Termite 54808 225967901 2008-07-16T06:48:48Z Dyanega 1357080 rv vandalism {{Taxobox | fossil_range = [[Late Triassic]] - Recent | image = Coptotermes formosanus shiraki USGov k8204-7.jpg | image_width = 200px | image_caption = [[Formosan subterranean termite]] soldiers (red colored heads) and workers (pale colored heads). | regnum = [[Animal]]ia | phylum = [[Arthropod]]a | classis = [[Insect]]a | subclassis = [[Pterygota]] | infraclassis = [[Neoptera]] | superordo = [[Dictyoptera]] | ordo = [[Isoptera]] | subdivision_ranks = Families | subdivision = [[Mastotermitidae]]<br> [[Kalotermitidae]]<br> [[Termopsidae]] <br> [[Hodotermitidae]]<br> [[Rhinotermitidae]]<br> [[Serritermitidae]]<br> [[Termitidae]]<br> }} '''Termites''', sometimes incorrectly called '''"white [[ant]]s"''', are a group of [[social]] [[insect]]s usually classified at the [[Taxonomy|taxonomic rank]] of [[Order (biology)|order]] '''Isoptera''' (but see also [[#taxonomy|taxonomy]] below). As truly social animals, they are termed ''[[eusocial]]'' along with the ants and some bees and wasps. Termites mostly feed on [[detritus|dead plant material]], generally in the form of wood, leaf litter, soil, or animal dung, and about 10% 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 [[detrivore]]s, particularly in the [[Subtropics|subtropical]] and [[Tropics|tropical]] regions, and their recycling of wood and other plant matter is of considerable [[ecological]] importance. As [[eusocial insect]]s, termites live in colonies that, at maturity, number from several hundred to several million individuals. They are a prime example of [[decentralised]], [[self-organised]] systems using [[swarm intelligence]] and use this cooperation to exploit food sources and environments that could not be available 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. ==Social organization== ===Reproductives=== A female that has flown, mated, and is producing eggs, is called a "queen". Similarly, a male that has flown, mated, and remains in proximity to a queen, is termed a "King". These anthropocentric terms have caused great misunderstanding of colony dynamics. Research using genetic techniques to determine relatedness of colony members is showing that the idea that colonies are only ever headed by a [[monogamous]] royal pair is wrong. Multiple pairs of reproductives within a colony are not uncommon. In the families Rhinotermitidae and Termitidae, and possibly others, [[sperm competition]] 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 [[ecdysis|moult]], resulting in a greatly distended [[abdomen]] and increased [[fecundity]], often reported to reach a production of more than two thousand 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]]s useful in colony integration, and these are thought to be spread through shared feeding ([[trophallaxis]]). 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, and 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 sub-caste 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, 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 windspeeds of less than 2 [[kilometres per hour|kph]], 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=== [[Image:Workertermite1.jpg|350px|right|thumb|Worker termite]] Worker termites undertake the labours of foraging, food storage, brood, nest maintenance, and some of the defence effort in certain species. Workers are the main caste in the colony for the digestion of [[cellulose]] 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]] [[protist]]s in the gut that assist in cellulose digestion. However, in the Termitidae, which account for approximately 60% 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]] textbooks for decades, is complicated by the finding that all studied termites can produce their own [[cellulase]] [[enzyme]]s, and therefore can digest wood in the absence of their symbiotic microbes. 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 or [[anus]]. This process of feeding of one colony member by another is known as [[trophallaxis]] 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 moulting into a separate caste. Termite workers are generally blind due to undeveloped eyes. Despite this limitation, they are able to create elaborate nests and tunnel systems using a combination of soil, chewed wood/cellulose, saliva, and faeces. Some species have been known to create such durable walls that industrial machinery has been damaged in an attempt to break their tall mounds{{Fact|date=March 2008}}. Some African and Australian species have mounds more than 4 metres high. The nest is created and maintained by workers with many distinct features such as housing the brood, water collection through condensation, reproductive chambers, and tunnel networks that effectively provide air conditioning and control the CO<sub>2</sub>/O<sub>2</sub> balance. A few species even practice agriculture, with elaborate fungal gardens which are fed on collected plant matter, providing a nutritious [[mycelium]] on which the colony then feeds (''see "Diet", below''). ===Soldiers=== [[Image:Nasutes.jpg|thumb|Termites with some nasute 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 pan-tropical sub-family Nasutitermitinae (''The South American species of which are under review and are likely to deserve a separate taxon'')have soldiers with the ability to exude noxious liquids through either a horn-like nozzle (nasus) or simple hole in the head ([[fontanelle]]). Fontanelles which exude defensive secretions are also a feature of the family Rhinotermitidae. Many species are readily identified using the characteristics of the soldiers' heads, [[mandible]]s, 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, soldiers developing from the reproductive line may have at least partly functional eyes. [[Image:NasuteImms.png|thumb|A nasute]] It's generally accepted that 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 which 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, defence requires special formations where soldiers form a [[Phalanx formation|phalanx]]-like formation around the breach and blindly bite at intruders or shoot toxic glue from the [[nasus]]. This formation involves self-sacrifice because once the workers have repaired the breach during fighting, no return is provided, thus causing the death of all the defenders. Termites undergo incomplete [[metamorphosis (biology)|metamorphosis]], with their freshly hatched young taking the form of tiny termites that grow without significant morphological changes (other than wings and soldier specializations). Some species of termite have [[dimorphic]] 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 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]] 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]]s) such as ''[[Trichonympha]]'', and other [[microbe]]s in their gut to digest the cellulose for them and absorb the end products for their own use. Gut protozoa, such as ''[[Trichonympha]]'', in turn rely on symbiotic [[bacterium|bacteria]] embedded on their surfaces to produce some of the necessary digestive enzymes. This relationship is one of the finest examples of mutualism among animals. Most so called "higher termites", especially in the [[Family]] Termitidae, can produce their own [[cellulase]] enzymes. However, they still retain a rich gut fauna and primarily rely upon the bacteria. Due to closely related bacterial species, it is strongly presumed that the termites' gut flora are descended from the gut flora of the ancestral wood-eating [[cockroach]]es, like those of the genus ''[[Cryptocercus]]''. Some species of termite practice [[fungiculture]]. They maintain a 'garden' of specialized 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.<ref>[http://www.pnas.org/cgi/content/full/99/23/14887 The evolution of fungus-growing termites and their mutualistic fungal symbionts] by Duur K. Aanen, Paul Eggleton, Corinne Rouland-Lefèvre, Tobias Guldberg-Frøslev, Søren Rosendahl & Jacobus J. Boomsma </ref><ref>[http://www.pnas.org/cgi/content/full/99/24/15247 Fungus-farming insects: Multiple origins and diverse evolutionary histories] by Ulrich G. Mueller & Nicole Gerardo</ref> They are also well known for eating smaller insects in a last resort environment. === Mounds <!--"termite mounds" in Ant-hill article links here -->=== [[Image:Termite mound Tanzania.jpg|thumb|right| Termite mound in Tanzania]]Termites build nests to house their colonies. Nests are commonly located in larger timber or in the soil in locations such as growing trees, inside fallen trees, underground, and in above-ground mounds which they construct, commonly called "anthills" in Africa and Australia, despite the technical incorrectness of that name. Mounds occur when the nest grows beyond its initially concealing surface. In [[tropical savanna]]s 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,<ref>[http://library.eb.com.au/all/eb/article-39592?query=Amitermes%20meridionalis&ct=null "Termite."] ''Encyclopædia Britannica'' Online Library Edition. Retrieved 19 November 2007.]]</ref>. Two to three metres, however, would be typical for the largest mounds in most [[savanna]]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. This orientation has been experimentally shown to help in [[thermoregulation]]. The column of hot air rising in the above ground mounds helps drive air circulation currents inside the subterranean network. The structure of these mounds can be quite complex. The temperature control 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 one degree C over a day. In some parts of the African savanna, a high density of above-ground 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.<ref>[[Google Earth]], at lat -14.6565° long 25.8337°. The smaller termite mounds are the light patches; the larger ones are clumps of bushes with lighter patches of bare earth. Retrieved 19 November 2007.]]</ref>. <gallery> Image:Termite_Cathedral_DSC03570.jpg|Cathedral Mounds Image:Termite Magnetic DSC03613.jpg|Magnetic Mounds (nearly North-South Axis) Image:RayNorris_termite_cathedral_mounds.jpg|Termite cathedral mounds in the [[Northern Territory]] of [[Australia]] Image:Termite cathedral mounds in a bushfire blackened tropical savanna.jpg|Two cathedral mounds in a [[tropical savanna]] blackened by [[Kakadu National Park]]'s annual winter [[bushfire]]s. <!--Image:Termite_mound_with_comparison.jpg|Humans and trees for size comparison--> </gallery> == Human interaction == [[Image:Termite damaged wood.jpg|thumb|200px|right|The result of an infestation is severe wood damage.]] Because of their wood-eating habits, termites sometimes do great damage to 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]], [[cloth]], [[carpet]]s, and other cellulosic materials. Often, other soft materials are damaged and may be used for construction. Particles taken from soft plastics, plaster, rubber, and sealants such as silicon rubber and acrylics are often employed in construction. Termites usually avoid exposure to unfavourable environmental conditions. They tend to remain hidden in tunnels in earth and wood. Where they need to cross an impervious or unfavourable substrate, they cover their tracks with tubing made of faeces, plant matter, and soil. Sometimes these shelter tubes will extend for many metres, such as up the outside of a tree reaching from the soil to dead branches. Termite barrier systems used for protecting buildings aim to prevent concealed termite access, thus forcing the termites out into the open where they must form clearly visible shelter tubes to gain entry. 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. In many cultures, termites are used for food (particularly the alates), and termite nests are used widely in construction (the dirt is often dust-free) and as a soil amendment. Humans have moved many wood-eating species between continents, but have also caused drastic population decline in others through habitat loss and pesticide application. === Avoiding termite troubles === [[Image:Termite damage.JPG|thumb|right|200px|Termite damage on external structure]] [[Image:Termite Nest In Wall Cavity.jpg|thumb|right|200px|Coptotermes acinaciformis Bivouac]] [[Image:Termite Bivouac After Treatment.jpg|thumb|right|200px|Coptotermes acinaciformis Bivouac (after treatment)]] Precautions: * Avoid contact of susceptible timber with ground by using termite-resistant [[concrete]], [[steel]], or [[masonry]] 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, 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]], 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, and thus, they can be clearly visible both to the building occupants and a range of predators. Regular inspection by a competent (trained and experienced) inspector is the best defence. * [[Timber treatment]]. * Use of timber that is naturally resistant to termites such as ''Canarium australianum'' (Turpentine Tree), ''Callitris glaucophylla'' (White Cypress), or one of the ''Sequoias''. 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]]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]], insect growth regulators ([[hormone]]s) such as triflumuron and, more recently, [[fipronil]]. Blowing dusts into termite workings is a highly skilled process. All these slow-acting [[poison]]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]], [[hexaflumuron]], and novaflumuron as bait toxicants and [[fipronil]] and [[imidacloprid]] as soil poisons. Soil poisons are the least-preferred method of control as this requires much larger doses of toxin and results in uncontrollable release to the environment. ===Termites in the human diet=== 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 [[Central Africa|Central]] and [[Southern Africa]] 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; [[cooking oil|oil]] 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. == Ecology == Ecologically, termites are important in [[nutrient]] recycling, [[Habitat (ecology)|habitat]] creation, [[soil]] 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 [[floodplain|seasonally inundated]] 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]]s, [[lizard]]s, snakes, small mammals, and birds live in abandoned or weathered mounds, and [[aardvark]]s dig substantial caves and burrows in them, which then become homes for larger animals such as [[hyena]]s and [[mongoose]]s. As [[detrivore]]s, 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 USA. Globally, termites are found roughly between 50 degrees North & South, with the greatest [[biomass]] 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]], one of the prime [[greenhouse gas]]es. Termites have been common since at least the [[Cretaceous]] 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. <ref>[http://209.85.173.104/search?q=cache:skbJ_ftgqZ8J:geologyindy.byu.edu/students/Anne/Termites%2520and%2520Dinosaurs%2520ms.pdf+termite+paleoecology&hl=en&ct=clnk&cd=7&gl=us]</ref> ===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]]s, [[resin]]s, and [[lignin]]s) into the woody cell walls. This reduces the ability of termites to efficiently digest the [[cellulose]]. 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 (plant)|Eremophila]] that repels termites.<ref>[http://www.abc.net.au/news/newsitems/200511/s1507502.htm Plant extract stops termites dead]</ref> Tests have shown that termites are strongly repelled by the toxic material to the extent that they will starve rather than cross treated samples. When kept in close proximity to the extract, they become disoriented and eventually die. Scientists hope to use this toxic compound commercially to prevent termite feeding. ==Taxonomy, evolution and systematics== [[Image:Mastotermes darwiniensis.jpg|thumb|left|The famous [[Giant Northern Termite]] ''Mastotermes darwiniensis'' attests to the close relationship of termites and cockroaches.]] Recent DNA evidence{{Verify source|date=March 2008}}<!-- 1. What DNA? 2. Cladistic analysis is inference not evidence; if in doubt use terms like "data", "studies" or "analyses" --> has supported the nearly 120-year-old hypothesis, originally based on morphology, that termites are most closely related to the wood-eating [[cockroach]]es (genus ''[[Cryptocercus]]''), 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 [[Blattaria]], which contains cockroaches <ref>{{cite news | title = Termites are 'social cockroaches' | language = English | publisher = BBC News | date = [[13 April]] [[2007]] | url = http://news.bbc.co.uk/1/hi/sci/tech/6553219.stm | accessdate = }}</ref><ref>Eggleton, P. &al. (2007), ''Biological Letters'', [[June 7]], cited in ''Science News'' vol. 171, p. 318 </ref>. However, most researchers advocate the less drastic measure of retaining the termites as Isoptera but as a group subordinate to true roaches, preserving the internal classification of termites <ref> Lo, N. &al. (2007), ''Biology Letters'', [[14 August]] [[2007]], doi 10.1098/rsbl.2007.0264 </ref>. <!-- Phylogeny predicted by Deitz in 2003: http://globiz.sachsen.de/snsd/publikationen/ArthropodSystematicsPhylogeny/EA_61_1/EA_61_1-69-91_Deitz.pdf --> [[Image:Copal with insects close-up.jpg|300px|thumb|Termites and other insects in [[copal]]]] === Evolutionary history === The oldest unambiguous termite [[fossils]] date to the early [[Cretaceous]], although structures from the late Triassic have been interpreted as fossilized termite nests.<ref>Gay and Calaby 1970 Termites of the Australian region. in; Krishna K Weesner FM eds. Biology of Termites, Vol. II Academic Press NY p401</ref> Given the diversity of Cretaceous termites, it is likely that they had their origin at least sometime in the [[Jurassic]]. It has long been accepted that termites are closely related to [[cockroach]]es and [[mantid]]s, and they are classified in the same superorder ([[Dictyoptera]]), but new research has shed light on the details of termite evolution.<ref>[http://mbe.oxfordjournals.org/cgi/reprint/20/6/907.pdf Evidence for Cocladogenesis Between Diverse Dictyopteran Lineages and Their Intracellular Endosymbionts]</ref> 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]]'', 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. As mentioned above, the primitive [[Giant Northern Termite]] (''Mastotermes darwiniensis'') exhibits numerous cockroach-like characteristics that are not shared with other termites. ===Systematics=== As of 1996, about 2,800 termite [[species]] 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 (evolution)|basal]] to the most advanced: * [[Mastotermitidae]] (1 species, ''[[Mastotermes darwiniensis]]'') * [[Hodotermitidae]] (3 genera, 19 species) ** [[Hodotermitinae]] * [[Kalotermitidae]] (22 genera, 419 species) * [[Termopsidae]] (5 genera, 20 species) ** [[Termopsinae]] ** [[Porotermitinae]] ** [[Stolotermitinae]] * [[Rhinotermitidae]] (14 genera, 343 species) ** [[Coptotermitinae]] <small>Holmgren</small> ** [[Heterotermitinae]] <small>Froggatt</small> ** [[Prorhinoterminae]] <small>Quennedey & Deligne, 1975</small> ** [[Psammotermitinae]] <small>Holmgren</small> ** [[Rhinotermitinae]] <small>Froggatt</small> ** [[Stylotermitinae]] <small>Holmgren, K & N, 1917</small> ** [[Termitogetoninae]] <small>Holmgren</small> * [[Serritermitidae]] (1 species, ''[[Serritermes serrifer]]'') * [[Termitidae]] (236 genera, 1958 species) ** [[Macrotermitinae]] (14 genera, 349 species) ** [[Nasutitermitinae]] (91 genera, 663 species) ** [[Amitermitinae]] (17 genera, 295 species) ** [[Apicotermitinae]] (43 genera, 202 species) ** [[Cubitermitinae]] (28 genera, 161 species) ** [[Termitinae]] (43 genera, 288 species) The most current classification of termites is summarized by Engel & Krishna (2004). ==Termites as a source of power== The [[US 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]].<ref>[http://www.jgi.doe.gov JGI - Organization responsible for sequencing the termite.]</ref> Termites may produce up to two litres of [[hydrogen]] from digesting a single sheet of paper, making them one of the planet's most efficient [[bioreactor]]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]]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 [[Termite Hindgut Microbiology|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]], in commercial quantities. Sceptics regard this as unlikely to become a carbon-neutral commercial process due to the energy inputs required to maintain the system. For decades, researchers have sought to house termites on a commercial scale (like [[worm farm]]s) 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]] have not been overcome.<ref>[http://www.jgi.doe.gov/education/bioenergy/bioenergy_4.html Original article on termites as bioreactors]</ref> ==See also== *[[Decompiculture]] *[[Stigmergy]] *[[Xylophagy]] ==References== *{{cite book|author=[[David Grimaldi|Grimaldi, D.]] and [[Michael S. Engel|Engel, M.S.]] |title=Evolution of the Insects|year=[[2005]]|publisher=[[Cambridge University Press]]|id=ISBN 0-521-82149-5}} * {{cite journal | quotes=no |author=[[Michael S. Engel|Engel, M.S.]] and K. Krishna |year=2004 |title=Family-group names for termites (Isoptera) |journal=[[American Museum Novitates]] |volume=3432 |pages=1–9 |doi=10.1206/0003-0082(2004)432<0001:FNFTI>2.0.CO;2}} * [http://www.earthlife.net/insects/isoptera.html Earthlife] * [http://www.drdons.net/Dr_Dons_Termite_Glossary.htm Termite terms] * [http://www.gsajournals.org/gsaonline/?request=get-abstract&doi=10.1130%2F0091-7613(1986)14%3C87:OTNFTU%3E2.0.CO%3B2 Cretaceous termites] {{reflist}} ==Further reading== {{cite book|author=Abe T., Bignell D.E., Higashi M. (eds.)|title=[http://books.google.com/books?hl=en&lr=&id=Q_-KUDz2-sYC&oi=fnd&pg=PR11&dq=Paul+Eggleton+termites+evolution&ots=DS5sbRP5Qb&sig=6mOUUJ43YxE0ZqD8Dpi1TYUk3dI#PPR8,M1 Termites: evolution, sociality, symbioses, ecology]|year=2000|publisher=Kluwer academic publishers|id=ISBN 0792363612}} ==External links== {{wikispecies|Isoptera}} {{commonscat|Isoptera}} *[http://ianrpubs.unl.edu/Insects/g1062.htm University of Nebraska page on Termites] *[http://drdons.net/control.htm A summary of termite control methods] *[http://www.ipm.ucdavis.edu/PMG/PESTNOTES/pn7440.html University of California advice on Drywood Termites] *[http://npic.orst.edu/pest2.htm#termites Termite Pest Control Information - USA National Pesticide Information Center] *[http://www.unb.br/ib/zoo/docente/constant/catal/catnew.html Catalogue of the termites of the World] *[http://lancaster.unl.edu/pest/termite.shtml Pictures of termites] *[http://darwinblog.blogspot.com/2005/12/transitional-species-in-insect_24.html Transitional Species in Insect Evolution] *[http://members.tripod.com/~charles_W/laterite.html Did the Termites Cause Laterites?] *[http://www.angelfire.com/nc/isoptera/soil.html Cretaceous Termites and Soil Phosphorus] *[http://www.utoronto.ca/forest/termite/benefice.htm Beneficial Uses of Termites] *[http://termite.tamu.edu/index.html Texas A&M University Department of Entomology - Center for Urban & Structural Entomology] *[http://www.zpluspartners.com/zblog/archive/2004_01_24_zblogarchive.html Biomimicry used in Eastgate Building, Zimbabwe] *[http://journeytoforever.org/farm_library/Marais1/whiteantToC.html The Soul of the White Ant - Eugène N. Marais] *[http://www.utoronto.ca/forest/termite/termite.html Urban Entomology Program University of Toronto] *[http://www.uos.harvard.edu/ehs/pes_termites.shtml Harvard University fact sheet on Eastern Subterranean Termites] *[http://www.ento.csiro.au/education/insects/isoptera.html Isoptera: termites (CSIRO Australia Entomology)]. *[http://www.physorg.com/news3700.html 'Termite guts can save the planet', says Nobel laureate] *[http://www.pestworld.org/database/FactSheet.asp?ID=135&UserType= USA Pest Management Association's fact sheet on termites] *[http://www.bugfacts.org/category.php?category=Termites BugFacts --Information from a US chemical supply company] *[http://www.termiteinformation.info --Termite related opinion with a USA bias] [[Category:Termites| ]] [[Category:Insects]] [[Category:Pest insects]] [[Category:Building defects]] [[Category:Invasive animal species]] [[ar:نمل أبيض]] [[br:Laou-koad]] [[ca:Tèrmit]] [[cs:Termiti]] [[da:Termit (insekt)]] [[de:Termiten]] [[et:Termiidilised]] [[es:Isoptera]] [[eo:Termito]] [[fa:موریانه]] [[fr:Isoptera]] [[ko:흰개미]] [[io:Termito]] [[it:Isoptera]] [[he:טרמיטאים]] [[ht:Tèmit]] [[lt:Termitai]] [[ln:Ndɔngɛ]] [[hu:Termeszek]] [[nl:Termieten]] [[ja:シロアリ]] [[no:Termitter]] [[nn:Termittar]] [[oc:Isoptera]] [[pl:Termity]] [[pt:Cupim]] [[ru:Термиты]] [[simple:Termite]] [[sk:Termity]] [[sl:Termiti]] [[sr:Термит]] [[su:Rinyuh]] [[fi:Termiitit]] [[sv:Termiter]] [[vi:Mối]] [[tr:Termit (zooloji)]] [[ur:دیمک]] [[zh:白蟻]]