Termites are small
insect
Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s that live in colonies and have distinct castes (
eusocial
Eusociality (from Greek εὖ ''eu'' "good" and social), the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping generat ...
) and feed on wood or other dead plant matter. Termites comprise the
infraorder
Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...
Isoptera, or alternatively the
epifamily
In biological classification, taxonomic rank is the relative level of a group of organisms (a taxon) in an ancestral or hereditary hierarchy. A common system consists of species, genus, family, order, class, phylum, kingdom, domain. While olde ...
Termitoidae, within the order
Blattodea
Blattodea is an order of insects that contains cockroaches and termites. Formerly, termites were considered a separate order, Isoptera, but genetic and molecular evidence suggests they evolved from within the cockroach lineage, cladistically ...
(along with
cockroach
Cockroaches (or roaches) are a paraphyletic group of insects belonging to Blattodea, containing all members of the group except termites. About 30 cockroach species out of 4,600 are associated with human habitats. Some species are well-known as ...
es). Termites were once classified in a separate
order from cockroaches, but recent
phylogenetic
In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
studies indicate that they evolved from cockroaches, as they are deeply nested within the group, and the
sister group
In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree.
Definition
The expression is most easily illustrated by a cladogram:
Taxon A and t ...
to wood eating cockroaches of the genus ''
Cryptocercus
''Cryptocercus'' is a genus of Dictyoptera (cockroaches and allies) and the sole member of its own family Cryptocercidae. Species are known as wood roaches or brown-hooded cockroaches. These roaches are subsocial, their young requiring considera ...
''. Previous estimates suggested the divergence took place during the
Jurassic
The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
or
Triassic
The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
. More recent estimates suggest that they have an origin during the
Late Jurassic
The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987.
In European lithostratigraphy, the name ...
,
with the first fossil records in the
Early Cretaceous
The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145 Ma to 100.5 Ma.
Geology
Pro ...
. About 3,106
species
In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
are currently described, with a few hundred more left to be described. Although these insects are often called "white ants", they are not
ant
Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22 ...
s, and are not closely related to ants.
Like ants and some
bees and
wasp
A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder. Th ...
s from the separate order
Hymenoptera
Hymenoptera is a large order (biology), order of insects, comprising the sawfly, sawflies, wasps, bees, and ants. Over 150,000 living species of Hymenoptera have been described, in addition to over 2,000 extinct ones. Many of the species are Par ...
, termites divide as "workers" and "soldiers" that are usually sterile. All colonies have fertile males called "kings" and one or more fertile females called "queens". Termites mostly feed on
dead plant material and
cellulose
Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell wall ...
, generally in the form of wood,
leaf litter
Plant litter (also leaf litter, tree litter, soil litter, litterfall or duff) is dead plant material (such as leaves, bark, needles, twigs, and cladodes) that have fallen to the ground. This detritus or dead organic material and its constituent ...
, soil, or animal dung. Termites are major
detritivore
Detritivores (also known as detrivores, detritophages, detritus feeders, or detritus eaters) are heterotrophs that obtain nutrients by consuming detritus (decomposing plant and animal parts as well as feces). There are many kinds of invertebrates, ...
s, particularly in the
subtropical
The subtropical zones or subtropics are geographical zone, geographical and Köppen climate classification, climate zones to the Northern Hemisphere, north and Southern Hemisphere, south of the tropics. Geographically part of the Geographical z ...
and
tropical
The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in
the Southern Hemisphere at S. The tropics are also referred to ...
regions, and their recycling of wood and plant matter is of considerable ecological importance.
Termites are among the most successful groups of insects on Earth, colonising most landmasses except
Antarctica
Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean, it contains the geographic South Pole. Antarctica is the fifth-largest contine ...
. Their colonies range in size from a few hundred individuals to enormous societies with several million individuals. Termite queens have the longest known lifespan of any insect, with some queens reportedly living up to 30 to 50 years. Unlike ants, which undergo a complete metamorphosis, each individual termite goes through an
incomplete metamorphosis
Hemimetabolism or hemimetaboly, also called incomplete metamorphosis and paurometabolism,McGavin, George C. ''Essential Entomology: An Order-by-Order Introduction''. Oxford: Oxford University Press, 2001. pp. 20. is the mode of development of certa ...
that proceeds through egg,
nymph
A nymph ( grc, νύμφη, nýmphē, el, script=Latn, nímfi, label=Modern Greek; , ) in ancient Greek folklore is a minor female nature deity. Different from Greek goddesses, nymphs are generally regarded as personifications of nature, are ty ...
, and adult stages. Colonies are described as
superorganism
A superorganism or supraorganism is a group of synergetically interacting organisms of the same species. A community of synergetically interacting organisms of different species is called a holobiont.
Concept
The term superorganism is used mo ...
s because the termites form part of a self-regulating entity: the colony itself.
Termites have several effects on humans. They are a delicacy in the diet of some human cultures and are used in many traditional medicines. Several hundred species are economically significant as pests that can cause serious damage to buildings, crops, or plantation forests. Some species, such as the
West Indian drywood termite
''Cryptotermes brevis'' is a species of termite in the family Kalotermitidae, commonly known as the West Indian drywood termite or the powderpost termite. It is able to live completely inside timber structures or articles made of wood such as fu ...
(''Cryptotermes brevis''), are regarded as
invasive species
An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species ad ...
.
Etymology
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
Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
and
Late Latin
Late Latin ( la, Latinitas serior) is the scholarly name for the form of Literary Latin of late antiquity.Roberts (1996), p. 537. English dictionary definitions of Late Latin date this period from the , and continuing into the 7th century in t ...
word ''termes'' ("woodworm, white ant"), altered by the influence of Latin ''terere'' ("to rub, wear, erode") from the earlier word ''tarmes''. A termite nest is also known as a ''termitary'' or ''termitarium'' (plural ''termitaria'' or ''termitariums''). In earlier English, termites were known as "wood ants" or "white ants".
The modern term was first used in 1781.
Taxonomy and evolution
![Mastotermes darwiniensis](https://upload.wikimedia.org/wikipedia/commons/a/a3/Mastotermes_darwiniensis.jpg)
Termites were formerly placed in the order Isoptera. As early as 1934 suggestions were made that they were closely related to wood-eating cockroaches (genus ''
Cryptocercus
''Cryptocercus'' is a genus of Dictyoptera (cockroaches and allies) and the sole member of its own family Cryptocercidae. Species are known as wood roaches or brown-hooded cockroaches. These roaches are subsocial, their young requiring considera ...
'', 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
A nymph ( grc, νύμφη, nýmphē, el, script=Latn, nímfi, label=Modern Greek; , ) in ancient Greek folklore is a minor female nature deity. Different from Greek goddesses, nymphs are generally regarded as personifications of nature, are typ ...
. In 2008
DNA analysis from
16S rRNA 16S rRNA may refer to:
* 16S ribosomal RNA
16 S ribosomal RNA (or 16 S rRNA) is the RNA component of the 30S subunit of a prokaryotic ribosome ( SSU rRNA). It binds to the Shine-Dalgarno sequence and provides most of the SSU structure.
The g ...
sequences supported the position of termites being nested within the evolutionary tree containing the order
Blattodea
Blattodea is an order of insects that contains cockroaches and termites. Formerly, termites were considered a separate order, Isoptera, but genetic and molecular evidence suggests they evolved from within the cockroach lineage, cladistically ...
, 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
Social behavior is behavior among two or more organisms within the same species, and encompasses any behavior in which one member affects the other. This is due to an interaction among those members. Social behavior can be seen as similar to an ...
such as
trophallaxis
Trophallaxis () is the transfer of food or other fluids among members of a community through mouth-to-mouth ( stomodeal) or anus-to-mouth ( proctodeal) feeding. Along with nutrients, trophallaxis can involve the transfer of molecules such as pher ...
and
allogrooming
Social grooming is a behavior in which social animals, including humans, clean or maintain one another's body or appearance. A related term, allogrooming, indicates social grooming between members of the same species. Grooming is a major soci ...
. Termites are thought to be the descendants of the genus ''Cryptocercus''.
Some researchers have suggested a more conservative measure of retaining the termites as the Termitoidae, an
epifamily
In biological classification, taxonomic rank is the relative level of a group of organisms (a taxon) in an ancestral or hereditary hierarchy. A common system consists of species, genus, family, order, class, phylum, kingdom, domain. While olde ...
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
Mantidae is one of the largest families in the order of praying mantises, based on the type species '' Mantis religiosa''; however, most genera are tropical or subtropical. Historically, this was the only family in the order, and many referen ...
, and they are classified in the same superorder (
Dictyoptera
Dictyoptera (from Greek δίκτυον ''diktyon'' "net" and πτερόν ''pteron'' "wing") is an insect superorder that includes two extant orders of polyneopterous insects: the order Blattodea (termites and cockroaches together) and the o ...
).
The oldest unambiguous termite
fossils
A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in ...
date to the early
Cretaceous
The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
, but given the diversity of Cretaceous termites and early fossil records showing mutualism between microorganisms and these insects, they possibly originated earlier in the Jurassic or Triassic. Possible evidence of a Jurassic origin is the assumption that the extinct ''
Fruitafossor
''Fruitafossor'' was a termite-eating mammal endemic to North America during the Late Jurassic epoch (around 150 mya).
The description is based on a complete skeleton of a chipmunk-sized animal.
It was discovered on March 31, 2005, in Fruita, ...
'' consumed termites, judging from its morphological similarity to modern termite-eating mammals. The oldest termite nest discovered is believed to be from the
Upper Cretaceous
The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', the ...
in
West Texas
West Texas is a loosely defined region in the U.S. state of Texas, generally encompassing the arid and semiarid lands west of a line drawn between the cities of Wichita Falls, Abilene, and Del Rio.
No consensus exists on the boundary betwee ...
, 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
Mastotermitidae is a family of termites with one sole living species, ''Mastotermes darwiniensis'' which is found only in northern Australia. The remaining genera of this family are only known from the fossil record.
Fossil record
Numerous foss ...
termites may go back to the
Late Permian
Late may refer to:
* LATE, an acronym which could stand for:
** Limbic-predominant age-related TDP-43 encephalopathy, a proposed form of dementia
** Local-authority trading enterprise, a New Zealand business law
** Local average treatment effect, ...
, 251 million years ago, and fossil wings that have a close resemblance to the wings of ''Mastotermes'' of the Mastotermitidae, the most primitive living termite, have been discovered in the
Permian
The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleoz ...
layers in Kansas.
It is even possible that the first termites emerged during the
Carboniferous
The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...
. The folded wings of the fossil wood roach ''
Pycnoblattina'', arranged in a convex pattern between segments 1a and 2a, resemble those seen in ''Mastotermes'', the only living insect with the same pattern.
Krishna ''et al.'', though, consider that all of the Paleozoic and Triassic insects tentatively classified as termites are in fact unrelated to termites and should be excluded from the Isoptera.
Other studies suggest that the origin of termites is more recent, having diverged from ''Cryptocercus'' sometime during the
Early Cretaceous
The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145 Ma to 100.5 Ma.
Geology
Pro ...
.
![Isoptera](https://upload.wikimedia.org/wikipedia/commons/7/71/Isoptera.jpg)
The primitive
giant northern termite (''Mastotermes darwiniensis'') exhibits numerous cockroach-like characteristics that are not shared with other termites, such as laying its eggs in rafts and having anal lobes on the wings. It has been proposed that the Isoptera and Cryptocercidae be grouped in the clade "
Xylophagodea". Termites are sometimes called "white ants" but the only resemblance to the ants is due to their sociality which is due to
convergent evolution
Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
with termites being the first social insects to evolve a caste system more than 100 million years ago.
Termite genomes are generally relatively large compared to those of other insects; the first fully sequenced termite genome, of ''
Zootermopsis nevadensis'', which was published in the journal
Nature Communications
''Nature Communications'' is a peer-reviewed, open access, scientific journal published by Nature Portfolio since 2010. It is a multidisciplinary journal and it covers the natural sciences, including physics, chemistry, earth sciences, medicine, ...
, consists of roughly 500Mb, while two subsequently published genomes, ''
Macrotermes natalensis
''Macrotermes natalensis'' is a fungus-growing termite species that belongs to the genus ''Macrotermes.'' This species is associated with the '' Termitomyces'' fungal genus. ''M. natalensis'' has domesticated ''Termitomyces'' to produce food for ...
'' and ''
Cryptotermes secundus'', are considerably larger at around 1.3Gb.
External phylogeny
Internal phylogeny
As of 2013, about 3,106 living and fossil termite
species
In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
are recognised, classified in 12 families; reproductive and/or soldier castes are usually required for identification. The infraorder Isoptera is divided into the following clade and family groups, showing the subfamilies in their respective classification:
Basal termite families
: Infraorder Isoptera (= Epifamily Termitoidae)
:: Family
Cratomastotermitidae
''Cratomastotermes'' is an extinct genus of termites in the family Cratomastotermitidae, the sole genus of the family. There is one described species in ''Cratomastotermes'', ''C. wolfschwenningeri''.
References
Termites
Taxa named ...
:: Family
Mastotermitidae
Mastotermitidae is a family of termites with one sole living species, ''Mastotermes darwiniensis'' which is found only in northern Australia. The remaining genera of this family are only known from the fossil record.
Fossil record
Numerous foss ...
: Parvorder
Euisoptera
:: Family
Arceotermitidae
:: Family
Archotermopsidae
Archotermopsidae is a family of termites in the order Blattodea, known as dampwood termites, formerly included within the family Termopsidae. They constitute a small and rather primitive family with five extant genera and 13–20 living species. ...
:: Family
Hodotermitidae
The harvester termites (from Greek ὁδός (hodós), travelling; Latin '' termes'', woodworm) are an ancient, Old World family of termites, the Hodotermitidae. They are distinguished by the serrated inner edge of their mandibles, and their func ...
:: Family
Kalotermitidae
Kalotermitidae is a family of termites, commonly known as drywood termites. Kalotermitidae includes 21 genera and 419 species. The family has a cosmopolitan circumtropical distribution, and is found in functionally arid environments.
Biology
The ...
:: Family
Krishnatermitidae
:: Family
Melqartitermitidae
:: Family
Mylacrotermitidae
:: Family
Stolotermitidae
Stolotermitidae is a family of termites in the order Blattodea
Blattodea is an order of insects that contains cockroaches and termites. Formerly, termites were considered a separate order, Isoptera, but genetic and molecular evidence suggests ...
:: Family
Tanytermitidae
:: Family
Termopsidae
Termopsidae is an extinct family of termites in the order Blattodea. The five extant genera formerly included in Termopsidae ('' Archotermopsis'', '' Hodotermopsis'', '' Porotermes'', '' Stolotermes'', and ''Zootermopsis'') have recently been tre ...
Neoisoptera
The
Neoisoptera, literally meaning "newer termites" (in an evolutionary sense), are a recently coined
nanorder that include families commonly referred-to as "higher termites", although some authorities only apply this term to the largest family
Termitidae
Termitidae is the largest family of termites whose members are commonly known as the higher termites. They are evolutionarily the most specialised termite group, with their highly compartmentalized hindgut lacking the flagellated protozoans comm ...
. The latter characteristically do not have Pseudergate nymphs (many "lower termite" worker nymphs have the capacity to develop into reproductive castes: see
below
Below may refer to:
*Earth
*Ground (disambiguation)
*Soil
*Floor
*Bottom (disambiguation)
Bottom may refer to:
Anatomy and sex
* Bottom (BDSM), the partner in a BDSM who takes the passive, receiving, or obedient role, to that of the top or ...
). Cellulose digestion in "higher termites" has co-evolved with
eukaryotic
Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
gut microbiota and many genera have symbiotic relationships with fungi such as ''
Termitomyces
''Termitomyces'' is a genus of basidiomycete fungi belonging to the family Lyophyllaceae. There are 30-40 species in the genus, all of which are completely dependent on termites to survive. They are the food source for a subfamily of termites, th ...
''; in contrast, "lower termites" typically have
flagellate
A flagellate is a cell or organism with one or more whip-like appendages called flagella. The word ''flagellate'' also describes a particular construction (or level of organization) characteristic of many prokaryotes and eukaryotes and their ...
s and
prokaryote
A prokaryote () is a single-celled organism that lacks a nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Greek πρό (, 'before') and κάρυον (, 'nut' or 'kernel').Campbell, N. "Biology:Concepts & Connec ...
s in their hindguts. Five families are now included here:
:: Family
Archeorhinotermitidae
''Archeorhinotermes'' is an extinct genus of termites in the family Archeorhinotermitidae, and is the sole genus of the family. There is one described species in ''Archeorhinotermes'', ''A. rossi''. It was discovered in Burmese amber.
Refe ...
:: Family
Rhinotermitidae
Rhinotermitidae is a family of termites (Isoptera). 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 such as ''Coptotermes formosanus'', ''Cop ...
:::
Coptotermitinae
Rhinotermitidae is a family of termites (Isoptera). 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 such as '' Coptotermes formosanus'', '' ...
:::
Heterotermitinae
The Heterotermitinae are a termite subfamily in the family Rhinotermitidae
Rhinotermitidae is a family of termites (Isoptera). They feed on wood and can cause extensive damage to buildings or other wooden structures. About 345 species are reco ...
:::
Prorhinoterminae
:::
Psammotermitinae
Rhinotermitidae is a family of termites (Isoptera). 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 such as '' Coptotermes formosanus'', '' ...
:::
Rhinotermitinae
Rhinotermitidae is a family of termites (Isoptera). 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 such as '' Coptotermes formosanus'', '' ...
:::
Termitogetoninae
:: Family
Serritermitidae
Serritermitidae is a family of termites belonging to the infraorder Isoptera in the order Blattodea
Blattodea is an order of insects that contains cockroaches and termites. Formerly, termites were considered a separate order, Isoptera, but ...
:: Family
Stylotermitidae
Stylotermitidae is a family of termites in the order Blattodea
Blattodea is an order of insects that contains cockroaches and termites. Formerly, termites were considered a separate order, Isoptera, but genetic and molecular evidence sugge ...
:: Family
Termitidae
Termitidae is the largest family of termites whose members are commonly known as the higher termites. They are evolutionarily the most specialised termite group, with their highly compartmentalized hindgut lacking the flagellated protozoans comm ...
:::
Apicotermitinae
:::
Cubitermitinae
:::
Foraminitermitinae
:::
Macrotermitinae
The Macrotermitinae, the fungus-growing termites, constitute a subfamily of the family Termitidae that is only found within the Old World tropics.
This subfamily consists of 14 genera and about 350 species and are distinguished by the fact t ...
:::
Nasutitermitinae
The Nasutitermitinae are a subfamily of higher termites that includes more than 80 genera. They are most recognisable by the more highly derived soldier caste which exhibits vestigial mandibles and a protruding fontanellar process on the head ...
:::
Sphaerotermitinae
:::
Syntermitinae
:::
Termitinae
Termitinae is a subfamily of termites; ''Amitermes'' and certain other genera (indicated here with a *) have previously been placed in the Amitermitinae
Amitermitinae is a disputed subfamily of "higher termites" now often merged with the sub ...
Distribution and diversity
Termites are found on all continents except
Antarctica
Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean, it contains the geographic South Pole. Antarctica is the fifth-largest contine ...
. The diversity of termite species is low in
North America
North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
and
Europe
Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...
(10 species known in Europe and 50 in North America), but is high in
South America
South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
, where over 400 species are known.
Of the 3,000 termite species currently classified, 1,000 are found in
Africa
Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
, where mounds are extremely abundant in certain regions. Approximately 1.1 million active termite mounds can be found in the northern
Kruger National Park
Kruger National Park is a South African National Park and one of the largest game reserves in Africa. It covers an area of in the provinces of Limpopo and Mpumalanga in northeastern South Africa, and extends from north to south and from ea ...
alone. In
Asia
Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...
, there are 435 species of termites, which are mainly distributed in
China
China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
. Within China, termite species are restricted to mild
tropical
The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in
the Southern Hemisphere at S. The tropics are also referred to ...
and
subtropical
The subtropical zones or subtropics are geographical zone, geographical and Köppen climate classification, climate zones to the Northern Hemisphere, north and Southern Hemisphere, south of the tropics. Geographically part of the Geographical z ...
habitats south of the Yangtze River.
[ In ]Australia
Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
, all ecological groups of termites (dampwood, drywood, subterranean) are endemic
Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsew ...
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
Biomass is plant-based material used as a fuel for heat or electricity production. It can be in the form of wood, wood residues, energy crops, agricultural residues, and waste from industry, farms, and households. Some people use the terms bi ...
. Termites and ant
Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22 ...
s comprise about 1% of insect species, but represent more than 50% of insect biomass.
Due to their soft cuticles, termites do not inhabit cool or cold habitats. There are three ecological groups of termites: dampwood, drywood and subterranean. Dampwood termites are found only in coniferous forests, and drywood termites are found in hardwood forests; subterranean termites live in widely diverse areas.[ One species in the drywood group is the West Indian drywood termite ''(]Cryptotermes brevis
''Cryptotermes brevis'' is a species of termite in the family Kalotermitidae, commonly known as the West Indian drywood termite or the powderpost termite. It is able to live completely inside timber structures or articles made of wood such as f ...
)'', which is an invasive species in Australia.
Description
Termites are usually small, measuring between in length.[ The largest of all extant termites are the queens of the species '']Macrotermes bellicosus
''Macrotermes bellicosus'' is a species of ''Macrotermes''. It is the largest termite known, with queens measuring about long, workers about and soldiers are slightly larger. ' means "combative" in Latin. The species is a member of a genus indi ...
'', measuring up to over 10 centimetres (4 in) in length. Another giant termite, the extinct ''Gyatermes styriensis
Termites are small insects that live in colonies and have distinct castes (eusocial) and feed on wood or other dead plant matter. Termites comprise the infraorder Isoptera, or alternatively the epifamily Termitoidae, within the order Blattod ...
'', flourished in Austria
Austria, , bar, Östareich officially the Republic of Austria, is a country in the southern part of Central Europe, lying in the Eastern Alps. It is a federation of nine states, one of which is the capital, Vienna, the most populous ...
during the Miocene
The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
and had a wingspan of and a body length of .
Most worker and soldier termites are completely blind as they do not have a pair of eyes. However, some species, such as ''Hodotermes mossambicus
''Hodotermes'' (from Greek ὁδός (hodós), travelling; Latin '' termes'', woodworm) is a genus of African harvester termites in the Hodotermitidae. They range from Palaearctic North Africa, through the East African savannas to the karroid ...
'', have compound eyes
A compound eye is a visual organ found in arthropods such as insects and crustaceans. It may consist of thousands of ommatidia, which are tiny independent photoreception units that consist of a cornea, lens, and photoreceptor cells which distin ...
which they use for orientation and to distinguish sunlight from moonlight. The alate
Alate (Latin ''ālātus'', from ''āla'' (“wing”)) is an adjective and noun used in entomology and botany to refer to something that has wings or winglike structures.
In entomology
In entomology, "alate" usually refers to the winged form o ...
s (winged males and females) have eyes along with lateral ocelli
A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-l ...
. Lateral ocelli, however, are not found in all termites, absent in the families Hodotermitidae
The harvester termites (from Greek ὁδός (hodós), travelling; Latin '' termes'', woodworm) are an ancient, Old World family of termites, the Hodotermitidae. They are distinguished by the serrated inner edge of their mandibles, and their func ...
, Termopsidae
Termopsidae is an extinct family of termites in the order Blattodea. The five extant genera formerly included in Termopsidae ('' Archotermopsis'', '' Hodotermopsis'', '' Porotermes'', '' Stolotermes'', and ''Zootermopsis'') have recently been tre ...
, and Archotermopsidae
Archotermopsidae is a family of termites in the order Blattodea, known as dampwood termites, formerly included within the family Termopsidae. They constitute a small and rather primitive family with five extant genera and 13–20 living species. ...
.[ Like other insects, termites have a small tongue-shaped labrum and a clypeus; the clypeus is divided into a postclypeus and anteclypeus. Termite antennae have a number of functions such as the sensing of touch, taste, odours (including pheromones), heat and vibration. The three basic segments of a termite antenna include a scape, a pedicel (typically shorter than the scape), and the flagellum (all segments beyond the scape and pedicel). The mouth parts contain a ]maxillae
The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
, a labium, and a set of mandibles
In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
. The maxillae and labium have palps
Pedipalps (commonly shortened to palps or palpi) are the second pair of appendages of chelicerates – a group of arthropods including spiders, scorpions, horseshoe crabs, and sea spiders. The pedipalps are lateral to the chelicerae ("jaws") a ...
that help termites sense food and handling.
Consistent with all insects, the anatomy of the termite thorax
The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the cre ...
consists of three segments: the prothorax
The prothorax is the foremost of the three segments in the thorax of an insect, and bears the first pair of legs. Its principal sclerites (exoskeletal plates) are the pronotum (dorsal), the prosternum (ventral), and the propleuron (lateral) on ea ...
, the mesothorax
The mesothorax is the middle of the three segments of the thorax of hexapods, and bears the second pair of legs. Its principal sclerites (exoskeletal plates) are the mesonotum (dorsal), the mesosternum (ventral), and the mesopleuron (lateral) on ...
and the metathorax
The metathorax is the posterior of the three segments in the thorax of an insect, and bears the third pair of legs. Its principal sclerites ( exoskeletal plates) are the metanotum (dorsal), the metasternum (ventral), and the metapleuron (lateral) ...
. Each segment contains a pair of leg
A leg is a weight-bearing and locomotive anatomical structure, usually having a columnar shape. During locomotion, legs function as "extensible struts". The combination of movements at all joints can be modeled as a single, linear element ca ...
s. On alates, the wings are located at the mesothorax and metathorax, which is consistent with all four-winged insects. The mesothorax and metathorax have well-developed exoskeletal plates; the prothorax has smaller plates.
Termites have a ten-segmented abdomen with two plates, the tergites
A ''tergum'' (Latin for "the back"; plural ''terga'', associated adjective tergal) is the dorsal ('upper') portion of an arthropod segment other than the head. The anterior edge is called the 'base' and posterior edge is called the 'apex' or 'mar ...
and the sternites
The sternum (pl. "sterna") is the ventral portion of a segment of an arthropod thorax or abdomen.
In insects, the sterna are usually single, large sclerites, and external. However, they can sometimes be divided in two or more, in which case the ...
. The tenth abdominal segment has a pair of short cerci. There are ten tergites, of which nine are wide and one is elongated. The reproductive organs are similar to those in cockroaches but are more simplified. For example, the intromittent organ
An intromittent organ is any external organ of a male organism that is specialized to deliver sperm during copulation. Intromittent organs are found most often in terrestrial species, as most non-mammalian aquatic species fertilize their eggs e ...
is not present in male alates, and the sperm is either immotile or aflagellate. However, Mastotermitidae termites have multiflagellate sperm with limited motility
Motility is the ability of an organism to move independently, using metabolic energy.
Definitions
Motility, the ability of an organism to move independently, using metabolic energy, can be contrasted with sessility, the state of organisms th ...
. The genitals in females are also simplified. Unlike in other termites, Mastotermitidae females have an ovipositor
The ovipositor is a tube-like organ used by some animals, especially insects, for the laying of eggs. In insects, an ovipositor consists of a maximum of three pairs of appendages. The details and morphology of the ovipositor vary, but typical ...
, a feature strikingly similar to that in female cockroaches.
The non-reproductive castes of termites are wingless and rely exclusively on their six legs for locomotion. The alates fly only for a brief amount of time, so they also rely on their legs. The appearance of the legs is similar in each caste, but the soldiers have larger and heavier legs. The structure of the legs is consistent with other insects: the parts of a leg include a coxa, trochanter
A trochanter is a tubercle of the femur near its joint with the hip bone. In humans and most mammals, the trochanters serve as important muscle attachment sites. Humans are known to have three trochanters, though the anatomic "normal" includes ...
, femur
The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates with ...
, tibia
The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
and the tarsus. The number of tibial spurs on an individual's leg varies. Some species of termite have an arolium, located between the claws
A claw is a curved, pointed appendage found at the end of a toe or finger in most amniotes (mammals, reptiles, birds). Some invertebrates such as beetles and spiders have somewhat similar fine, hooked structures at the end of the leg or tarsus ...
, which is present in species that climb on smooth surfaces but is absent in most termites.
Unlike in ants, the hind-wings and fore-wings are of equal length.[ Most of the time, the alates are poor flyers; their technique is to launch themselves in the air and fly in a random direction. Studies show that in comparison to larger termites, smaller termites cannot fly long distances. When a termite is in flight, its wings remain at a right angle, and when the termite is at rest, its wings remain parallel to the body.
]
Caste system
Worker termites undertake the most labour within the colony, being responsible for foraging, food storage, and brood and nest maintenance. Workers are tasked with the digestion of cellulose
Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell wall ...
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 the transfer of food or other fluids among members of a community through mouth-to-mouth ( stomodeal) or anus-to-mouth ( proctodeal) feeding. Along with nutrients, trophallaxis can involve the transfer of molecules such as pher ...
. Trophallaxis is an effective nutritional tactic to convert and recycle nitrogenous components. It frees the parents from feeding all but the first generation of offspring, allowing for the group to grow much larger and ensuring that the necessary gut symbionts are transferred from one generation to another. Some termite species may rely on nymphs to perform work without differentiating as a separate caste. Workers may be male or female and are usually sterile, especially in termites that have a nest site that is separate from their foraging site. Sterile workers are sometimes termed as true workers while those that are fertile, as in the wood-nesting Archotermopsidae, are termed as false workers.
The soldier caste has anatomical and behavioural specialisations, and their sole purpose is to defend the colony. Many soldiers have large heads with highly modified powerful jaws so enlarged they cannot feed themselves. Instead, like juveniles, they are fed by workers.[ ]Fontanelle
A fontanelle (or fontanel) (colloquially, soft spot) is an anatomical feature of the infant human skull comprising soft membranous gaps ( sutures) between the cranial bones that make up the calvaria of a fetus or an infant. Fontanelles allow f ...
s, simple holes in the forehead that exude defensive secretions, are a feature of the family Rhinotermitidae. Many species are readily identified using the characteristics of the soldiers' larger and darker head and large mandibles.[ Among certain termites, soldiers may use their globular (phragmotic) heads to block their narrow tunnels. Different sorts of soldiers include minor and major soldiers, and nasutes, which have a horn-like nozzle frontal projection (a nasus).][ These unique soldiers are able to spray noxious, sticky secretions containing ]diterpene
Diterpenes are a class of chemical compounds composed of four isoprene units, often with the molecular formula C20H32. They are biosynthesized by plants, animals and fungi via the HMG-CoA reductase pathway, with geranylgeranyl pyrophosphate being ...
s at their enemies. Nitrogen fixation
Nitrogen fixation is a chemical process by which molecular nitrogen (), with a strong triple covalent bond, in the air is converted into ammonia () or related nitrogenous compounds, typically in soil or aquatic systems but also in industry. Atmo ...
plays an important role in nasute nutrition. Soldiers are usually sterile but some species of Archotermopsidae are known to have neotenic forms with soldier-like heads while also having sexual organs.
The primary reproductive caste of a colony consists of the adult form (imago) female and male individuals, colloquially known as the queen and king. The queen of the colony is responsible for egg production for the colony. Unlike in ants, the king mates with her for life.[ In some species, the abdomen of the queen swells up dramatically to increase ]fecundity
Fecundity is defined in two ways; in human demography, it is the potential for reproduction of a recorded population as opposed to a sole organism, while in population biology, it is considered similar to fertility, the natural capability to pr ...
, 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
Nuptial flight is an important phase in the reproduction of most ant, termite, and some bee species. It is also observed in some fly species, such as ''Rhamphomyia longicauda''.
During the flight, virgin queens mate with males and then land to ...
begins. These swarms attract a wide variety of predators.[
]
Life cycle
Termites are often compared with the social
Social organisms, including human(s), live collectively in interacting populations. This interaction is considered social whether they are aware of it or not, and whether the exchange is voluntary or not.
Etymology
The word "social" derives from ...
Hymenoptera (ants and various species of bees and wasps), but their differing evolutionary origins result in major differences in life cycle. In the eusocial Hymenoptera, the workers are exclusively female. Males (drones) are haploid and develop from unfertilised eggs, while females (both workers and the queen) are diploid and develop from fertilised eggs. In contrast, worker termites, which constitute the majority in a colony, are diploid
Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
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
An egg is an organic vessel grown by an animal to carry a possibly fertilized egg cell (a zygote) and to incubate from it an embryo within the egg until the embryo has become an animal fetus that can survive on its own, at which point the a ...
, but is different from that of a bee or ant in that it goes through a developmental process called incomplete metamorphosis
Hemimetabolism or hemimetaboly, also called incomplete metamorphosis and paurometabolism,McGavin, George C. ''Essential Entomology: An Order-by-Order Introduction''. Oxford: Oxford University Press, 2001. pp. 20. is the mode of development of certa ...
, with egg, nymph and adult stages. Nymphs resemble small adults, and go through a series of moults as they grow. In some species, eggs go through four moulting stages and nymphs go through three. Nymphs first moult into workers, and then some workers go through further moulting and become soldiers or alates; workers become alates only by moulting into alate nymphs.
The development of nymphs into adults can take months; the time period depends on food availability, temperature, and the general population of the colony. Since nymphs are unable to feed themselves, workers must feed them, but workers also take part in the social life of the colony and have certain other tasks to accomplish such as foraging, building or maintaining the nest or tending to the queen. Pheromones regulate the caste system in termite colonies, preventing all but a very few of the termites from becoming fertile queens.
Queens of the eusocial
Eusociality (from Greek εὖ ''eu'' "good" and social), the highest level of organization of sociality, is defined by the following characteristics: cooperative brood care (including care of offspring from other individuals), overlapping generat ...
termite ''Reticulitermes
''Reticulitermes'' is a termite genus in the family Rhinotermitidae. They are found in most temperate regions on Earth including much of Asia and the Middle East, Western Europe, and all of North America.
Caste descriptions
''Reticulitermes'' ...
speratus'' are capable of a long lifespan without sacrificing fecundity
Fecundity is defined in two ways; in human demography, it is the potential for reproduction of a recorded population as opposed to a sole organism, while in population biology, it is considered similar to fertility, the natural capability to pr ...
. 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
Catalase is a common enzyme found in nearly all living organisms exposed to oxygen (such as bacteria, plants, and animals) which catalyzes the decomposition of hydrogen peroxide to water and oxygen. It is a very important enzyme in protecting t ...
, an enzyme that protects against oxidative stress
Oxidative stress reflects an imbalance between the systemic manifestation of reactive oxygen species and a biological system's ability to readily Detoxification, detoxify the reactive intermediates or to repair the resulting damage. Disturbances ...
.
Reproduction
Termite alates only leave the colony when a nuptial flight
Nuptial flight is an important phase in the reproduction of most ant, termite, and some bee species. It is also observed in some fly species, such as ''Rhamphomyia longicauda''.
During the flight, virgin queens mate with males and then land to ...
takes place. Alate males and females pair up together and then land in search of a suitable place for a colony. A termite king and queen do not mate until they find such a spot. When they do, they excavate a chamber big enough for both, close up the entrance and proceed to mate.[ After mating, the pair never go outside and spend the rest of their lives in the nest. Nuptial flight time varies in each species. For example, alates in certain species emerge during the day in summer while others emerge during the winter.] The nuptial flight may also begin at dusk, when the alates swarm around areas with many lights. The time when nuptial flight begins depends on the environmental conditions, the time of day, moisture, wind speed and precipitation.[ The number of termites in a colony also varies, with the larger species typically having 100–1,000 individuals. However, some termite colonies, including those with many individuals, can number in the millions.][
The queen only lays 10–20 eggs in the very early stages of the colony, but lays as many as 1,000 a day when the colony is several years old.][ At maturity, a primary queen has a great capacity to lay eggs. In some species, the mature queen has a greatly distended abdomen and may produce 40,000 eggs a day. The two mature ovaries may have some 2,000 ]ovariole
An ovariole is a tubular component of the insect ovary, and the basic unit of egg production. Each ovariole is composed of a germarium (the germline stem cell niche) at the anterior tip, a set of developing oocytes contained within follicles, and ...
s each. The abdomen increases the queen's body length to several times more than before mating and reduces her ability to move freely; attendant workers provide assistance.
The king grows only slightly larger after initial mating and continues to mate with the queen for life (a termite queen can live between 30 to 50 years); this is very different from ant colonies, in which a queen mates once with the males and stores the gametes for life, as the male ants die shortly after mating.[ If a queen is absent, a termite king produces pheromones which encourage the development of replacement termite queens. As the queen and king are monogamous, sperm competition does not occur.
Termites going through incomplete metamorphosis on the path to becoming alates form a subcaste in certain species of termite, functioning as potential supplementary reproductives. These supplementary reproductives only mature into primary reproductives upon the death of a king or queen, or when the primary reproductives are separated from the colony.][ Supplementaries have the ability to replace a dead primary reproductive, and there may also be more than a single supplementary within a colony.][ Some queens have the ability to switch from sexual reproduction to ]asexual reproduction
Asexual reproduction is a type of reproduction that does not involve the fusion of gametes or change in the number of chromosomes. The offspring that arise by asexual reproduction from either unicellular or multicellular organisms inherit the fu ...
. Studies show that while termite queens mate with the king to produce colony workers, the queens reproduce their replacements (neotenic
Neoteny (), also called juvenilization,Montagu, A. (1989). Growing Young. Bergin & Garvey: CT. is the delaying or slowing of the physiological, or somatic, development of an organism, typically an animal. Neoteny is found in modern humans compared ...
queens) parthenogenetically.
The neotropical termite ''Embiratermes neotenicus'' and several other related species produce colonies that contain a primary king accompanied by a primary queen or by up to 200 neotenic
Neoteny (), also called juvenilization,Montagu, A. (1989). Growing Young. Bergin & Garvey: CT. is the delaying or slowing of the physiological, or somatic, development of an organism, typically an animal. Neoteny is found in modern humans compared ...
queens that had originated through thelytokous parthenogenesis
Thelytoky (from the Greek ''thēlys'' "female" and ''tokos'' "birth") is a type of parthenogenesis in which females are produced from unfertilized eggs, as for example in aphids. Thelytokous parthenogenesis is rare among animals and reported in a ...
of a founding primary queen. The form of parthenogenesis
Parthenogenesis (; from the Greek grc, παρθένος, translit=parthénos, lit=virgin, label=none + grc, γένεσις, translit=génesis, lit=creation, label=none) is a natural form of asexual reproduction in which growth and development ...
likely employed maintains heterozygosity
Zygosity (the noun, zygote, is from the Greek "yoked," from "yoke") () is the degree to which both copies of a chromosome or gene have the same genetic sequence. In other words, it is the degree of similarity of the alleles in an organism.
Mo ...
in the passage of the genome
In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ge ...
from mother to daughter, thus avoiding inbreeding depression
Inbreeding depression is the reduced biological fitness which has the potential to result from inbreeding (the breeding of related individuals). Biological fitness refers to an organism's ability to survive and perpetuate its genetic material. In ...
.
Behaviour and ecology
Diet
Termites are detritivore
Detritivores (also known as detrivores, detritophages, detritus feeders, or detritus eaters) are heterotrophs that obtain nutrients by consuming detritus (decomposing plant and animal parts as well as feces). There are many kinds of invertebrates, ...
s, consuming dead plants at any level of decomposition. They also play a vital role in the ecosystem by recycling waste material such as dead wood, faeces and plants. Many species eat cellulose
Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell wall ...
, having a specialised midgut that breaks down the fibre. Termites are considered to be a major source (11%) of atmospheric methane
Atmospheric methane is the methane present in Earth's atmosphere. Atmospheric methane concentrations are of interest because it is one of the most potent greenhouse gases in Earth's atmosphere. Atmospheric methane is rising.
The 20-year glob ...
, one of the prime greenhouse gas
A greenhouse gas (GHG or GhG) is a gas that Absorption (electromagnetic radiation), absorbs and Emission (electromagnetic radiation), emits radiant energy within the thermal infrared range, causing the greenhouse effect. The primary greenhouse ...
es, produced from the breakdown of cellulose. Termites rely primarily upon symbiotic protozoa (metamonad
The metamonads are microscopic eukaryotic organisms, a large group of flagellate amitochondriate Loukozoa. Their composition is not entirely settled, but they include the retortamonads, diplomonads, and possibly the parabasalids and oxymonads a ...
s) and other microbe
A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in olde ...
s such as flagellate
A flagellate is a cell or organism with one or more whip-like appendages called flagella. The word ''flagellate'' also describes a particular construction (or level of organization) characteristic of many prokaryotes and eukaryotes and their ...
protist
A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the exc ...
s in their guts to digest the cellulose for them, allowing them to absorb the end products for their own use. The microbial ecosystem present in the termite gut contains many species found nowhere else on Earth. Termites hatch without these symbionts present in their guts, and develop them after fed a culture from other termites. Gut protozoa, such as ''Trichonympha'', in turn, rely on symbiotic bacteria
Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among ...
embedded on their surfaces to produce some of the necessary digestive enzyme
Digestive enzymes are a group of enzymes that break down polymeric macromolecules into their smaller building blocks, in order to facilitate their absorption into the cells of the body. Digestive enzymes are found in the digestive tracts of anim ...
s. Most higher termites, especially in the family Termitidae, can produce their own cellulase
Cellulase (EC 3.2.1.4; systematic name 4-β-D-glucan 4-glucanohydrolase) is any of several enzymes produced chiefly by fungi, bacteria, and protozoans that catalyze cellulolysis, the decomposition of cellulose and of some related polysaccharide ...
enzymes, but they rely primarily upon the bacteria. The flagellates have been lost in Termitidae. Researches have found species of spirochetes
A spirochaete () or spirochete is a member of the phylum Spirochaetota (), (synonym Spirochaetes) which contains distinctive diderm (double-membrane) gram-negative bacteria, most of which have long, helically coiled (corkscrew-shaped or ...
living in termite guts capable of fixing atmospheric nitrogen to a form usable by the insect. Scientists' understanding of the relationship between the termite digestive tract and the microbial endosymbionts is still rudimentary; what is true in all termite species, however, is that the workers feed the other members of the colony with substances derived from the digestion of plant material, either from the mouth
In animal anatomy, the mouth, also known as the oral cavity, or in Latin cavum oris, is the opening through which many animals take in food and issue vocal sounds. It is also the cavity lying at the upper end of the alimentary canal, bounded on ...
or anus.[Geetha Iyer ''Scroll.in'' (Mar 09, 2017) Why Indians worship the mound of the much-hated termite]
"he soldier termites
He or HE may refer to:
Language
* He (pronoun), an English pronoun
* He (kana), the romanization of the Japanese kana へ
* He (letter), the fifth letter of many Semitic alphabets
* He (Cyrillic), a letter of the Cyrillic script called ''He'' ...
and the reproductive castes obtain their nutrients from the workers through oral or anal trophallaxis." Judging from closely related bacterial species, it is strongly presumed that the termites' and cockroach's gut microbiota
Gut microbiota, gut microbiome, or gut flora, are the microorganisms, including bacteria, archaea, fungi, and viruses that live in the digestive tracts of animals. The gastrointestinal metagenome is the aggregate of all the genomes of the gut m ...
derives from their dictyoptera
Dictyoptera (from Greek δίκτυον ''diktyon'' "net" and πτερόν ''pteron'' "wing") is an insect superorder that includes two extant orders of polyneopterous insects: the order Blattodea (termites and cockroaches together) and the o ...
n ancestors.
Certain species such as '' Gnathamitermes tubiformans'' have seasonal food habits. For example, they may preferentially consume Red three-awn (''Aristida longiseta
''Aristida'' is a very nearly cosmopolitan genus of plants in the grass family. ''Aristida'' is distinguished by having three awns (bristles) on each lemma of each floret. The genus includes about 300 species found worldwide, often in arid warm ...
'') during the summer, Buffalograss ('' Buchloe dactyloides'') from May to August, and blue grama ''Bouteloua gracilis
''Bouteloua gracilis'', the blue grama, is a long-lived, warm-season (C4 carbon fixation, C4) Perennial plant, perennial grass, native to North America.
It is most commonly found from Alberta, Canada, east to Manitoba and south across the Rocky ...
'' during spring, summer and autumn. Colonies of ''G. tubiformans'' consume less food in spring than they do during autumn when their feeding activity is high.
Various woods differ in their susceptibility to termite attack; the differences are attributed to such factors as moisture content, hardness, and resin and lignin content. In one study, the drywood termite ''Cryptotermes brevis'' strongly preferred poplar and maple
''Acer'' () is a genus of trees and shrubs commonly known as maples. The genus is placed in the family Sapindaceae.Stevens, P. F. (2001 onwards). Angiosperm Phylogeny Website. Version 9, June 2008 nd more or less continuously updated since http ...
woods to other woods that were generally rejected by the termite colony. These preferences may in part have represented conditioned or learned behaviour.
Some species of termite practice fungiculture
Fungiculture is the cultivation of fungi such as mushrooms. Cultivating fungi can yield foods (which include mostly mushrooms), medicine, construction materials and other products. A ''mushroom farm'' is involved in the business of growing fun ...
. They maintain a "garden" of specialised fungi of genus ''Termitomyces
''Termitomyces'' is a genus of basidiomycete fungi belonging to the family Lyophyllaceae. There are 30-40 species in the genus, all of which are completely dependent on termites to survive. They are the food source for a subfamily of termites, th ...
'', which are nourished by the excrement of the insects. When the fungi are eaten, their spores pass undamaged through the intestines of the termites to complete the cycle by germinating in the fresh faecal pellets. Molecular evidence suggests that the family Macrotermitinae
The Macrotermitinae, the fungus-growing termites, constitute a subfamily of the family Termitidae that is only found within the Old World tropics.
This subfamily consists of 14 genera and about 350 species and are distinguished by the fact t ...
developed agriculture about 31 million years ago. It is assumed that more than 90 percent of dry wood in the semiarid savannah ecosystems of Africa and Asia are reprocessed by these termites. Originally living in the rainforest, fungus farming allowed them to colonise the African savannah and other new environments, eventually expanding into Asia.
Depending on their feeding habits, termites are placed into two groups: the lower termites and higher termites. The lower termites predominately feed on wood. As wood is difficult to digest, termites prefer to consume fungus-infected wood because it is easier to digest and the fungi are high in protein. Meanwhile, the higher termites consume a wide variety of materials, including faeces, humus
In classical soil science, humus is the dark organic matter in soil that is formed by the decomposition of plant and animal matter. It is a kind of soil organic matter. It is rich in nutrients and retains moisture in the soil. Humus is the Lati ...
, grass, leaves and roots. The gut of the lower termites contains many species of bacteria along with protozoa
Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Histo ...
and '' Holomastigotoides'', while the higher termites only have a few species of bacteria with no protozoa.
Predators
Termites are consumed by a wide variety of predators
Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...
. One termite species alone, ''Hodotermes mossambicus
''Hodotermes'' (from Greek ὁδός (hodós), travelling; Latin '' termes'', woodworm) is a genus of African harvester termites in the Hodotermitidae. They range from Palaearctic North Africa, through the East African savannas to the karroid ...
'', was found in the stomach contents of 65 bird
Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
s and 19 mammal
Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
s. Arthropod
Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...
s such as ants
Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22,00 ...
, centipede
Centipedes (from New Latin , "hundred", and Latin , " foot") are predatory arthropods belonging to the class Chilopoda (Ancient Greek , ''kheilos'', lip, and New Latin suffix , "foot", describing the forcipules) of the subphylum Myriapoda, an ...
s, cockroach
Cockroaches (or roaches) are a paraphyletic group of insects belonging to Blattodea, containing all members of the group except termites. About 30 cockroach species out of 4,600 are associated with human habitats. Some species are well-known as ...
es, crickets
Crickets are orthopteran insects which are related to bush crickets, and, more distantly, to grasshoppers. In older literature, such as Imms,Imms AD, rev. Richards OW & Davies RG (1970) ''A General Textbook of Entomology'' 9th Ed. Methuen 8 ...
, dragonflies
A dragonfly is a flying insect belonging to the infraorder Anisoptera below the order Odonata. About 3,000 extant species of true dragonfly are known. Most are tropical, with fewer species in temperate regions. Loss of wetland habitat threa ...
, scorpion
Scorpions are predatory arachnids of the order Scorpiones. They have eight legs, and are easily recognized by a pair of grasping pincers and a narrow, segmented tail, often carried in a characteristic forward curve over the back and always end ...
s and spider
Spiders ( order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...
s, reptile
Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates (lizards and snakes) and rhynchocephalians ( ...
s such as lizard
Lizards are a widespread group of squamate reptiles, with over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The group is paraphyletic since it excludes the snakes and Amphisbaenia alt ...
s, and amphibian
Amphibians are tetrapod, four-limbed and ectothermic vertebrates of the Class (biology), class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terres ...
s such as frog
A frog is any member of a diverse and largely Carnivore, carnivorous group of short-bodied, tailless amphibians composing the order (biology), order Anura (ανοὐρά, literally ''without tail'' in Ancient Greek). The oldest fossil "proto-f ...
s and toad
Toad is a common name for certain frogs, especially of the family Bufonidae, that are characterized by dry, leathery skin, short legs, and large bumps covering the parotoid glands.
A distinction between frogs and toads is not made in scientif ...
s consume termites, with two spider
Spiders ( order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...
s in the family Ammoxenidae
Ground spiders comprise Gnaphosidae, the seventh largest spider family with over 2,000 described species in over 100 genera distributed worldwide. There are 105 species known to central Europe, and common genera include ''Gnaphosa'', ''Drassodes' ...
being specialist termite predators. Other predators include aardvark
The aardvark ( ; ''Orycteropus afer'') is a medium-sized, burrowing, nocturnal mammal native to Africa. It is the only living species of the order Tubulidentata, although other prehistoric species and genera of Tubulidentata are known. Unlike ...
s, aardwolves
The aardwolf (''Proteles cristata'') is an insectivorous species of hyena, native to East and Southern Africa. Its name means "earth-wolf" in Afrikaans and Dutch. It is also called maanhaar-jackal (Afrikaans for " mane-jackal"), termite-eatin ...
, anteater
Anteater is a common name for the four extant mammal species of the suborder Vermilingua (meaning "worm tongue") commonly known for eating ants and termites. The individual species have other names in English and other languages. Together with ...
s, bat
Bats are mammals of the order Chiroptera.''cheir'', "hand" and πτερόν''pteron'', "wing". With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most ...
s, bear
Bears are carnivoran mammals of the family Ursidae. They are classified as caniforms, or doglike carnivorans. Although only eight species of bears are extant, they are widespread, appearing in a wide variety of habitats throughout the Nor ...
s, bilbies, many bird
Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...
s, echidnas
Echidnas (), sometimes known as spiny anteaters, are quill-covered monotremes (egg-laying mammals) belonging to the family Tachyglossidae . The four extant species of echidnas and the platypus are the only living mammals that lay eggs and the ...
, fox
Foxes are small to medium-sized, omnivorous mammals belonging to several genera of the family Canidae. They have a flattened skull, upright, triangular ears, a pointed, slightly upturned snout, and a long bushy tail (or ''brush'').
Twelve sp ...
es, galago
Galagos , also known as bush babies, or ''nagapies'' (meaning "night monkeys" in Afrikaans), are small nocturnal primates native to continental, sub-Sahara Africa, and make up the family Galagidae (also sometimes called Galagonidae). They ar ...
s, numbat
The numbat (''Myrmecobius fasciatus''), also known as the noombat or walpurti, is an insectivorous marsupial. It is diurnal and its diet consists almost exclusively of termites.
The species was once widespread across southern Australia, but ...
s, mice
A mouse ( : mice) is a small rodent. Characteristically, mice are known to have a pointed snout, small rounded ears, a body-length scaly tail, and a high breeding rate. The best known mouse species is the common house mouse (''Mus musculus' ...
and pangolin
Pangolins, sometimes known as scaly anteaters, are mammals of the order Pholidota (, from Ancient Greek ϕολιδωτός – "clad in scales"). The one extant family, the Manidae, has three genera: '' Manis'', '' Phataginus'', and '' Smut ...
s. The aardwolf
The aardwolf (''Proteles cristata'') is an insectivorous species of hyena, native to East and Southern Africa. Its name means "earth-wolf" in Afrikaans and Dutch. It is also called maanhaar-jackal (Afrikaans for " mane-jackal"), termite-eatin ...
is an insectivorous
A robber fly eating a hoverfly
An insectivore is a carnivorous animal or plant that eats insects. An alternative term is entomophage, which can also refer to the human practice of eating insects.
The first vertebrate insectivores were ...
mammal
Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
that primarily feeds on termites; it locates its food by sound and also by detecting the scent secreted by the soldiers; a single aardwolf
The aardwolf (''Proteles cristata'') is an insectivorous species of hyena, native to East and Southern Africa. Its name means "earth-wolf" in Afrikaans and Dutch. It is also called maanhaar-jackal (Afrikaans for " mane-jackal"), termite-eatin ...
is capable of consuming thousands of termites in a single night by using its long, sticky tongue. Sloth bears
The sloth bear (''Melursus ursinus'') is a myrmecophagous bear species native to the Indian subcontinent. It feeds on fruits, ants and termites. It is listed as Vulnerable species, vulnerable on the IUCN Red List, mainly because of habitat loss ...
break open mounds to consume the nestmates, while chimpanzees
The chimpanzee (''Pan troglodytes''), also known as simply the chimp, is a species of great ape native to the forest and savannah of tropical Africa. It has four confirmed subspecies and a fifth proposed subspecies. When its close relative the ...
have developed tools to "fish" termites from their nest. Wear pattern analysis of bone tools used by the early hominin
The Hominini form a taxonomic tribe of the subfamily Homininae ("hominines"). Hominini includes the extant genera ''Homo'' (humans) and '' Pan'' (chimpanzees and bonobos) and in standard usage excludes the genus ''Gorilla'' (gorillas).
The t ...
''Paranthropus robustus
''Paranthropus robustus'' is a species of robustness (morphology), robust australopithecine from the Early Pleistocene, Early and possibly Middle Pleistocene, Middle Pleistocene of the Cradle of Humankind, South Africa, about 2.27 to 0.87 (or, mo ...
'' suggests that they used these tools to dig into termite mounds.
Among all predators, ants are the greatest enemy to termites. Some ant genera are specialist predators of termites. For example, ''Megaponera
''Megaponera analis'' is the sole species of the genus ''Megaponera''. They are a strictly termite-eating (termitophagous) ponerine ant species widely distributed in Sub-Saharan Africa and most commonly known for their column-like raiding format ...
'' is a strictly termite-eating (termitophagous) genus that perform raiding activities, some lasting several hours. '' Paltothyreus tarsatus'' is another termite-raiding species, with each individual stacking as many termites as possible in its mandible
In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
s before returning home, all the while recruiting additional nestmates to the raiding site through chemical trails.[ The Malaysian basicerotine ants '' Eurhopalothrix heliscata'' uses a different strategy of termite hunting by pressing themselves into tight spaces, as they hunt through rotting wood housing termite colonies. Once inside, the ants seize their prey by using their short but sharp mandibles.][ '' Tetramorium uelense'' is a specialised predator species that feeds on small termites. A scout recruits 10–30 workers to an area where termites are present, killing them by immobilising them with their stinger. '']Centromyrmex
''Centromyrmex'' is a pantropical, though mainly Afrotropical, genus of ants in the subfamily Ponerinae. This ponerine ant was recorded for the first time in French Guiana and the most northerly point of recording was in Costa Rica. The specime ...
'' and ''Iridomyrmex
''Iridomyrmex'' is a genus of ants called rainbow ants (referring to their blue-green iridescent sheen) first described by Austrian entomologist Gustav Mayr in 1862. He placed the genus in the subfamily Dolichoderinae of the family Formicida ...
'' colonies sometimes nest in termite mound
Mound-building termites are a group of termite species that live in mounds. These termites live in Africa, Australia and South America. The mounds sometimes have a diameter of . Most of the mounds are in well-drained areas. Termite mounds usually ...
s, and so the termites are preyed on by these ants. No evidence for any kind of relationship (other than a predatory one) is known. Other ants, including ''Acanthostichus
''Acanthostichus'' is a predatory and predominantly subterranean genus of ant in the subfamily Dorylinae. They are found in the New World, from the southern United States to Uruguay, Paraguay and northern Argentina
Argentina (), offici ...
'', '' Camponotus'', ''Crematogaster
''Crematogaster'' is an ecologically diverse genus of ants found worldwide, which are characterised by a distinctive heart-shaped gaster (abdomen), which gives them one of their common names, the Saint Valentine ant. Members of this genus are ...
'', ''Cylindromyrmex
''Cylindromyrmex'' is a Neotropical genus of ants in the subfamily Dorylinae.
Distribution and habitat
The genus is known from the New World, from Central America to Brazil, Bolivia and the Galápagos Islands, and currently contains 10 extant ...
'', ''Leptogenys
''Leptogenys'' is a genus of ants in the subfamily Ponerinae. ''Leptogenys'' is the most diverse ponerine ant genus in the world; it is widespread throughout tropical and subtropical regions and there are over 260 extant species described. Mos ...
'', ''Odontomachus
''Odontomachus'' is a genus of ants commonly called trap-jaw ants found in the tropics and subtropics throughout the world.
Overview
Commonly known as trap-jaw ants, species in ''Odontomachus'' have a pair of large, straight mandibles capable ...
'', ''Ophthalmopone
''Ophthalmopone'' is a ponerine genus of ant
Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous ...
'', ''Pachycondyla
''Pachycondyla'' is a ponerine genus of ants found in the Neotropics.
Distribution
''Pachycondyla'' is currently distributed from southern United States to northern Argentina, but some fossil species (e.g. '' P. eocenica'' and '' P. lutzi'') a ...
'', ''Rhytidoponera
''Rhytidoponera'' is a large genus of ants in the subfamily Ectatomminae. The genus is known from Australia and Melanesia, with New Caledonia as the most eastern limit.
Some ''Rhytidoponera'' species have both winged alate
Alate (Latin ''āl ...
'', '' Solenopsis'' and ''Wasmannia
''Wasmannia'' is a genus of ants. ''Wasmannia auropunctata'' is known as the ''electric ant''[Dorylus
''Dorylus'', also known as driver ants, safari ants, or siafu, is a large genus of army ants found primarily in central and east Africa, although the range also extends to southern Africa and tropical Asia. The term siafu is a loanword from Swah ...](_blank)
'' ants rarely consume termites.
Ants are not the only invertebrates that perform raids. Many sphecoid wasps and several species including '' Polybia'' and '' Angiopolybia'' are known to raid termite mounds during the termites' nuptial flight.
Parasites, pathogens and viruses
Termites are less likely to be attacked by parasites than bees, wasps and ants, as they are usually well protected in their mounds. Nevertheless, termites are infected by a variety of parasites. Some of these include dipteran flies, '' Pyemotes'' mites, and a large number of nematode
The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-Parasitism, parasitic nematodes also known as eelworms. They are a diverse animal phylum inhab ...
parasites. Most nematode parasites are in the order Rhabditida
Rhabditida is an order of free-living, zooparasitic, and phytoparasitic microbivorous nematodes living in soil.
The Cephalobidae, Panagrolaimidae, Steinernematidae, and Strongyloididae seem to be closer to the Tylenchia, regardless of whet ...
; others are in the genus ''Mermis
''Mermis'' is a genus of nematodes
The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-parasitic nematodes also known as eelworms. They are a d ...
'', '' Diplogaster aerivora'' and '' Harteria gallinarum''. Under imminent threat of an attack by parasites, a colony may migrate to a new location. Certain fungal pathogens such as ''Aspergillus nomius
''Aspergillus nomius'' is a species of fungus in the genus ''Aspergillus''. It is from the ''Flavi'' section. The species was first described in 1987.Kurtzman, C.P.; Horn, B.W.; Hesseltine, C.W. 1987. Aspergillus nomius, a new aflatoxin-producin ...
'' and ''Metarhizium anisopliae
''Metarhizium robertsii'' formerly known as ''M. anisopliae'', and even earlier as ''Entomophthora anisopliae'' (basionym) is a fungus that grows naturally in soils throughout the world and causes disease in various insects by acting as a paras ...
'' are, however, major threats to a termite colony as they are not host-specific and may infect large portions of the colony; transmission usually occurs via direct physical contact. ''M. anisopliae'' is known to weaken the termite immune system. Infection with ''A. nomius'' only occurs when a colony is under great stress. Over 34 fungal species are known to live as parasites on the exoskeleton of termites, with many being host-specific and only causing indirect harm to their host.
Termites are infected by viruses including Entomopoxvirinae
''Entomopoxvirinae'' is a subfamily of viruses, in the family ''Poxviridae''. Insects, human, vertebrates, and arthropods serve as natural hosts. There are currently 31 species in this subfamily, divided among 4 genera with one species unassign ...
and the Nuclear Polyhedrosis Virus.
Locomotion and foraging
Because the worker and soldier castes lack wings and thus never fly, and the reproductives use their wings for just a brief amount of time, termites predominantly rely upon their legs to move about.
Foraging behaviour depends on the type of termite. For example, certain species feed on the wood structures they inhabit, and others harvest food that is near the nest. Most workers are rarely found out in the open, and do not forage unprotected; they rely on sheeting and runways to protect them from predators. Subterranean termites construct tunnels and galleries to look for food, and workers who manage to find food sources recruit additional nestmates by depositing a phagostimulant pheromone that attracts workers. Foraging workers use semiochemicals to communicate with each other,[ and workers who begin to forage outside of their nest release trail pheromones from their sternal glands. In one species, '' Nasutitermes costalis'', there are three phases in a foraging expedition: first, soldiers scout an area. When they find a food source, they communicate to other soldiers and a small force of workers starts to emerge. In the second phase, workers appear in large numbers at the site. The third phase is marked by a decrease in the number of soldiers present and an increase in the number of workers. Isolated termite workers may engage in ]Lévy flight
A Lévy flight is a random walk in which the step-lengths have a Lévy distribution, a probability distribution that is heavy-tailed. When defined as a walk in a space of dimension greater than one, the steps made are in isotropic random direct ...
behaviour as an optimised strategy for finding their nestmates or foraging for food.
Competition
Competition between two colonies always results in agonistic behaviour
Agonistic behaviour is any social behaviour related to fighting. The term has broader meaning than aggressive behaviour because it includes threats, displays, retreats, placation, and conciliation. The term "agonistic behaviour" was first implemen ...
towards each other, resulting in fights. These fights can cause mortality on both sides and, in some cases, the gain or loss of territory. "Cemetery pits" may be present, where the bodies of dead termites are buried.
Studies show that when termites encounter each other in foraging areas, some of the termites deliberately block passages to prevent other termites from entering.[ Dead termites from other colonies found in exploratory tunnels leads to the isolation of the area and thus the need to construct new tunnels. Conflict between two competitors does not always occur. For example, though they might block each other's passages, colonies of ''Macrotermes bellicosus'' and ''Macrotermes subhyalinus'' are not always aggressive towards each other. Suicide cramming is known in '' Coptotermes formosanus''. Since ''C. formosanus'' colonies may get into physical conflict, some termites squeeze tightly into foraging tunnels and die, successfully blocking the tunnel and ending all agonistic activities.
Among the reproductive caste, neotenic queens may compete with each other to become the dominant queen when there are no primary reproductives. This struggle among the queens leads to the elimination of all but a single queen, which, with the king, takes over the colony.
Ants and termites may compete with each other for nesting space. In particular, ants that prey on termites usually have a negative impact on arboreal nesting species.]
Communication
Most termites are blind, so communication primarily occurs through chemical, mechanical and pheromonal cues. These methods of communication are used in a variety of activities, including foraging, locating reproductives, construction of nests, recognition of nestmates, nuptial flight, locating and fighting enemies, and defending the nests.[ The most common way of communicating is through antennation.][ A number of pheromones are known, including contact pheromones (which are transmitted when workers are engaged in trophallaxis or grooming) and ]alarm
An alarm device is a mechanism that gives an audible, visual or other kind of alarm signal to alert someone to a problem or condition that requires urgent attention.
Alphabetical musical instruments
Etymology
The word ''alarm'' comes from th ...
, trail
A trail, also known as a path or track, is an unpaved lane or small road usually passing through a natural area. In the United Kingdom and the Republic of Ireland, a path or footpath is the preferred term for a pedestrian or hiking trail. Th ...
and sex pheromone
Sex pheromones are pheromones released by an organism to attract an individual of the same species, encourage them to mate with them, or perform some other function closely related with sexual reproduction.
Sex pheromones specifically focus on ind ...
s. The alarm pheromone and other defensive chemicals are secreted from the frontal gland. Trail pheromones are secreted from the sternal gland, and sex pheromones derive from two glandular sources: the sternal and tergal glands.[ When termites go out to look for food, they forage in columns along the ground through vegetation. A trail can be identified by the faecal deposits or runways that are covered by objects. Workers leave pheromones on these trails, which are detected by other nestmates through olfactory receptors.][ Termites can also communicate through mechanical cues, vibrations, and physical contact.][ These signals are frequently used for alarm communication or for evaluating a food source.][
When termites construct their nests, they use predominantly indirect communication. No single termite would be in charge of any particular construction project. Individual termites react rather than think, but at a group level, they exhibit a sort of collective cognition. Specific structures or other objects such as pellets of soil or pillars cause termites to start building. The termite adds these objects onto existing structures, and such behaviour encourages building behaviour in other workers. The result is a self-organised process whereby the information that directs termite activity results from changes in the environment rather than from direct contact among individuals.][
Termites can distinguish nestmates and non-nestmates through chemical communication and gut symbionts: chemicals consisting of hydrocarbons released from the cuticle allow the recognition of alien termite species. Each colony has its own distinct odour. This odour is a result of genetic and environmental factors such as the termites' diet and the composition of the bacteria within the termites' intestines.
]
Defence
Termites rely on alarm communication to defend a colony.[ Alarm pheromones can be released when the nest has been breached or is being attacked by enemies or potential pathogens. Termites always avoid nestmates infected with '']Metarhizium anisopliae
''Metarhizium robertsii'' formerly known as ''M. anisopliae'', and even earlier as ''Entomophthora anisopliae'' (basionym) is a fungus that grows naturally in soils throughout the world and causes disease in various insects by acting as a paras ...
'' spores, through vibrational signals released by infected nestmates. Other methods of defence include intense jerking and secretion of fluids from the frontal gland and defecating faeces containing alarm pheromones.[
In some species, some soldiers block tunnels to prevent their enemies from entering the nest, and they may deliberately rupture themselves as an act of defence. In cases where the intrusion is coming from a breach that is larger than the soldier's head, soldiers form a ]phalanx
The phalanx ( grc, φάλαγξ; plural phalanxes or phalanges, , ) was a rectangular mass military formation, usually composed entirely of heavy infantry armed with spears, pikes, sarissas, or similar pole weapons. The term is particularly use ...
-like formation around the breach and bite at intruders.[ If an invasion carried out by '' Megaponera analis'' is successful, an entire colony may be destroyed, although this scenario is rare.]
To termites, any breach of their tunnels or nests is a cause for alarm. When termites detect a potential breach, the soldiers usually bang their heads, apparently to attract other soldiers for defence and to recruit additional workers to repair any breach.[ Additionally, an alarmed termite bumps into other termites which causes them to be alarmed and to leave pheromone trails to the disturbed area, which is also a way to recruit extra workers.][
]
The pantropical subfamily Nasutitermitinae
The Nasutitermitinae are a subfamily of higher termites that includes more than 80 genera. They are most recognisable by the more highly derived soldier caste which exhibits vestigial mandibles and a protruding fontanellar process on the head ...
has a specialised caste of soldiers, known as nasutes, that have the ability to exude noxious liquids through a horn-like frontal projection that they use for defence. Nasutes have lost their mandibles through the course of evolution and must be fed by workers.[ A wide variety of ]monoterpene
Monoterpenes are a class of terpenes that consist of two isoprene units and have the molecular formula C10H16. Monoterpenes may be linear (acyclic) or contain rings (monocyclic and bicyclic). Modified terpenes, such as those containing oxygen funct ...
hydrocarbon solvent
A solvent (s) (from the Latin '' solvō'', "loosen, untie, solve") is a substance that dissolves a solute, resulting in a solution. A solvent is usually a liquid but can also be a solid, a gas, or a supercritical fluid. Water is a solvent for ...
s have been identified in the liquids that nasutes secrete. Similarly, Formosan subterranean termite
The Formosan termite (''Coptotermes formosanus'') is a species of termite local to southern China and introduced to Taiwan (formerly known as Formosa, where it gets its name), Japan, South Africa, Sri Lanka, Hawaii, and the continental United Sta ...
s have been known to secrete naphthalene
Naphthalene is an organic compound with formula . It is the simplest polycyclic aromatic hydrocarbon, and is a white crystalline solid with a characteristic odor that is detectable at concentrations as low as 0.08 ppm by mass. As an aromati ...
to protect their nests.
Soldiers of the species ''Globitermes sulphureus
''Globitermes sulphureus'' (suicidal attack ants) 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 Cambodia, Thailand, and Peninsular Malaysia. They live in ...
'' commit suicide by autothysis – rupturing a large gland just beneath the surface of their cuticles. The thick, yellow fluid in the gland becomes very sticky on contact with the air, entangling ants or other insects that are trying to invade the nest. Another termite, '' Neocapriterme taracua'', also engages in suicidal defence. Workers physically unable to use their mandibles while in a fight form a pouch full of chemicals, then deliberately rupture themselves, releasing toxic chemicals that paralyse and kill their enemies. The soldiers of the neotropical
The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropical terrestrial ecoregions of the Americas and the entire South American temperate zone.
Definition
In bioge ...
termite family Serritermitidae
Serritermitidae is a family of termites belonging to the infraorder Isoptera in the order Blattodea
Blattodea is an order of insects that contains cockroaches and termites. Formerly, termites were considered a separate order, Isoptera, but ...
have a defence strategy which involves front gland autothysis, with the body rupturing between the head and abdomen. When soldiers guarding nest entrances are attacked by intruders, they engage in autothysis, creating a block that denies entry to any attacker.
Workers use several different strategies to deal with their dead, including burying, cannibalism, and avoiding a corpse altogether. To avoid pathogens
In biology, a pathogen ( el, πάθος, "suffering", "passion" and , "producer of") in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a germ ...
, termites occasionally engage in necrophoresis
Necrophoresis is a sanitation behavior found in social insects – such as ants, bees, wasps, and termites – in which they carry away the dead bodies of members of their colony from the nest or hive area. The term was introduced in 1958 by E.O. ...
, in which a nestmate carries away a corpse from the colony to dispose of it elsewhere. Which strategy is used depends on the nature of the corpse a worker is dealing with (i.e. the age of the carcass).[
]
Relationship with other organisms
A species of fungus
A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from th ...
is known to mimic termite eggs, successfully avoiding its natural predators. These small brown balls, known as "termite balls", rarely kill the eggs, and in some cases the workers tend to them. This fungus mimics these eggs by producing a cellulose-digesting enzyme known as glucosidases
Glucosidases are the glycoside hydrolase enzymes categorized under the EC number 3.2.1.
Function
Alpha-glucosidases are enzymes involved in breaking down complex carbohydrates such as starch and glycogen into their monomers.
They catalyze the ...
. A unique mimicking behaviour exists between various species of '' Trichopsenius'' beetles and certain termite species within ''Reticulitermes
''Reticulitermes'' is a termite genus in the family Rhinotermitidae. They are found in most temperate regions on Earth including much of Asia and the Middle East, Western Europe, and all of North America.
Caste descriptions
''Reticulitermes'' ...
''. The beetles share the same cuticle
A cuticle (), or cuticula, is any of a variety of tough but flexible, non-mineral outer coverings of an organism, or parts of an organism, that provide protection. Various types of "cuticle" are non- homologous, differing in their origin, structu ...
hydrocarbon
In organic chemistry, a hydrocarbon is an organic compound consisting entirely of hydrogen and carbon. Hydrocarbons are examples of group 14 hydrides. Hydrocarbons are generally colourless and hydrophobic, and their odors are usually weak or ex ...
s as the termites and even biosynthesize them. This chemical mimicry allows the beetles to integrate themselves within the termite colonies. The developed appendages
An appendage (or outgrowth) is an external body part, or natural prolongation, that protrudes from an organism's body.
In arthropods, an appendage refers to any of the homologous body parts that may extend from a body segment, including anten ...
on the physogastric abdomen of '' Austrospirachtha mimetes'' allows the beetle to mimic a termite worker.
Some species of ant are known to capture termites to use as a fresh food source later on, rather than killing them. For example, '' Formica nigra'' captures termites, and those that try to escape are immediately seized and driven underground. Certain species of ants in the subfamily Ponerinae
Ponerinae is a subfamily of ants in the Poneromorph subfamilies group, with about 1,600 species in 47 extant genera, including ''Dinoponera gigantea'' - one of the world's largest species of ant. Mated workers have replaced the queen as the fun ...
conduct these raids although other ant species go in alone to steal the eggs or nymphs.[ Ants such as ''Megaponera analis'' attack the outside of mounds and ]Dorylinae
Dorylinae is an ant subfamily, with distributions in both the Old World and New World. Brady ''et al.'' (2014) synonymized the previous dorylomorph subfamilies (Aenictinae, Aenictogitoninae, Cerapachyinae, Ecitoninae, and Leptanilloidinae) unde ...
ants attack underground.[ Despite this, some termites and ants can coexist peacefully. Some species of termite, including '']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, Pa ...
'', form associations with certain ant species to keep away predatory ant species. The earliest known association between '' Azteca'' ants and ''Nasutitermes'' termites date back to the Oligocene to Miocene period.
54 species of ants are known to inhabit ''Nasutitermes'' mounds, both occupied and abandoned ones. One reason many ants live in ''Nasutitermes'' mounds is due to the termites' frequent occurrence in their geographical range; another is to protect themselves from floods.[ ''Iridomyrmex'' also inhabits termite mounds although no evidence for any kind of relationship (other than a predatory one) is known.][ In rare cases, certain species of termites live inside active ant colonies. Some invertebrate organisms such as beetles, caterpillars, flies and millipedes are termitophiles and dwell inside termite colonies (they are unable to survive independently).][ As a result, certain beetles and flies have evolved with their hosts. They have developed a gland that secrete a substance that attracts the workers by licking them. Mounds may also provide shelter and warmth to birds, lizards, snakes and scorpions.][
Termites are known to carry pollen and regularly visit flowers,] so are regarded as potential pollinators for a number of flowering plants. One flower in particular, '' Rhizanthella gardneri'', is regularly pollinated by foraging workers, and it is perhaps the only Orchidaceae
Orchids are plants that belong to the family Orchidaceae (), a diverse and widespread group of flowering plants with blooms that are often colourful and fragrant.
Along with the Asteraceae, they are one of the two largest families of flowering ...
flower in the world to be pollinated by termites.[
Many plants have developed effective defences against termites. However, seedlings are vulnerable to termite attacks and need additional protection, as their defence mechanisms only develop when they have passed the seedling stage. Defence is typically achieved by secreting antifeedant chemicals into the woody cell walls.][ This reduces the ability of termites to efficiently digest the ]cellulose
Cellulose is an organic compound with the formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of β(1→4) linked D-glucose units. Cellulose is an important structural component of the primary cell wall ...
. A commercial product, "Blockaid", has been developed in Australia that uses a range of plant extracts to create a paint-on nontoxic termite barrier Termite barriers are materials that have been specifically designed to prevent subterranean termites from gaining access to a structure. Physical termite barriers are free of pesticides and physically block termite foraging activity. Development of ...
for buildings. An extract of a species of Australian figwort, '' Eremophila'', has been shown to repel termites; tests have shown that termites are strongly repelled by the toxic material to the extent that they will starve rather than consume the food. When kept close to the extract, they become disoriented and eventually die.[
]
Relationship with the environment
Termite populations can be substantially impacted by environmental changes including those caused by human intervention. A Brazilian study investigated the termite assemblages of three sites of Caatinga
Caatinga (, ) is a type of semi-arid tropical vegetation, and an ecoregion characterized by this vegetation in interior northeastern Brazil. The name "Caatinga" is a Tupi word meaning "white forest" or "white vegetation" (''caa'' = forest, v ...
under different levels of anthropogenic disturbance in the semi-arid region of northeastern Brazil
Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
were sampled using 65 x 2 m transects. A total of 26 species of termites were present in the three sites, and 196 encounters were recorded in the transects. The termite assemblages were considerably different among sites, with a conspicuous reduction in both diversity and abundance with increased disturbance, related to the reduction of tree density and soil cover, and with the intensity of trampling by cattle and goats. The wood-feeders were the most severely affected feeding group.
Nests
A termite nest can be considered as being composed of two parts, the inanimate and the animate. The animate is all of the termites living inside the colony, and the inanimate part is the structure itself, which is constructed by the termites. Nests can be broadly separated into three main categories: subterranean (completely below ground), epigeal (protruding above the soil surface), and arboreal (built above ground, but always connected to the ground via shelter tubes).[ Epigeal nests (mounds) protrude from the earth with ground contact and are made out of earth and mud. A nest has many functions such as providing a protected living space and providing shelter against predators. Most termites construct underground colonies rather than multifunctional nests and mounds.] Primitive termites of today nest in wooden structures such as logs, stumps and the dead parts of trees, as did termites millions of years ago.
To build their nests, termites primarily use faeces, which have many desirable properties as a construction material. Other building materials include partly digested plant material, used in carton nests (arboreal nests built from faecal elements and wood), and soil, used in subterranean nest and mound construction. Not all nests are visible, as many nests in tropical forests are located underground.[ Species in the subfamily Apicotermitinae are good examples of subterranean nest builders, as they only dwell inside tunnels. Other termites live in wood, and tunnels are constructed as they feed on the wood. Nests and mounds protect the termites' soft bodies against desiccation, light, pathogens and parasites, as well as providing a fortification against predators. Nests made out of carton are particularly weak, and so the inhabitants use counter-attack strategies against invading predators.
Arboreal carton nests of ]mangrove swamp
Mangrove forests, also called mangrove swamps, mangrove thickets or mangals, are productive wetlands that occur in coastal intertidal zones. Mangrove forests grow mainly at tropical and subtropical latitudes because mangroves cannot withstand fre ...
-dwelling ''Nasutitermes'' are enriched in lignin
Lignin is a class of complex organic polymers that form key structural materials in the support tissues of most plants. Lignins are particularly important in the formation of cell walls, especially in wood and bark, because they lend rigidity ...
and depleted in cellulose and xylans. This change is caused by bacterial decay in the gut of the termites: they use their faeces as a carton building material. Arboreal termites nests can account for as much as 2% of above ground carbon storage in Puerto Rica
Puerto, a Spanish word meaning ''seaport'', may refer to:
Places
*El Puerto de Santa María, Andalusia, Spain
*Puerto, a seaport town in Cagayan de Oro City, Philippines
*Puerto Colombia, Colombia
*Puerto Cumarebo, Venezuela
*Puerto Galera, Orient ...
n mangrove swamps. These ''Nasutitermes'' nests are mainly composed of partially biodegraded wood material from the stems and branches of mangrove trees, namely, ''Rhizophora mangle
''Rhizophora mangle'', the red mangrove, is distributed in Estuary, estuarine ecosystems throughout the tropics. Its Vivipary, viviparous "seeds", in actuality called propagules, become fully mature plants before dropping off the parent tree. Th ...
'' (red mangrove), ''Avicennia germinans
''Avicennia germinans'', the black mangrove, is a shrub or small tree growing up to 12 meters (39 feet) in the acanthus family, Acanthaceae. It grows in tropical and subtropical regions of the Americas, on both the Atlantic and Pacific Coasts, ...
'' (black mangrove) and ''Laguncularia racemosa
''Laguncularia racemosa'', the white mangrove, is a species of flowering plant in the leadwood tree family, Combretaceae. It is native to the coasts of western Africa from Senegal to Cameroon, the Atlantic Coast of the Americas from Bermuda and ...
'' (white mangrove).
Some species build complex nests called polycalic nests; this habitat is called polycalism. Polycalic species of termites form multiple nests, or calies, connected by subterranean chambers.[ The termite genera '' Apicotermes'' and '']Trinervitermes
''Trinervitermes'' is a termite genus belonging to family Termitidae. Members are native to the Old World. They inhabit grasslands and store grass in their nests or mounds, just below the ground surface. Their grass-collecting activities are main ...
'' are known to have polycalic species. Polycalic nests appear to be less frequent in mound-building species although polycalic arboreal nests have been observed in a few species of ''Nasutitermes''.[
]
Mounds
Nests are considered mounds if they protrude from the earth's surface. A mound provides termites the same protection as a nest but is stronger. Mounds located in areas with torrential and continuous rainfall are at risk of mound erosion due to their clay-rich construction. Those made from carton can provide protection from the rain, and in fact can withstand high precipitation. Certain areas in mounds are used as strong points in case of a breach. For example, '' Cubitermes'' colonies build narrow tunnels used as strong points, as the diameter of the tunnels is small enough for soldiers to block. A highly protected chamber, known as the "queen's cell", houses the queen and king and is used as a last line of defence.
Species in the genus ''Macrotermes'' arguably build the most complex structures in the insect world, constructing enormous mounds. These mounds are among the largest in the world, reaching a height of 8 to 9 metres (26 to 29 feet), and consist of chimneys, pinnacles and ridges.[ Another termite species, '']Amitermes meridionalis
''Amitermes meridionalis'', commonly known as the magnetic termite or compass termite, is a species of eusocial insect in the family Termitidae. It is endemic to northern Australia and the common names derive from the fact that the wedge-shaped ...
'', can build nests 3 to 4 metres (9 to 13 feet) high and 2.5 metres (8 feet) wide. The tallest mound ever recorded was 12.8 metres (42 ft) long found in the Democratic Republic of the Congo.
The sculptured mounds sometimes have elaborate and distinctive forms, such as those of the compass termite (''Amitermes meridionalis'' and ''A. laurensis''), which builds tall, wedge-shaped mounds with the long axis oriented approximately north–south, which gives them their common name. This orientation has been experimentally shown to assist thermoregulation
Thermoregulation is the ability of an organism to keep its body temperature within certain boundaries, even when the surrounding temperature is very different. A thermoconforming organism, by contrast, simply adopts the surrounding temperature ...
. The north–south orientation causes the internal temperature of a mound to increase rapidly during the morning while avoiding overheating from the midday sun. The temperature then remains at a plateau for the rest of the day until the evening.
File:RayNorris termite cathedral mounds.jpg, alt=. These termite mounds have a base shaped like the base of a tree, about two meters wide and a meter high. From this base, rounded chimneys from half a meter to a meter in diameter rise to a total height of about four or five meters. The chimneys are fused together with ridges between, and terminate in rounded pinnacles at the top., Cathedral mounds in the Northern Territory
The Northern Territory (commonly abbreviated as NT; formally the Northern Territory of Australia) is an states and territories of Australia, Australian territory in the central and central northern regions of Australia. The Northern Territory ...
, Australia
Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
File:Termite Magnetic DSC03613.jpg, alt=. Hundreds of compass termite mounds are visible in this photo of a field in northern Australia. The chisel-shaped mounds range from several centimeters to several meters in height., Mounds of "compass" or "magnetic" termites (''Amitermes'') oriented north–south, thereby avoiding mid-day heat
File:Termitenhuegel.jpg, alt=. This termite mound is about three meters in height and four meters across. The mound chimneys are about a meter in diameter and fuse together to form a rounded top., Termite mound in Queensland
)
, nickname = Sunshine State
, image_map = Queensland in Australia.svg
, map_caption = Location of Queensland in Australia
, subdivision_type = Country
, subdivision_name = Australia
, established_title = Before federation
, established_ ...
, Australia
File:Termites in a mound.jpg, alt=. The photographer has broken off a piece of a mound to show the mound's interior. Dozens of tunnels have been exposed, and hundreds of soldiers have emerged to guard the breech in the wall., Termites in a mound, Analamazoatra Reserve
Analamazaotra National Park is a national park of Madagascar. The park is in the eastern portion of Madagascar's Central Highlands. The neighbouring Analamazaotra Forest Station is a local reforestation effort. It adjoins Andasibe-Mantadia Nati ...
, Madagascar
Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Africa ...
File:Termitenhügel Namibia.jpg, Termite mound in Namibia
Namibia (, ), officially the Republic of Namibia, is a country in Southern Africa. Its western border is the Atlantic Ocean. It shares land borders with Zambia and Angola to the north, Botswana to the east and South Africa to the south and ea ...
Shelter tubes
Termites construct shelter tubes, also known as earthen tubes or mud tubes, that start from the ground. These shelter tubes can be found on walls and other structures. Constructed by termites during the night, a time of higher humidity, these tubes provide protection to termites from potential predators, especially ants. Shelter tubes also provide high humidity and darkness and allow workers to collect food sources that cannot be accessed in any other way.[ These passageways are made from soil and faeces and are normally brown in colour. The size of these shelter tubes depends on the number of food sources that are available. They range from less than 1 cm to several cm in width, but may be dozens of metres in length.][
]
Relationship with humans
As pests
Owing to their wood-eating habits, many termite species can do significant damage to unprotected buildings and other wooden structures.[ Termites play an important role as decomposers of wood and vegetative material, and the conflict with humans occurs where structures and landscapes containing structural wood components, cellulose derived structural materials and ornamental vegetation provide termites with a reliable source of food and moisture. Their habit of remaining concealed often results in their presence being undetected until the timbers are severely damaged, with only a thin exterior layer of wood remaining, which protects them from the environment. Of the 3,106 species known, only 183 species cause damage; 83 species cause significant damage to wooden structures.] In North America, 18 subterranean species are pests; in Australia, 16 species have an economic impact; in the Indian subcontinent 26 species are considered pests, and in tropical Africa, 24. In Central America and the West Indies, there are 17 pest species.[ Among the termite genera, ''Coptotermes'' has the highest number of pest species of any genus, with 28 species known to cause damage.][ Less than 10% of drywood termites are pests, but they infect wooden structures and furniture in tropical, subtropical and other regions. Dampwood termites only attack lumber material exposed to rainfall or soil.][
Drywood termites thrive in warm climates, and human activities can enable them to invade homes since they can be transported through contaminated goods, containers and ships.][ Colonies of termites have been seen thriving in warm buildings located in cold regions. Some termites are considered invasive species. ''Cryptotermes brevis'', the most widely introduced invasive termite species in the world, has been introduced to all the islands in the West Indies and to Australia.][
]
In addition to causing damage to buildings, termites can also damage food crops. Termites may attack trees whose resistance to damage is low but generally ignore fast-growing plants. Most attacks occur at harvest time; crops and trees are attacked during the dry season.[
The damage caused by termites costs the southwestern United States approximately $1.5 billion each year in wood structure damage, but the true cost of damage worldwide cannot be determined.][ Drywood termites are responsible for a large proportion of the damage caused by termites. The goal of termite control is to keep structures and susceptible ornamental plants free from termites.; Structures may be homes or business, or elements such as wooden fence posts and telephone poles. Regular and thorough inspections by a trained professional may be necessary to detect termite activity in the absence of more obvious signs like termite swarmers or alates inside or adjacent to a structure. Termite monitors made of wood or cellulose adjacent to a structure may also provide indication of termite foraging activity where it will be in conflict with humans. Termites can be controlled by application of ]Bordeaux mixture
Bordeaux mixture (also called ''Bordo Mix'') is a mixture of copper(II) sulphate (CuSO4) and quicklime ( Ca O) used as a fungicide. It is used in vineyards, fruit-farms and gardens to prevent infestations of downy mildew, powdery mildew and other ...
or other substances that contain copper
Copper is a chemical element with the symbol Cu (from la, cuprum) and atomic number 29. It is a soft, malleable, and ductile metal with very high thermal and electrical conductivity. A freshly exposed surface of pure copper has a pinkis ...
such as chromated copper arsenate Chromated copper arsenate (CCA) is a wood preservative containing compounds of chromium, copper, and arsenic, in various proportions. It is used to impregnate timber and other wood products, especially those intended for outdoor use, in order to pr ...
. In the United states, application of a soil termiticide with the active ingredient Fipronil
Fipronil is a broad-spectrum insecticide that belongs to the phenylpyrazole chemical family. Fipronil disrupts the insect central nervous system by blocking the ligand-gated ion channel of the GABAA receptor and glutamate-gated chloride (GluCl ...
, such as Termidor SC or Taurus SC, by a licensed professional, is a common remedy approved by the Environmental Protection Agency for economically significant subterranean termites. A growing demand for alternative, green, and "more natural" extermination methods has increased demand for mechanical and biological control methods such as Orange Oil
Orange oil is an essential oil produced by cells within the rind of an orange fruit (''Citrus sinensis'' fruit). In contrast to most essential oils, it is extracted as a by-product of orange juice production by centrifugation, producing a co ...
.
To better control the population of termites, various methods have been developed to track termite movements. One early method involved distributing termite bait laced with immunoglobulin G
Immunoglobulin G (Ig G) is a type of antibody. Representing approximately 75% of serum antibodies in humans, IgG is the most common type of antibody found in blood circulation. IgG molecules are created and released by plasma B cells. Each IgG a ...
(IgG) marker proteins from rabbits or chickens. Termites collected from the field could be tested for the rabbit-IgG markers using a rabbit-IgG-specific assay
An assay is an investigative (analytic) procedure in laboratory medicine, mining, pharmacology, environmental biology and molecular biology for qualitatively assessing or quantitatively measuring the presence, amount, or functional activity of a ...
. More recently developed, less expensive alternatives include tracking the termites using egg white, cow milk, or soy milk proteins, which can be sprayed on termites in the field. Termites bearing these proteins can be traced using a protein-specific ELISA
The enzyme-linked immunosorbent assay (ELISA) (, ) is a commonly used analytical biochemistry assay, first described by Eva Engvall and Peter Perlmann in 1971. The assay uses a solid-phase type of enzyme immunoassay (EIA) to detect the presence ...
test.[
In 1994, termites, of the species ]Reticulitermes
''Reticulitermes'' is a termite genus in the family Rhinotermitidae. They are found in most temperate regions on Earth including much of Asia and the Middle East, Western Europe, and all of North America.
Caste descriptions
''Reticulitermes'' ...
grassei, were identified in two bungalows in Saunton
Saunton is a village located approximately two miles from Braunton on the North Devon coast in the South West of England.
Several kilometres long, the village borders Braunton Burrows, the heart of North Devon's Biosphere Reserve, the first ...
, Devon
Devon ( , historically known as Devonshire , ) is a ceremonial and non-metropolitan county in South West England. The most populous settlement in Devon is the city of Plymouth, followed by Devon's county town, the city of Exeter. Devon is ...
. Anecdotal evidence suggests the infestation could date back 70 years before the official identification. There are reports that gardeners had seen white ants and that a greenhouse had had to be replaced in the past. The Saunton infestation was the first and only colony ever recorded in the UK. In 1998 Termite Eradication Programme was set-up, with the intention of containing and eradicating the colony. The TEP was managed by the Ministry of Housing, Communities & Local Government (now the Department for Levelling Up, Housing and Communities
The Department for Levelling Up, Housing and Communities (DLUHC), formerly the Ministry for Housing, Communities and Local Government (MHCLG), is a department of His Majesty's Government responsible for housing, communities, local government ...
.) The TEP used 'insect growth regulators' to prevent the termites from reaching maturity and reproducing. In 2021 the UK’s Termite Eradication Programme announced the eradication of the colony, the first time a country has eradicated termites.
As food
43 termite species are used as food by humans or are fed to livestock. These insects are particularly important in impoverished countries where malnutrition is common, as the protein
Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, respo ...
from termites can help improve the human diet. Termites are consumed in many regions globally, but this practice has only become popular in developed nations in recent years.[
Termites are consumed by people in many different cultures around the world. In many parts of Africa, the ]alate
Alate (Latin ''ālātus'', from ''āla'' (“wing”)) is an adjective and noun used in entomology and botany to refer to something that has wings or winglike structures.
In entomology
In entomology, "alate" usually refers to the winged form o ...
s are an important factor in the diets of native populations. Groups have different ways of collecting or cultivating insects; sometimes collecting soldiers from several species. Though harder to acquire, queens are regarded as a delicacy. Termite alates are high in nutrition with adequate levels of fat
In nutrition science, nutrition, biology, and chemistry, fat usually means any ester of fatty acids, or a mixture of such chemical compound, compounds, most commonly those that occur in living beings or in food.
The term often refers spec ...
and protein. They are regarded as pleasant in taste, having a nut-like flavour after they are cooked.[
Alates are collected when the rainy season begins. During a nuptial flight, they are typically seen around lights to which they are attracted, and so nets are set up on lamps and captured alates are later collected. The wings are removed through a technique that is similar to ]winnowing
Winnowing is a process by which chaff is separated from grain. It can also be used to remove pests from stored grain. Winnowing usually follows threshing in grain preparation. In its simplest form, it involves throwing the mixture into the ...
. The best result comes when they are lightly roasted on a hot plate or fried until crisp. Oil
An oil is any nonpolar chemical substance that is composed primarily of hydrocarbons and is hydrophobic (does not mix with water) & lipophilic (mixes with other oils). Oils are usually flammable and surface active. Most oils are unsaturated ...
is not required as their bodies usually contain sufficient amounts of oil. Termites are typically eaten when livestock is lean and tribal crops have not yet developed or produced any food, or if food stocks from a previous growing season are limited.[
In addition to Africa, termites are consumed in local or tribal areas in Asia and North and South America. In Australia, ]Indigenous Australians
Indigenous Australians or Australian First Nations are people with familial heritage from, and membership in, the ethnic groups that lived in Australia before British colonisation. They consist of two distinct groups: the Aboriginal peoples ...
are aware that termites are edible but do not consume them even in times of scarcity; there are few explanations as to why.[ Termite mounds are the main sources of soil consumption (]geophagy
Geophagia (), also known as geophagy (), is the intentional practice of eating earth or soil-like substances such as clay, chalk, or termite mounds. It is a behavioural adaptation that occurs in many non-human animals and has been documented in ...
) in many countries including Kenya
)
, national_anthem = "Ee Mungu Nguvu Yetu"()
, image_map =
, map_caption =
, image_map2 =
, capital = Nairobi
, coordinates =
, largest_city = Nairobi
, ...
, Tanzania
Tanzania (; ), officially the United Republic of Tanzania ( sw, Jamhuri ya Muungano wa Tanzania), is a country in East Africa within the African Great Lakes region. It borders Uganda to the north; Kenya to the northeast; Comoro Islands and ...
, Zambia
Zambia (), officially the Republic of Zambia, is a landlocked country at the crossroads of Central Africa, Central, Southern Africa, Southern and East Africa, although it is typically referred to as being in Southern Africa at its most cent ...
, Zimbabwe
Zimbabwe (), officially the Republic of Zimbabwe, is a landlocked country located in Southeast Africa, between the Zambezi and Limpopo Rivers, bordered by South Africa to the south, Botswana to the south-west, Zambia to the north, and Mozam ...
and South Africa
South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring countri ...
. Researchers have suggested that termites are suitable candidates for human consumption and space agriculture, as they are high in protein and can be used to convert inedible waste to consumable products for humans.
In agriculture
Termites can be major agricultural pests, particularly in East Africa and North Asia, where crop losses can be severe (3–100% in crop loss in Africa). Counterbalancing this is the greatly improved water infiltration where termite tunnels in the soil allow rainwater to soak in deeply, which helps reduce runoff and consequent soil erosion through bioturbation
Bioturbation is defined as the reworking of soils and sediments by animals or plants. It includes burrowing, ingestion, and defecation of sediment grains. Bioturbating activities have a profound effect on the environment and are thought to be a pr ...
. In South America, cultivated plants such as eucalyptus, upland rice and sugarcane
Sugarcane or sugar cane is a species of (often hybrid) tall, Perennial plant, perennial grass (in the genus ''Saccharum'', tribe Andropogoneae) that is used for sugar Sugar industry, production. The plants are 2–6 m (6–20 ft) tall with ...
can be severely damaged by termite infestations, with attacks on leaves, roots and woody tissue. Termites can also attack other plants, including cassava
''Manihot esculenta'', common name, commonly called cassava (), manioc, or yuca (among numerous regional names), is a woody shrub of the spurge family, Euphorbiaceae, native to South America. Although a perennial plant, cassava is extensively ...
, coffee
Coffee is a drink prepared from roasted coffee beans. Darkly colored, bitter, and slightly acidic, coffee has a stimulant, stimulating effect on humans, primarily due to its caffeine content. It is the most popular hot drink in the world.
S ...
, cotton
Cotton is a soft, fluffy staple fiber that grows in a boll, or protective case, around the seeds of the cotton plants of the genus ''Gossypium'' in the mallow family Malvaceae. The fiber is almost pure cellulose, and can contain minor perce ...
, fruit trees, maize
Maize ( ; ''Zea mays'' subsp. ''mays'', from es, maíz after tnq, mahiz), also known as corn (North American and Australian English), is a cereal grain first domesticated by indigenous peoples in southern Mexico about 10,000 years ago. Th ...
, peanut
The peanut (''Arachis hypogaea''), also known as the groundnut, goober (US), pindar (US) or monkey nut (UK), is a legume crop grown mainly for its edible Seed, seeds. It is widely grown in the tropics and subtropics, important to both small ...
s, soybeans and vegetables.[ Mounds can disrupt farming activities, making it difficult for farmers to operate farming machinery; however, despite farmers' dislike of the mounds, it is often the case that no net loss of production occurs.][ Termites can be beneficial to agriculture, such as by boosting ]crop yield
In agriculture, the yield is a measurement of the amount of a crop grown, or product such as wool, meat or milk produced, per unit area of land. The seed ratio is another way of calculating yields.
Innovations, such as the use of fertilizer, the c ...
s and enriching the soil. Termites and ants can re-colonise untilled land that contains crop stubble, which colonies use for nourishment when they establish their nests. The presence of nests in fields enables larger amounts of rainwater to soak into the ground and increases the amount of nitrogen in the soil, both essential for the growth of crops.
In science and technology
The termite gut has inspired various research efforts aimed at replacing fossil fuels
A fossil fuel is a hydrocarbon-containing material formed naturally in the Earth's crust from the remains of dead plants and animals that is extracted and burned as a fuel. The main fossil fuels are coal, oil, and natural gas. Fossil fuels ...
with cleaner, renewable energy sources.[ Termites are efficient ]bioreactor
A bioreactor refers to any manufactured device or system that supports a biologically active environment. In one case, a bioreactor is a vessel in which a chemical reaction, chemical process is carried out which involves organisms or biochemistry, ...
s, capable of producing two litres of hydrogen
Hydrogen is the chemical element with the symbol H and atomic number 1. Hydrogen is the lightest element. At standard conditions hydrogen is a gas of diatomic molecules having the formula . It is colorless, odorless, tasteless, non-toxic, an ...
from a single sheet of paper. Approximately 200 species of microbes live inside the termite hindgut, releasing the hydrogen that was trapped inside wood and plants that they digest. Through the action of unidentified enzymes in the termite gut, lignocellulose
Lignocellulose refers to plant dry matter (biomass), so called lignocellulosic biomass. It is the most abundantly available raw material on the Earth for the production of biofuels. It is composed of two kinds of carbohydrate polymers, cellulose a ...
polymer
A polymer (; Greek '' poly-'', "many" + ''-mer'', "part")
is a substance or material consisting of very large molecules called macromolecules, composed of many repeating subunits. Due to their broad spectrum of properties, both synthetic a ...
s are broken down into sugars and are transformed into hydrogen. The bacteria within the gut turns the sugar and hydrogen into cellulose acetate
In biochemistry, cellulose acetate refers to any acetate ester of cellulose, usually cellulose diacetate. It was first prepared in 1865. A bioplastic, cellulose acetate is used as a film base in photography, as a component in some coatings, and ...
, an acetate
An acetate is a salt (chemistry), salt formed by the combination of acetic acid with a base (e.g. Alkali metal, alkaline, Alkaline earth metal, earthy, Transition metal, metallic, nonmetallic or radical Radical (chemistry), base). "Acetate" als ...
ester
In chemistry, an ester is a compound derived from an oxoacid (organic or inorganic) in which at least one hydroxyl group () is replaced by an alkoxy group (), as in the substitution reaction of a carboxylic acid and an alcohol. Glycerides ar ...
of cellulose on which termites rely for energy.[ Community DNA sequencing of the microbes in the termite hindgut has been employed to provide a better understanding of the ]metabolic pathway
In biochemistry, a metabolic pathway is a linked series of chemical reactions occurring within a cell. The reactants, products, and intermediates of an enzymatic reaction are known as metabolites, which are modified by a sequence of chemical reac ...
.[ Genetic engineering may enable hydrogen to be generated in bioreactors from woody biomass.][
The development of ]autonomous robot
An autonomous robot is a robot that acts without recourse to human control. The first autonomous robots environment were known as Elmer and Elsie, which were constructed in the late 1940s by W. Grey Walter. They were the first robots in history th ...
s capable of constructing intricate structures without human assistance has been inspired by the complex mounds that termites build. These robots work independently and can move by themselves on a tracked grid, capable of climbing and lifting up bricks. Such robots may be useful for future projects on Mars, or for building levee
A levee (), dike (American English), dyke (English in the Commonwealth of Nations, Commonwealth English), embankment, floodbank, or stop bank is a structure that is usually soil, earthen and that often runs parallel (geometry), parallel to ...
s to prevent flooding.
Termites use sophisticated means to control the temperatures of their mounds. As discussed above, the shape and orientation of the mounds of the Australian compass termite stabilises their internal temperatures during the day. As the towers heat up, the solar chimney
:''This article refers to a device for ventilation. For the power generation technology, see Solar updraft tower.''
A solar chimney often referred to as a thermal chimney is a way of improving the natural ventilation of buildings by using convecti ...
effect (stack effect
The stack effect or chimney effect is the movement of air into and out of buildings through unsealed openings, chimneys, flue-gas stacks, or other containers, resulting from air buoyancy. Buoyancy occurs due to a difference in indoor-to-outdoor a ...
) creates an updraft of air within the mound. Wind blowing across the tops of the towers enhances the circulation of air through the mounds, which also include side vents in their construction. The solar chimney effect has been in use for centuries in the Middle East
The Middle East ( ar, الشرق الأوسط, ISO 233: ) is a geopolitical region commonly encompassing Arabian Peninsula, Arabia (including the Arabian Peninsula and Bahrain), Anatolia, Asia Minor (Asian part of Turkey except Hatay Pro ...
and Near East
The ''Near East''; he, המזרח הקרוב; arc, ܕܢܚܐ ܩܪܒ; fa, خاور نزدیک, Xāvar-e nazdik; tr, Yakın Doğu is a geographical term which roughly encompasses a transcontinental region in Western Asia, that was once the hist ...
for passive cooling, as well as in Europe by the Romans
Roman or Romans most often refers to:
*Rome, the capital city of Italy
* Ancient Rome, Roman civilization from 8th century BC to 5th century AD
*Roman people, the people of ancient Rome
*''Epistle to the Romans'', shortened to ''Romans'', a lette ...
. It is only relatively recently, however, that climate responsive construction techniques have become incorporated into modern architecture. Especially in Africa, the stack effect has become a popular means to achieve natural ventilation and passive cooling in modern buildings.[
]
In culture
The Eastgate Centre is a shopping centre and office block in central Harare
Harare (; formerly Salisbury ) is the capital and most populous city of Zimbabwe. The city proper has an area of 940 km2 (371 mi2) and a population of 2.12 million in the 2012 census and an estimated 3.12 million in its metropolitan ...
, Zimbabwe, whose architect, Mick Pearce, used passive cooling
Passive cooling is a building design approach that focuses on heat gain control and heat dissipation in a building in order to improve the indoor thermal comfort with low or no energy consumption. This approach works either by preventing heat from ...
inspired by that used by the local termites. It was the first major building exploiting termite-inspired cooling techniques to attract international attention. Other such buildings include the Learning Resource Center at the Catholic University of Eastern Africa
Catholic University of Eastern Africa (CUEA), is a multi-campus private university in Kenya. It is accredited by the Commission for University Education in Kenya.
Location
The main campus of the university is located along Bogani East Roa ...
and the Council House 2
Council House 2 (also known as CH2), is an office building located at 240 Little Collins Street in the Melbourne central business district, Australia. It is used by the City of Melbourne council, and in April 2005, became the first purpose-built ...
building in Melbourne
Melbourne ( ; Boonwurrung/Woiwurrung: ''Narrm'' or ''Naarm'') is the capital and most populous city of the Australian state of Victoria, and the second-most populous city in both Australia and Oceania. Its name generally refers to a met ...
, Australia.[
Few zoos hold termites, due to the difficulty in keeping them captive and to the reluctance of authorities to permit potential pests. One of the few that do, the ]Zoo Basel
Zoo Basel is a non-profit zoo in the city of Basel, Switzerland. Its official name is ''Zoologischer Garten Basel'' — or in English: Basel Zoological Garden. Basel residents affectionately call it ''Zolli''. in Switzerland
). Swiss law does not designate a ''capital'' as such, but the federal parliament and government are installed in Bern, while other federal institutions, such as the federal courts, are in other cities (Bellinzona, Lausanne, Luzern, Neuchâtel ...
, has two thriving ''Macrotermes bellicosus'' populations – resulting in an event very rare in captivity: the mass migrations of young flying termites. This happened in September 2008, when thousands of male termites left their mound each night, died, and covered the floors and water pits of the house holding their exhibit.
African tribes in several countries have termites as totem
A totem (from oj, ᑑᑌᒼ, italics=no or ''doodem'') is a spirit being, sacred object, or symbol that serves as an emblem of a group of people, such as a family, clan, lineage, or tribe, such as in the Anishinaabe clan system.
While ''the wo ...
s, and for this reason tribe members are forbidden to eat the reproductive alates. Termites are widely used in traditional popular medicine; they are used as treatments for diseases and other conditions such as asthma, bronchitis
Bronchitis is inflammation of the bronchi (large and medium-sized airways) in the lungs that causes coughing. Bronchitis usually begins as an infection in the nose, ears, throat, or sinuses. The infection then makes its way down to the bronchi. ...
, hoarseness
A hoarse voice, also known as dysphonia or hoarseness, is when the voice involuntarily sounds breathy, raspy, or strained, or is softer in volume or lower in pitch. A hoarse voice, can be associated with a feeling of unease or scratchiness in the ...
, influenza, sinusitis
Sinusitis, also known as rhinosinusitis, is inflammation of the nasal mucosa, mucous membranes that line the paranasal sinuses, sinuses resulting in symptoms that may include thick Mucus#Respiratory system, nasal mucus, a nasal congestion, plugg ...
, tonsillitis
Tonsillitis is inflammation of the tonsils in the upper part of the throat. It can be acute or chronic. Acute tonsillitis typically has a rapid onset. Symptoms may include sore throat, fever, enlargement of the tonsils, trouble swallowing, and en ...
and whooping cough.[ In Nigeria, '' Macrotermes nigeriensis'' is used for spiritual protection and to treat wounds and sick pregnant women. In Southeast Asia, termites are used in ritual practices. In Malaysia, Singapore and Thailand, termite mounds are commonly worshiped among the populace.] Abandoned mounds are viewed as structures created by spirits, believing a local guardian dwells within the mound; this is known as Keramat
In Sunni Islam, ''karamat'' ( ar, کرامات ''karāmāt'', pl. of ''karāmah'', lit. generosity, high-mindedness) refers to supernatural wonders performed by Muslim saints. In the technical vocabulary of Islamic religious sciences, the sin ...
and Datok Kong. In urban areas, local residents construct red-painted shrines over mounds that have been abandoned, where they pray for good health, protection and luck.[
]
See also
*Stigmergy
Stigmergy ( ) is a mechanism of indirect coordination, through the environment, between agents or actions. The principle is that the trace left in the environment by an individual action stimulates the performance of a succeeding action by the sam ...
*Termite shield
A termite shield is a sheet metal fabrication used in light frame construction to reduce the movement of termites
Termites are small insects that live in colonies and have distinct castes ( eusocial) and feed on wood or other dead plant mat ...
*Xylophagy
Xylophagy is a term used in ecology to describe the habits of an herbivorous animal whose diet consists primarily (often solely) of wood. The word derives from Greek ''ξυλοφάγος'' (''xulophagos'') "eating wood", from ''ξύλον'' (') ...
Notes
References
Cited literature
*
*
External links
*
Isoptera: termites
at CSIRO Australia Entomology
Jared Leadbetter seminar: Termites and Their Symbiotic Gut Microbes
{{Authority control
Articles containing video clips
Building defects
Household pest insects
Insects in culture
Lopingian first appearances
Symbiosis
Superorganisms