.

Skeptics regard this as unlikely to become a carbon-neutral commercial process owing to the energy inputs required to maintain the system. For decades, researchers have sought to house termites on a commercial scale (like worm farms) to break down woody debris and paper, but funding has been scarce and the problems of developing a continuous process that does not disrupt the termites' homeostasis
Homeostasis
Homeostasis is the property of a system that regulates its internal environment and tends to maintain a stable, constant condition of properties like temperature or pH...

 have not been overcome.

Ground water divining in Ancient India


Varaha Mihira (505 AD- 587 AD), the famous astronomer, mathematician, and astrologer of India
History of India
The history of India begins with evidence of human activity of Homo sapiens as long as 75,000 years ago, or with earlier hominids including Homo erectus from about 500,000 years ago. The Indus Valley Civilization, which spread and flourished in the northwestern part of the Indian subcontinent from...

, in his treatise "Brihat Samhita," also spelled "Vrahat Sanhita," refers to Dakargala (Sanskrit word meaning “science of underground water exploration”), wherein the role of termite knolls, as an indicator of underground water has been elaborately explained.

In Verse.S.54.9 of the Samhita, it is stated that sweet ground water would be found near a termite mound located east of a Jambu tree (botanical names - Eugenia Jambus,Engenia Jambolana), at a specific distance and a specific depth of 15 ft to the south of the tree.

The above verse has been justified with an explanation:


Without exception the water requirements of the insects are generally very high, and they need to protect themselves against fatal desiccation by living and working within the climatically sealed environment of their nest or within earth covered galleries. According to present level of research, the atmosphere within the nest has to be maintained practically saturation moisture level ( 99-100 % humidity). It is a matter of common observation that whenever a termite nest or runway, is damaged, the insects immediately rush to the breach and repairs it with wet soil brought up from within the nest. From an overall consideration of the evidence it seems to be safe to conclude that, while normally the insects use every readily available source of water close to the ground surface, under condition of severe climatic stress, they can and they probably do descend to the water table, no matter how deep it may be. Hence, a well-developed, active, permanent colony of mound-building termites can be taken as an indication of underground springs in proximity.


Two examples mentioned in the referred publication are, a) termiteries seen in the Katanga province (Congo Kinshasa) right up to the hill slopes where springs emerge, b) in the dry jungle uplands of coastal zone of Karnataka
Karnataka
Karnataka , the land of the Kannadigas, is a state in South West India. It was created on 1 November 1956, with the passing of the States Reorganisation Act and this day is annually celebrated as Karnataka Rajyotsava...

 state (old Mysore state) and c) in the Deccan Plateau
Deccan Plateau
The Deccan Plateau is a large plateau in India, making up the majority of the southern part of the country. It rises a hundred meters high in the north, rising further to more than a kilometers high in the south, forming a raised triangle nested within the familiar downward-pointing triangle of...

 area.

It is also asserted in the verse Vr.S.54.85 that among a group of termite mounds, a water vein is sure to be found below the taller of the mounds. Verse 52 mentions that in a desert region, if a group of five termite mounds are found, and if the middle one is in white colour, then water will be found within a depth of Fifty five Purushas (in Sanskrit one Purusha is equivalent to 7.5 ft) or 412.5 ft.

As a common observation of a combination of different symptoms, termite mounds are said to be found close to trees, and ancient Hindus exploited this knowledge in the exploration of underground springs.

In captivity


Few zoos hold termites, due to the difficulty in keeping them captive and the reluctance of authorities to permit potential pests. One of them is Zoo Basel
Zoo Basel
Zoo Basel is a non-profit zoo located within the city of Basel, Switzerland. Its official name is Zoologischer Garten Basel — or in English: Basel Zoological Garden. Basel residents, however, call their zoo affectionately Zolli...

 in Switzerland
Switzerland
Switzerland name of one of the Swiss cantons. ; ; ; or ), in its full name the Swiss Confederation , is a federal republic consisting of 26 cantons, with Bern as the seat of the federal authorities. The country is situated in Western Europe,Or Central Europe depending on the definition....

. At Zoo Basel, two African termite (Macrotermes bellicosus
Macrotermitinae
Macrotermitinae constitute a subfamily of the harvester termites and allies . The termites are traditionally classified as order Isoptera, but have been found to be a subgroup of the Blattaria and consequently should be treated as part to this group; what taxonomy entomologists eventually will...

) populations exist and thrive - resulting in very rare (in captivity) mass migrations of young flying termites. This happened last in September 2008, when thousands of male termites left their mound each night, died, and covered the floors and water pits of the house their exhibit is in.

Ecology



Ecologically, termites are important in nutrient
Nutrient
A nutrient is a chemical that an organism needs to live and grow or a substance used in an organism's metabolism which must be taken in from its environment. They are used to build and repair tissues, regulate body processes and are converted to and used as energy...

 recycling, habitat
Habitat (ecology)
A habitat is an ecological or environmental area that is inhabited by a particular species of animal, plant or other type of organism...

 creation, soil
Soil
Soil is a natural body consisting of layers of mineral constituents of variable thicknesses, which differ from the parent materials in their morphological, physical, chemical, and mineralogical characteristics...

 formation and quality and, particularly the winged reproductives, as food for countless predators. The role of termites in hollowing timbers and thus providing shelter and increased wood surface areas for other creatures is critical for the survival of a large number of timber-inhabiting species. Larger termite mounds play a role in providing a habitat for plants and animals, especially on plains in Africa that are seasonally inundated
Floodplain
A floodplain, or flood plain, is a flat or nearly flat land adjacent a stream or river that stretches from the banks of its channel to the base of the enclosing valley walls and experiences flooding during periods of high discharge...

 by a rainy season, providing a retreat above the water for smaller animals and birds, and a growing medium for woody shrubs with root systems that cannot withstand inundation for several weeks. In addition, scorpion
Scorpion
Scorpions are predatory arthropod animals of the order Scorpiones within the class Arachnida. They have eight legs and are easily recognized by the pair of grasping claws and the narrow, segmented tail, often carried in a characteristic forward curve over the back, ending with a venomous stinger...

s, lizard
Lizard
Lizards are a widespread group of squamate reptiles, with nearly 3800 species, ranging across all continents except Antarctica as well as most oceanic island chains...

s, snakes, small mammals, and birds live in abandoned or weathered mounds, and aardvark
Aardvark
The aardvark is a medium-sized, burrowing, nocturnal mammal native to Africa...

s dig substantial caves and burrows in them, which then become homes for larger animals such as hyena
Hyena
Hyenas or Hyaenas are the animals of the family Hyaenidae of suborder feliforms of the Carnivora. It is the fourth smallest biological family in the Carnivora , and one of the smallest in the mammalia...

s and mongoose
Mongoose
Mongoose are a family of 33 living species of small carnivorans from southern Eurasia and mainland Africa. Four additional species from Madagascar in the subfamily Galidiinae, which were previously classified in this family, are also referred to as "mongooses" or "mongoose-like"...

s.

As detrivores, termites clear away leaf and woody litter and so reduce the severity of the annual bush fires in African savannas, which are not as destructive as those in Australia and the U.S.A. Their role in bioturbation
Bioturbation
In oceanography, limnology, pedology, geology , and archaeology, bioturbation is the displacement and mixing of sediment particles and solutes by fauna or flora . The mediators of bioturbation are typically annelid worms , bivalves In oceanography, limnology, pedology, geology (especially...

 on the Khorat Plateau
Khorat Plateau
The Khorat Plateau also Korat Plateau, is a plateau in the northeastern region of Thailand, named for the short form of Nakhon Ratchasima, an historical stronghold controlling access to and from the plateau.-Geography:...

 is under investigation.

Globally, termites are found roughly between 50 degrees north & south, with the greatest biomass
Biomass
Biomass, as a renewable energy source, is biological material from living, or recently living organisms. As an energy source, biomass can either be used directly, or converted into other energy products such as biofuel....

 in the tropics and the greatest diversity in tropical forests and Mediterranean shrublands. Termites are also considered to be a major source of atmospheric methane
Methane
Methane is a chemical compound with the chemical formula . It is the simplest alkane, the principal component of natural gas, and probably the most abundant organic compound on earth. The relative abundance of methane makes it an attractive fuel...

, one of the prime greenhouse gas
Greenhouse gas
A greenhouse gas is a gas in an atmosphere that absorbs and emits radiation within the thermal infrared range. This process is the fundamental cause of the greenhouse effect. The primary greenhouse gases in the Earth's atmosphere are water vapor, carbon dioxide, methane, nitrous oxide, and ozone...

es. Termites have been common since at least the Cretaceous
Cretaceous
The Cretaceous , derived from the Latin "creta" , usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago. In the geologic timescale, the Cretaceous follows the Jurassic period and is followed by the Paleogene period of the...

 period. Termites also eat bone and other parts of carcasses, and their traces have been found on dinosaur bones from the middle Jurassic in China.

Plant defences against termites


Many plants have developed effective defences against termites, and in most ecosystems, there is an observable balance between the growth of plants and the feeding of termites. Defence is typically achieved by secreting anti-feedant chemicals (such as oil
Oil
An oil is any substance that is liquid at ambient temperatures and does not mix with water but may mix with other oils and organic solvents. This general definition includes vegetable oils, volatile essential oils, petrochemical oils, and synthetic oils....

s, resin
Resin
Resin in the most specific use of the term is a hydrocarbon secretion of many plants, particularly coniferous trees. Resins are valued for their chemical properties and associated uses, such as the production of varnishes, adhesives, and food glazing agents; as an important source of raw materials...

s, and lignin
Lignin
Lignin or lignen is a complex chemical compound most commonly derived from wood, and an integral part of the secondary cell walls of plants and some algae. The term was introduced in 1819 by de Candolle and is derived from the Latin word lignum, meaning wood...

s) into the woody cell walls. This reduces the ability of termites to efficiently digest the cellulose
Cellulose
Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to over ten thousand β linked D-glucose units....

. Many of the strongly termite-resistant tree species have heartwood timber that is extremely dense (such as Eucalyptus camaldulensis) due to accretion of these resins. Over the years there has been considerable research into these natural defensive chemicals with scientists seeking to add them to timbers from susceptible trees. A commercial product, “Blockaid,” has been developed in Australia and uses a range of plant extracts to create a paint-on nontoxic termite barrier for buildings. In 2005 a group of Australian scientists “discovered” (announced) a treatment based on an extract of a species of Eremophila
Eremophila (plant)
Eremophila is a genus of plants of the family Scrophulariaceae, with species known by the common names of Emu Bush, Poverty Bush or Fuchsia Bush. Currently, there are 215 recognised species, all of which are endemic to Australia...

 that repels termites. Tests have shown that termites are strongly repelled by the toxic material to the extent that they will starve rather than consume cross treated samples. When kept close to the extract, they become disoriented and eventually die. Scientists hoped to use this toxic compound commercially as a soil barrier but it failed field testing.

Taxonomy, evolution and systematics



Recent DNA evidence has supported the hypothesis, originally based on morphology, that termites are most closely related to the wood-eating cockroach
Cockroach
Cockroaches are insects of the order Blattaria or Blattodea, of which about 30 species out of 4,500 total are associated with human habitations...

es (genus Cryptocercus
Cryptocercus
Cryptocercus is a genus of Dictyoptera in the family Polyphagidae, of which this genus is the only member. Species are known as wood roaches or brown-hooded cockroaches....

), to which the singular and very primitive Mastotermes darwiniensis shows some telltale similarities. Most recently, this has led some authors to propose that termites be reclassified as a single family, Termitidae, within the order Blattodea, which contains cockroaches. However, most researchers advocate the less drastic measure of retaining the termites as Termitoidae, an epifamily of the cockroach Order, which preserves the classification of termites at family level and below.

Evolutionary history


The oldest unambiguous termite fossils date to the early Cretaceous
Cretaceous
The Cretaceous , derived from the Latin "creta" , usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago. In the geologic timescale, the Cretaceous follows the Jurassic period and is followed by the Paleogene period of the...

, although structures from the late Triassic have been interpreted as fossilized termite nests. Given the diversity of Cretaceous termites, it is likely that they had their origin at least sometime in the Jurassic
Jurassic
The Jurassic is a geologic period and system that extends from about Mya to  Mya, that is, from the end of the Triassic to the beginning of the Cretaceous. The Jurassic constitutes the middle period of the Mesozoic era, also known as the age of reptiles. The start of the period is marked by...

. Weesner believes that Mastotermitidae termites may go back to the Permian and fossil wings have been discovered in the Permian of Kansas which have a close resemblance to wings of Mastotermes of the Mastotermitidae, which is the most primitive living termite. It is thought to be the descendant of Cryptocercus
Cryptocercus
Cryptocercus is a genus of Dictyoptera in the family Polyphagidae, of which this genus is the only member. Species are known as wood roaches or brown-hooded cockroaches....

genus, the wood roach. This fossil is called Pycnoblattina. It folded its wings in a convex pattern between segments 1a and 2a. Mastotermes
Mastotermes
Mastotermes darwiniensis, common names Giant Northern Termite and Darwin Termite, is a termite species found only in northern Australia. It is a very peculiar animal, the most primitive termite alive. As such, it shows uncanny similarities to certain cockroaches, the termites' closest relatives...

 is the only living insect that does the same,

It has long been accepted that termites are closely related to cockroach
Cockroach
Cockroaches are insects of the order Blattaria or Blattodea, of which about 30 species out of 4,500 total are associated with human habitations...

es and mantids, and they are classified in the same superorder (Dictyoptera
Dictyoptera
Dictyoptera includes three groups of polyneopterous insects - cockroaches , termites and mantids...

), but new research has shed light on the details of termite evolution. There is now strong evidence suggesting that termites are really highly modified, social, wood-eating cockroaches. A study conducted by scientists has found that endosymbiotic bacteria from termites and a genus of cockroaches, Cryptocercus
Cryptocercus
Cryptocercus is a genus of Dictyoptera in the family Polyphagidae, of which this genus is the only member. Species are known as wood roaches or brown-hooded cockroaches....

, share the strongest phylogenetical similarities out of all other cockroaches. Both termites and Cryptocercus also share similar morphological and social features—most cockroaches do not show social characteristics, but Cryptocercus takes care of its young and exhibits other social behaviour
Social behavior
In physics, physiology and sociology, social behavior is behavior directed towards society, or taking place between, members of the same species. Behavior such as predation which involves members of different species is not social...

. As mentioned above, the primitive Giant Northern Termite (Mastotermes darwiniensis) exhibits numerous cockroach-like characteristics, such as laying its eggs in rafts and having anal lobes on the wings that are not shared with other termites.

Systematics


As of 1996, about 2,800 termite species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...

 are recognized, classified in seven families http://www.utoronto.ca/forest/termite/speclist.htm. These are arranged here in a phylogenetic sequence, from the most basal to the most advanced:
  • Mastotermitidae (1 species, Mastotermes darwiniensis)

  • Hodotermitidae (3 genera, 19 species)
    • Hodotermitinae

  • Kalotermitidae
    Kalotermitidae
    Kalotermitidae is a family of termites. Kalotermitidae include 22 genera and 419 species.-Genera:* Calcaritermes* Cryptotermes* Glyptotermes* Incisitermes* Kalotermes* Marginitermes* Neotermes* Paraneotermes* Procryptotermes* Pterotermes...

     (22 genera, 419 species)

  • Termopsidae
    Termopsidae
    Dampwood termites constitute a small and rather primitive family of termites . They contain a mere 4–5 extant genera with 13–20 living species, but can be divided into several subfamilies. They may be a nuisance but compared to e.g. the drywood termites usually do not cause extensive damage to...

     (5 genera, 20 species)
    • Termopsinae
    • Porotermitinae
    • Stolotermitinae

  • Rhinotermitidae
    Rhinotermitidae
    Rhinotermitidae is a family of termites . They feed on wood and can cause extensive damage to buildings or other wooden structures. About 345 species are recognized, among these are severe pests like Coptotermes formosanus, Coptotermes gestroi and Reticulitermes flavipes....

     (14 genera, 343 species)
    • Coptotermitinae Holmgren
    • Heterotermitinae Froggatt
    • Prorhinoterminae Quennedey & Deligne, 1975
    • Psammotermitinae Holmgren
    • Rhinotermitinae Froggatt
    • Stylotermitinae Holmgren, K & N, 1917
    • Termitogetoninae Holmgren

  • Serritermitidae (1 species, Serritermes serrifer)

  • Termitidae
    Termitidae
    Termitidae is a family of termite, containing the following subfamilies:*Apicotermitinae Grassé & Noirot, 1955*Foraminitermitinae Holmgren, 1912*Macrotermitinae Kemner, 1934*Nasutitermitinae Hare, 1937*Sphaerotermitinae Engel & Krishna, 2004...

     (236 genera, 1958 species)
    • Apicotermitinae (42 genera, 208 species)
    • Foraminitermitinae (2 genera, 9 species)
    • Macrotermitinae
      Macrotermitinae
      Macrotermitinae constitute a subfamily of the harvester termites and allies . The termites are traditionally classified as order Isoptera, but have been found to be a subgroup of the Blattaria and consequently should be treated as part to this group; what taxonomy entomologists eventually will...

       (13 genera, 362 species)
    • Nasutitermitinae (80 genera, 576 species)
    • Sphaerotermitinae (1 species)
    • Syntermitinae (13 genera, 99 species)
    • Termitinae (90 genera, 760 species)


The most current classification of termites is summarized by Engel & Krishna (2004).

See also


  • Coatonachthodes ovambolandicus
    Coatonachthodes ovambolandicus
    Coatonachthodes ovambolandicus is a termite-mimicking beetle which lives as a parasite in their nests. It looks roughly like a termite from above, though its legs look like "twisted balloons" rather than real insect legs. This less than accurate emulation can be explained by looking at the beetle...

  • Decompiculture
    Decompiculture
    The term decompiculture is a neologism coined by forestry professor Timothy Myles of the Urban Entomology Program at the University of Toronto and refers to how decomposing organisms, like termites, could be grown or cultured for a variety of uses....

  • International Union for the Study of Social Insects
    International Union for the Study of Social Insects
    International Union for the Study of Social Insects is an association of entomologists from different countries engaged in research of social insects...

  • Nasutitermes corniger
    Nasutitermes corniger
    Nasutitermes corniger is a species of arboreal termite that is endemic to the neotropics. It is very closely related to Nasutitermes ephratae.The species has been studied relatively intensively, particularly on Barro Colorado Island, Panama...

  • Stigmergy
    Stigmergy
    Stigmergy is a mechanism of indirect coordination between agents or actions. The principle is that the trace left in the environment by an action stimulates the performance of a next action, by the same or a different agent...

  • Xylophagy
    Xylophagy
    Xylophagy is a term used in ecology to describe the habits of an herbivorous animal whose diet consists primarily of wood. The word derives from Greek ξυλοφάγος "eating wood", from ξύλον "wood" and φαγεῖν "to eat", an ancient Greek name for a kind of a worm-eating bird...



Further reading



External links



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