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75-556: Odontotermes is a termite genus belonging to subfamily Macrotermitinae (fungus-growing termites), which is native to the Old World . They are most destructive in wooden homes, and are agricultural pests in the tropics and subtropics of Africa and Asia. It is the most diverse termite genus in Africa, with 78 species recorded (as of 2002). Their underground nests form a slight mound above ground, which may be covered in grass. In large colonies,

150-514: A 'superorganism'". He concludes that "the superorganism analogy is unwarranted". Some scientists have suggested that individual human beings can be thought of as "superorganisms"; as a typical human digestive system contains 10 to 10 microorganisms whose collective genome , the microbiome studied by the Human Microbiome Project , contains at least 100 times as many genes as the human genome itself. Salvucci wrote that superorganism

225-419: A brief amount of time, so they also rely on their legs. The appearance of the legs is similar in each caste, but the soldiers have larger and heavier legs. The structure of the legs is consistent with other insects: the parts of a leg include a coxa , trochanter , femur , tibia and the tarsus . The number of tibial spurs on an individual's leg varies. Some species of termite have an arolium, located between

300-447: A certain developmental path. Workers may also be fertile or sterile, however the term "worker" is normally reserved for the latter, having evolved in taxa that exhibit a bifurcated developmental pathway. As a result, sterile workers like in the family Termitidae are termed true workers and are the most derived, while those that are undifferentiated and fertile as in the wood-nesting Archotermopsidae are termed pseudergates , which are

375-472: A developmental process called incomplete metamorphosis , going through multiple gradual pre-adult molts that are highly developmentally plastic before becoming an adult. Unlike in other hemimetabolous insects, nymphs are more strictly defined in termites as immature young with visible wing buds, which often invariably go through a series of moults to become winged adults . Larvae , which are defined as early nymph instars with absent wing buds, exhibit

450-425: A few hundred individuals to enormous societies with several million individuals. Most species are rarely seen, having a cryptic life-history where they remain hidden within the galleries and tunnels of their nests for most of their lives. Termites' success as a group has led to them colonizing almost every global landmass, with the highest diversity occurring in the tropics where they are estimated to constitute 10% of

525-513: A fontanelle, which appears as a circular pore or series of pores in a depressed region within the middle of the head. The fontanelle connects to the frontal gland, a novel organ unique to Neoisopteran termites which evolved to excrete an array of defensive chemicals and secretions, and so is typically most developed in the soldier caste. Cellulose digestion in the family Termitidae has co-evolved with bacterial gut microbiota and many taxa have evolved additional symbiotic relationships such as with

600-511: A new nest site. In challenging environments, micro organisms collaborate and evolve together to process unlikely sources of nutrients such as methane. This process called syntrophy ("eating together") might be linked to the evolution of eukaryote cells and involved in the emergence or maintenance of life forms in challenging environments on Earth and possibly other planets. Superorganisms tend to exhibit homeostasis , power law scaling, persistent disequilibrium and emergent behaviours. The term

675-454: A pedicel (typically shorter than the scape), and the flagellum (all segments beyond the scape and pedicel). The mouth parts contain a maxillae , a labium, and a set of mandibles . The maxillae and labium have palps that help termites sense food and handling. The cuticle of most castes is soft and flexible due to a resulting lack of sclerotization, particularly of the abdomen which often appears translucent. Pigmentation and sclerotization of

750-404: A termite is in flight, its wings remain at a right angle, and when the termite is at rest, its wings remain parallel to the body. Due to termites being hemimetabolous insects , where the young go through multiple and gradual adultoid molts before becoming an adult, the advent of eusociality has significantly altered the developmental patterns of this group of insects of which, although similar,

825-466: A wide variety of predators. The queens can be particularly long-lived for insects, with some reportedly living as long as 30 or 50 years. In both the linear and bifurcated developmental pathways, the primary reproductives only develop from winged immatures (nymphs). These winged immatures are capable of regressively molting into a form known as brachypterous neotenics ( nymphoids ), which retain juvenile and adult characteristics. BN 's can be found in both

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900-590: Is a "difference between emergence and reductionism". The economist Carl Menger expanded upon the evolutionary nature of much social growth but never abandoned methodological individualism . Many social institutions arose, Menger argued, not as "the result of socially teleological causes, but the unintended result of innumerable efforts of economic subjects pursuing 'individual' interests". Both Spencer and Menger argued that because individuals choose and act, any social whole should be considered less than an organism, but Menger emphasized that more strongly. Spencer used

975-432: Is a mixture of benzoquinone and protein. Their only food is the fungus grown in the fungal garden at the center of the nest. The fungus is cultivated on a substrate of wood , bark , leaf litter , dry dung, and dead grass. These are plastered with cement where they are obtained, which facilitates diurnal foraging. Odontotermes species are major contributors to litter decomposition. The fungus Termitomyces reticulatus

1050-412: Is another level of integration that is observed in nature. These levels include the genomic, the organismal and the ecological levels. The genomic structure of organisms reveals the fundamental role of integration and gene shuffling along evolution. The 19th-century thinker Herbert Spencer coined the term super-organic to focus on social organization (the first chapter of his Principles of Sociology

1125-477: Is based on the idea that a biological system – in order to be effective – needs a sub-system of cybernetic communications and control. This is demonstrated in the way a mole rat colony uses functional synergy and cybernetic processes together. Joël de Rosnay also introduced a concept called "cybionte" to describe cybernetic superorganism. The notion associates superorganism with chaos theory , multimedia technology, and other new developments. If Col. Thorpe [of

1200-696: Is believed to be from the Upper Cretaceous in West Texas , where the oldest known faecal pellets were also discovered. Claims that termites emerged earlier have faced controversy. For example, F. M. Weesner indicated that the Mastotermitidae termites may go back to the Late Permian , 251 million years ago, and fossil wings that have a close resemblance to the wings of Mastotermes of the Mastotermitidae,

1275-468: Is consistent with all four-winged insects. The mesothorax and metathorax have well-developed exoskeletal plates; the prothorax has smaller plates. Termites have a ten-segmented abdomen with two plates, the tergites and the sternites . The tenth abdominal segment has a pair of short cerci . There are ten tergites, of which nine are wide and one is elongated. The reproductive organs are similar to those in cockroaches but are more simplified. For example,

1350-552: Is divided into the following clade and family groups, showing the subfamilies in their respective classification: The Neoisoptera , literally meaning "newer termites" (in an evolutionary sense), are a recently coined clade that include families such as the Rhinotermitidae and Termitidae . Neoisopterans have a bifurcated caste development with true workers, and so notably lack pseudergates (except in some basal taxa such as Serritermitidae : see below ). All Neoisopterans have

1425-446: Is entitled "Super-organic Evolution" ), though this was apparently a distinction between the organic and the social, not an identity: Spencer explored the holistic nature of society as a social organism while distinguishing the ways in which society did not behave like an organism. For Spencer, the super-organic was an emergent property of interacting organisms, that is, human beings. And, as has been argued by D. C. Phillips, there

1500-538: Is expected to increase over time as a consequence of climate change . Increased urbanization and connectivity is also predicted to expand the range of some pest termites. The infraorder name Isoptera is derived from the Greek words iso (equal) and ptera (winged), which refers to the nearly equal size of the fore and hind wings. "Termite" derives from the Latin and Late Latin word termes ("woodworm, white ant"), altered by

1575-628: Is found in association with O. badius and O. transvaalensis in Africa. Species include: Termite Termites are a group of detritophagous eusocial insects which consume a variety of decaying plant material , generally in the form of wood , leaf litter , and soil humus . They are distinguished by their moniliform antennae and the soft-bodied and often unpigmented worker caste for which they have been commonly termed " white ants "; however, they are not ants , to which they are only distantly related. About 2,972 extant species are currently described, 2,105 of which are members of

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1650-494: Is found mainly in the derived taxa (i.e. Neoisoptera ), and is believed to have evolved in tandem with the sterile worker caste as species moved to foraging for food beyond their nests, as opposed to the nest also being the food (such as in obligate wood-dwellers). There are three main castes which are discussed below: Worker termites undertake the most labor within the colony, being responsible for foraging, food storage, and brood and nest maintenance. Workers are tasked with

1725-422: Is not homologous to that of the eusocial Hymenoptera . Unlike ants, bees, and wasps which undergo a complete metamorphosis and as a result only exhibit developmental plasticity at the immobile larval stage , the mobile adultoid instars of termites remain developmentally flexible throughout all life stages up to the final molt , which has uniquely allowed for the evolution of distinct yet flexible castes amongst

1800-498: Is used most often to describe a social unit of eusocial animals in which division of labour is highly specialised and individuals cannot survive by themselves for extended periods. Ants are the best-known example of such a superorganism. A superorganism can be defined as "a collection of agents which can act in concert to produce phenomena governed by the collective", phenomena being any activity "the hive wants" such as ants collecting food and avoiding predators , or bees choosing

1875-521: The nymphal (winged) and apterous (wingless) lines. The bifurcation occurs early, either at the egg or the first two instars, and represents an irreversible and committed development to either the reproductive or non-reproductive lifestyles. As such, the apterous lineage consists mostly of wingless and truly altruistic sterile individuals (true workers, soldiers), whereas the nymphal lineage consists mainly of fertile individuals destined to become winged reproductives. The bifurcated developmental pathway

1950-469: The Archotermopsidae , which are able to retain high developmental plasticity even up to the late instars. In these basal taxa, the immatures are able to go through progressive ( nymph-to-imago ), regressive ( winged-to-wingless ) and stationary ( size increase, remains wingless ) molts, which typically indicates the developmental trajectory an individual follows. There is significant variation of

2025-657: The Nasutes , which have a horn-like nozzle projection ( nasus ) on the head. These unique soldiers are able to spray noxious, sticky secretions containing diterpenes at their enemies. Nitrogen fixation plays an important role in Nasute nutrition. Soldiers are normally a committed sterile caste and so do not molt into anything else, but in certain basal taxa like the Archotermopsidae they are known to rarely molt into neotenic forms that develop functional sexual organs. In species with

2100-403: The claws , which is present in species that climb on smooth surfaces but is absent in most termites. Unlike in ants, the hind-wings and fore-wings are of equal length. Most of the time, the alates are poor flyers; their technique is to launch themselves in the air and fly in a random direction. Studies show that in comparison to larger termites, smaller termites cannot fly long distances. When

2175-432: The eusocial termite Reticulitermes speratus are capable of a long lifespan without sacrificing fecundity . These long-lived queens have a significantly lower level of oxidative damage, including oxidative DNA damage , than workers, soldiers and nymphs. The lower levels of damage appear to be due to increased catalase , an enzyme that protects against oxidative stress . Superorganism The term superorganism

2250-503: The intromittent organ is not present in male alates, and the sperm is either immotile or aflagellate. However, Mastotermitidae termites have multiflagellate sperm with limited motility . The genitals in females are also simplified. Unlike in other termites, Mastotermitidae females have an ovipositor , a feature strikingly similar to that in female cockroaches. The non-reproductive castes of termites are wingless and rely exclusively on their six legs for locomotion. The alates fly only for

2325-436: The "king" and "queen" that together form a lifelong monogamous pair. Also unlike ants, which undergo a complete metamorphosis , termites undergo an incomplete metamorphosis that proceeds through egg, nymph , and adult stages. Termite colonies are commonly described as superorganisms due to the collective behaviors of the individuals which form a self-governing entity: the colony itself. Their colonies range in size from

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2400-582: The 2,972 extant termite species currently classified, 1,000 are found in Africa , where mounds are extremely abundant in certain regions. Approximately 1.1 million active termite mounds can be found in the northern Kruger National Park alone. In Asia , there are 435 species of termites, which are mainly distributed in China . Within China, termite species are restricted to mild tropical and subtropical habitats south of

2475-668: The Paleozoic and Triassic insects tentatively classified as termites are in fact unrelated to termites and should be excluded from the Isoptera. Other studies suggest that the origin of termites is more recent, having diverged from Cryptocercus sometime during the Early Cretaceous . The primitive giant northern termite ( Mastotermes darwiniensis ) exhibits numerous cockroach-like characteristics that are not shared with other termites, such as laying its eggs in rafts and having anal lobes on

2550-789: The Yangtze River. In Australia , all ecological groups of termites (dampwood, drywood, subterranean) are endemic to the country, with over 360 classified species. Because termites are highly social and abundant, they represent a disproportionate amount of the world's insect biomass . Termites and ants comprise about 1% of insect species, but represent more than 50% of insect biomass. Due to their soft cuticles, termites do not inhabit cool or cold habitats. There are three ecological groups of termites: dampwood, drywood and subterranean. Dampwood termites are found only in coniferous forests, and drywood termites are found in hardwood forests; subterranean termites live in widely diverse areas. One species in

2625-528: The animal biomass , particularly in Africa which has the richest diversity with more than 1000 described species. They are important decomposers of decaying plant matter in the subtropical and tropical regions of the world, and their recycling of wood and plant matter is of considerable ecological importance. Many species are ecosystem engineers capable of altering soil characteristics such as hydrology , decomposition, nutrient cycling , vegetative growth, and consequently surrounding biodiversity through

2700-458: The colony. Workers remain wingless and across many taxa become developmentally arrested, appearing to not change into any other caste until death. In some basal taxa, there is no distinction, with the "workers" (pseudergates) essentially being late instar larvae that retain the ability to change into all other castes. The development of larvae into adults can take months; the time period depends on food availability and nutrition, temperature, and

2775-431: The cuticle correlates with life history , with species that spend more time in the surface in the open tending to have a more sclerotized and pigmented exoskeleton. Consistent with all insects, the anatomy of the termite thorax consists of three segments: the prothorax , the mesothorax and the metathorax . Each segment contains a pair of legs . On alates, the wings are located at the mesothorax and metathorax, which

2850-439: The derived and basal termite taxa, and generally serve as supplementary reproductives. Termites are often compared with the social Hymenoptera (ants and various species of bees and wasps), but their differing evolutionary origins result in major differences in life cycle. In the eusocial Hymenoptera, the workers are exclusively female. Males (drones) are haploid and develop from unfertilised eggs, while females (both workers and

2925-527: The descendants of the genus Cryptocercus . Some researchers have suggested a more conservative measure of retaining the termites as the Termitoidae, an epifamily within the cockroach order, which preserves the classification of termites at family level and below. Termites have long been accepted to be closely related to cockroaches and mantids , and they are classified in the same superorder ( Dictyoptera ). The oldest unambiguous termite fossils date to

3000-496: The developmental patterns in termites even across closely related taxa, but can typically be generalized into the following two patterns: The first is the linear developmental pathway , where all immatures are capable of developing into winged adults ( Alates ), exhibit high phenotypic potency, and where there exists no true sterile caste other than the soldier. The second is the bifurcated developmental pathway , where immatures diverge into two distinct developmental lineages known as

3075-418: The digestion of cellulose in food and are thus the most likely caste to be found in infested wood. The process of worker termites feeding other nestmates is known as trophallaxis . Trophallaxis is an effective nutritional tactic to convert and recycle nitrogenous components. It frees the parents from feeding all but the first generation of offspring, allowing for the group to grow much larger and ensuring that

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3150-533: The drywood group is the West Indian drywood termite ( Cryptotermes brevis ) , which is an invasive species in Australia. Termites are usually small, measuring between 4 and 15 millimetres ( 3 ⁄ 16 and 9 ⁄ 16  in) in length. The largest of all extant termites are the queens of the species Macrotermes bellicosus , measuring up to over 10 centimetres (4 in) in length. Another giant termite,

3225-579: The dynamics of a complex system . The concept of a superorganism raises the question of what is to be considered an individual . Toby Tyrrell's critique of the Gaia hypothesis argues that Earth's climate system does not resemble an animal's physiological system. Planetary biospheres are not tightly regulated in the same way that animal bodies are: "planets, unlike animals, are not products of evolution. Therefore we are entitled to be highly skeptical (or even outright dismissive) about whether to expect something akin to

3300-620: The early Cretaceous , but given the diversity of Cretaceous termites and early fossil records showing mutualism between microorganisms and these insects, they possibly originated earlier in the Jurassic or Triassic. Possible evidence of a Jurassic origin is the assumption that the extinct mammaliaform Fruitafossor from Morrison Formation consumed termites, judging from its morphological similarity to modern termite-eating mammals. Morrison Formation also yields social insect nest fossils close to that of termites. The oldest termite nest discovered

3375-671: The extinct Gyatermes styriensis , flourished in Austria during the Miocene and had a wingspan of 76 millimetres (3 in) and a body length of 25 millimetres (1 in). Most worker and soldier termites are completely blind as they do not have a pair of eyes. However, some species, such as Hodotermes mossambicus , have compound eyes which they use for orientation and to distinguish sunlight from moonlight. The alates (winged males and females) have eyes along with lateral ocelli . Lateral ocelli, however, are not found in all termites, absent in

3450-409: The families Hodotermitidae , Termopsidae , and Archotermopsidae . Like other insects, termites have a small tongue-shaped labrum and a clypeus ; the clypeus is divided into a postclypeus and anteclypeus. Termite antennae have a number of functions such as the sensing of touch, taste, odours (including pheromones), heat and vibration. The three basic segments of a termite antenna include a scape ,

3525-424: The family Termitidae . Termites comprise the infraorder Isoptera , or alternatively the epifamily Termitoidae , within the order Blattodea (along with cockroaches ). Termites were once classified in a separate order from cockroaches, but recent phylogenetic studies indicate that they evolved from cockroaches, as they are deeply nested within the group, and the sister group to wood eating cockroaches of

3600-1011: The first fully sequenced termite genome, of Zootermopsis nevadensis , which was published in the journal Nature Communications , consists of roughly 500Mb, while two subsequently published genomes, Macrotermes natalensis and Cryptotermes secundus , are considerably larger at around 1.3Gb. External phylogeny showing relationship of termites with other insect groups:  (Mantises) Blaberoidea Corydiodea Blattoidae Lamproblattidae Cryptocercidae (brown-hooded cockroaches) Isoptera (Termites) Internal phylogeny showing relationship of extant termite families: Mastotermitidae Stolotermitidae Hodotermopsidae Hodotermitidae Archotermopsidae Kalotermitidae Stylotermitidae Serritermitidae Rhinotermitidae Termitidae There are currently 3,173 living and fossil termite species recognised, classified in 12 families; reproductive and/or soldier castes are usually required for identification. The infraorder Isoptera

3675-551: The forehead that lead to a gland which exudes defensive secretions, are a feature of the clade Neoisoptera and are present in all extant taxa such as Rhinotermitidae. The majority of termite species have mandibulate soldiers which are easily identified by the disproportionately large sclerotized head and mandibles. Among certain termites, the soldier caste has evolved globular (phragmotic) heads to block their narrow tunnels such as seen in Cryptotermes . Amongst mandibulate soldiers,

3750-677: The fungus Termitomyces ; in contrast, basal Neoisopterans and all other Euisoptera have flagellates and prokaryotes in their hindguts. Extant families and subfamilies are organized as follows: Termites are found on all continents except Antarctica . The diversity of termite species is low in North America and Europe (10 species known in Europe and 50 in North America), but is high in South America , where over 400 species are known. Of

3825-684: The genus Cryptocercus . Previous estimates suggested the divergence took place during the Jurassic or Triassic . More recent estimates suggest that they have an origin during the Late Jurassic , with the first fossil records in the Early Cretaceous . Similarly to ants and some bees and wasps from the separate order Hymenoptera , most termites have an analogous "worker" and "soldier" caste system consisting of mostly sterile individuals which are physically and behaviorally distinct. Unlike ants, most colonies begin from sexually mature individuals known as

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3900-598: The highest developmental potentiality and are able to molt into Alates , Soldiers , Neotenics , or Workers . Workers are believed to have evolved from larvae, sharing many similarities to the extent that workers can be regarded as "larval", in that both lack wings, eyes, and functional reproductive organs while maintaining varying levels of developmental flexibility, although usually to a much lesser extent in workers. The main distinction being that while larvae are wholly dependent on other nestmates to survive, workers are independent and are able to feed themselves and contribute to

3975-639: The highest phenotypic potency and can be described as totipotent ( able to molt into all alternative phenotypes ), whereas following instars can be pluripotent ( able to molt into reproductives and non-reproductives but cannot molt into at least one phenotype ), to multipotent ( able to molt into either reproductive or non-reproductive phenotypes ), to unipotent ( able to molt into developmentally close phenotypes ), and then finally committed ( no longer able to change phenotype, functionally an adult.) In most termites, phenotypic potency decreases with every successive molt. Notable exceptions are basal taxa such as

4050-436: The immatures. As a result the caste system of termites consists mostly of neotenous or juvenile individuals that undertake the most labor in the colony, which is in contrast to the eusocial Hymenoptera where work is strictly undertaken by the adults. The developmental plasticity in termites can be described similarly to cell potency , where each molt offers a varying level of phenotypic potency. Early instars typically exhibit

4125-536: The influence of Latin terere ("to rub, wear, erode") from the earlier word tarmes . A termite nest is also known as a termitary or termitarium (plural termitaria or termitariums ). The word was first used in English in 1781. Earlier attested designations were "wood ants" or "white ants", though these may never have been in wide use as termites do not exist in the British Isles . Termites were formerly placed in

4200-564: The issue may be terminological. The term superorganic was adopted by the anthropologist Alfred L. Kroeber in 1917. Social aspects of the superorganism concept are analysed by Alan Marshall in his 2002 book "The Unity of Nature". Finally, recent work in social psychology has offered the superorganism metaphor as a unifying framework to understand diverse aspects of human sociality, such as religion, conformity, and social identity processes. Superorganisms are important in cybernetics , particularly biocybernetics , since they are capable of

4275-399: The king will continue to mate with the queen throughout their lives. In some species, the abdomen of the queen swells up dramatically to increase fecundity , a characteristic known as physogastrism . Depending on the species, the queen starts producing reproductive alates at a certain time of the year, and huge swarms emerge from the colony when nuptial flight begins. These swarms attract

4350-568: The large mounds constructed by certain species. Termites have several impacts on humans. They are a delicacy in the diet of some human cultures such as the Makiritare in the Alto Orinoco province of Venezuela , where they are commonly used as a spice. They are also used in traditional medicinal treatments of various diseases and ailments, such as influenza, asthma, bronchitis, etc. Termites are most famous for being structural pests; however,

4425-829: The level of developmental plasticity even between closely related taxa, with many species having true workers that can molt into the other apterous castes such as ergatoids (worker reproductive; apterous neotenics), soldiers, or the other worker castes. Pseudergates sensu stricto are individuals which arise from the linear developmental pathway that have regressively molted and lost their wing buds, and are regarded as totipotent immatures. They are capable of performing work but are overall less involved in labor and considered more cooperative than truly altruistic. Pseudergates sensu lato , otherwise known as false workers , are most represented in basal lineages ( Kalotermitidae , Archotermopsidae , Hodotermopsidae , Serritermitidae ) and closely resemble true workers in which they also perform most of

4500-439: The linear developmental pathway, soldiers develop from apterous immatures and constitute the only true sterile caste in these taxa. The primary reproductive caste of a colony consists of the fertile adult ( imago ) female and male individuals, colloquially known as the queen and king. The queen of the colony is responsible for egg production of the colony. Unlike in ants, the male and female reproductives form lifelong pairs where

4575-444: The mandibles have been adapted for a variety of defensive strategies: Biting/crushing ( Incisitermes ), slashing ( Cubitermes ), slashing/snapping ( Dentispicotermes ), symmetrical snapping ( Termes ), asymmetrical snapping ( Neocapritermes ), and piercing ( Armitermes ). In the more derived termite taxa, the soldier caste can be polymorphic and include minor and major forms. Other morphologically specialized soldiers includes

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4650-412: The most basal. True workers are individuals which irreversibly develop from the apterous lineage and have completely forgo development into a winged adult. They display altruistic behaviors and either have terminal molts or exhibit a low level of phenotypical potency. True workers across different termite taxa ( Mastotermitidae , Hodotermitidae , Rhinotermitidae & Termitidae) can widely vary in

4725-641: The most primitive living termite, have been discovered in the Permian layers in Kansas. It is even possible that the first termites emerged during the Carboniferous . The folded wings of the fossil wood roach Pycnoblattina , arranged in a convex pattern between segments 1a and 2a, resemble those seen in Mastotermes , the only living insect with the same pattern. Kumar Krishna et al. , though, consider that all of

4800-624: The mounds may be up to 6 m (20 ft) in diameter, and may be covered by shrubs and trees. Some species construct open chimneys or vent holes that descend into the mound. The fungal garden is enveloped by a thick layer of clay. The queen lives in a clay cell in the midst of the fungal garden at the center of the hive. The African species have a single soldier caste, unlike the related genus Macrotermes . Alarmed soldier termites of Odontotermes badius Haviland 1898 expel an odoriferous brown defensive secretion between their mandibles that becomes sticky and rubbery on exposure to air. The secretion

4875-461: The necessary gut symbionts are transferred from one generation to another. Workers are believed to have evolved from older wingless immatures ( Larvae ) that evolved cooperative behaviors; and indeed in some basal taxa the late instar larvae are known to undertake the role of workers without differentiating as a true separate caste. Workers can either be male or female, although in some species with polymorphic workers either sex may be restricted to

4950-470: The order Isoptera. As early as 1934 suggestions were made that they were closely related to wood-eating cockroaches (genus Cryptocercus , the woodroach) based on the similarity of their symbiotic gut flagellates. In the 1960s additional evidence supporting that hypothesis emerged when F. A. McKittrick noted similar morphological characteristics between some termites and Cryptocercus nymphs . In 2008 DNA analysis from 16S rRNA sequences supported

5025-419: The organistic idea to engage in extended analysis of social structure and conceded that it was primarily an analogy. For Spencer, the idea of the super-organic best designated a distinct level of social reality above that of biology and psychology, not a one-to-one identity with an organism. Nevertheless, Spencer maintained that "every organism of appreciable size is a society", which has suggested to some that

5100-583: The position of termites being nested within the evolutionary tree containing the order Blattodea , which included the cockroaches. The cockroach genus Cryptocercus shares the strongest phylogenetical similarity with termites and is considered to be a sister-group to termites. Termites and Cryptocercus share similar morphological and social features: for example, most cockroaches do not exhibit social characteristics, but Cryptocercus takes care of its young and exhibits other social behaviour such as trophallaxis and allogrooming . Termites are thought to be

5175-415: The queen) are diploid and develop from fertilised eggs. In contrast, worker termites, which constitute the majority in a colony, are diploid individuals of both sexes and develop from fertilised eggs. Depending on species, male and female workers may have different roles in a termite colony. The life cycle of a termite begins with an egg , but is different from that of a bee or ant in that it goes through

5250-430: The size of the colony. Since larvae and nymphs are unable to feed themselves, workers must feed them, but workers also take part in the social life of the colony and have certain other tasks to accomplish such as foraging, building or maintaining the nest or tending to the queen. Pheromones regulate the caste system in termite colonies, preventing all but a very few of the termites from becoming fertile queens. Queens of

5325-472: The so-called " distributed intelligence ", a system composed of individual agents that have limited intelligence and information. They can pool resources and so can complete goals that are beyond reach of the individuals on their own. Existence of such behavior in organisms has many implications for military and management applications and is being actively researched. Superorganisms are also considered dependent upon cybernetic governance and processes. This

5400-460: The vast majority of termite species are innocuous, with the regional numbers of economically significant species being: North America , 9; Australia , 16; Indian subcontinent , 26; tropical Africa , 24; Central America and the West Indies , 17. Of known pest species, 28 of the most invasive and structurally damaging belong to the genus Coptotermes . The distribution of most known pest species

5475-447: The wings. It has been proposed that the Isoptera and Cryptocercidae be grouped in the clade " Xylophagodea ". Termites are sometimes called "white ants", but the only resemblance to the ants is due to their sociality which is due to convergent evolution with termites being the first social insects to evolve a caste system more than 100 million years ago. Termite genomes are generally relatively large compared to those of other insects;

5550-575: The work and are similarly altruistic, however differ in developing from the linear developmental pathway where they exist in a stationary molt; i.e they have halted development before the growth of wing buds, and are regarded as pluripotent immatures. The soldier caste is the most anatomically and behaviorally specialized, and their sole purpose is to defend the colony. Many soldiers have large heads with highly modified powerful jaws so enlarged that they cannot feed themselves. Instead, like juveniles, they are fed by workers. Fontanelles , simple holes in

5625-401: Was coined in 1789 by James Hutton , the "father of geology", to refer to Earth in the context of geophysiology . The Gaia hypothesis of James Lovelock , and Lynn Margulis as well as the work of Hutton, Vladimir Vernadsky and Guy Murchie , have suggested that the biosphere itself can be considered a superorganism, but that has been disputed. This view relates to systems theory and

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