Nasutitermes corniger
Encyclopedia
Nasutitermes corniger is a species of arboreal termite
that is endemic to the neotropics. It is very closely related to Nasutitermes ephratae.
The species has been studied relatively intensively, particularly on Barro Colorado Island
, Panama
. These studies and others have shown that the termite interacts with many different organisms including a bat
that roosts in its nest and various species of ant
s that cohabit with the termite.
but as they grow they become more ellipsical
. There may also be localised lobes on the surface of the nest. The queen lives in a chamber located in the centre of the nest, (often near the tree trunk or branch to which the nest is attached), that is up to 8cm wide and 1cm high and heavily reinforced. The thickness of the walls in the nest decreases away from the queen and towards the exterior although if the nest is attacked by predators then the walls will be reinforced. In one study of their nests the heaviest nest identified weighed 28 kilograms and measured 68cm by 46cm by 34cm.
Fertile individuals
of N. corniger have black wings, dark bodies, and ocelli which are located relatively far from the eyes.
s. In some years large colonies do not produce a fertile brood. Alate nymph
s develop through five instar
s and spend between 5 and 8 months within the colony before leaving to mate. When the alates are mature they typically account for 35% of the colony's biomass
. More males than females are produced from each colony but because females are heavier (by between 20 and 40%) the energy investment in each sex is similar. Newly formed colonies tend to have multiple queens and kings all living in the same royal chamber. Slightly older colonies tend to consist of multiple queens (up to 33) but only one king, in these cases the species can be considered polygynous. Over several years the species turns to being monogamous, having only one queen and one king. Being polygynous in the early stages of the colony is advantageous as it allows the colony to produce many workers in a short period of time and allows the production of female alates more quickly than if they were monogamous from the start.
, Guatemala
, Honduras
, Costa Rica
, Panama
, Venezuela
, Trinidad
, Tobago
and Bolivia
.
s cohabit the nests of N. corniger or colonise them once the termites have abandoned them. Some species prey on the termites but others do not. Studies with radioactive tracer
s have shown that when cohabiting nutrient
s flow both ways between the ants and the termites. Monacis bispinosa, also known as Dolichoderus bispinosus is one of the most common ant species to cohabit with the termites but is susceptible to their chemical defences and cannot prey on live termites. Camponotus abdominalis
associates with termites less often but is an aggressive predator of the termites. Camponotus species and Dolichoderus diversus have been found to inhabit N. corniger nests that have been abandoned.
Crematogaster brevispinosa rochai is one subspecies of ant whose interaction with N. corniger has been studied. C. b. rochai lives in areas of caatinga
in Brazil
. No queens of the ant have been found in the nests but their larvae of all castes and sexes have been. Nests that contain C. b. rochai do not have a termite queen in either. It can therefore be concluded that both the ants and termites are members of polydomous colonies that each have multiple nest sites. The ants and termites are segregated within the nest and do not normally come into contact with each other. C. b. rochai plug channels at the boundary of the areas they occupy to cause this segregation. On occasions when the ants and termites do come into contact with each other (e.g. if the nest is broken into) they are rarely aggressive and tend to avoid each other instead. It has been hypothesised that the hydrocarbon
content of their cuticle
s may have changed to allow them to live together relatively peacefully.
The termites are thought to benefit from the association as the ants leave debris in the nest containing nitrogen
and that this increases the availability of this important nutrient in an environment where it is scarce. They may also benefit from the ants protecting the nest from predators. The ants benefit as the termite nests provide an ideal location to raise broods, particularly of reproductive castes. The climate is suitable for this and the nests are easily defended against predators. Other Nasutitermes species have been found to produce anti-fungal compounds and these would also be beneficial to the ants although it is not known if N. corniger do produce such compounds.
The White-throated Round-eared Bat, Lophostoma silvicolum, roosts inside the nests of N.corniger. Males excavate the roost themselves, expending considerable energy whilst doing so. They consequently gain reproductive success as a harem
of females will join them in the roost. The termite nest is an ideal temperature for raising young and provides protection from predators of the bats. N. corniger repair the damage made to the nest by the bat meaning that the males have to constantly maintain the roost. Once the bats leave the cavity is filled by the termites within a few weeks. Scientists are currently investigating how the bats are able to create the roosts without being attacked by the termites.
Several bird species including trogon
s, puffbird
s and parakeet
s also form nests in termite nests. These can be distinguished from those made by bats as they have a horizontal entrance whereas those made by bats have a vertical entrance at the base of the nest.
of a termite very closely related to N. corniger has been analysed using metagenomics
to determine the function of different microbes in their gut.
Symbiotic bacteria
in their gut are able to fix nitrogen
at a rate of 0.25-1.0 mg N per colony per hour. This suggests a nitrogen doubling time of 200-500 days making it possible for the whole population of the colony to be replaced once or twice each year.
that fills most of their head. The secretion contains pinene
, limonene
and other high molecular weight non-volatile compounds. The secretion is effective in repelling an anteater
predator, of the Tamandua
genus. When the anteater pierces the nest soldiers swarm around the break for several minutes and later just guard the break.
Termite
Termites are a group of eusocial insects that, until recently, were classified at the taxonomic rank of order Isoptera , but are now accepted as the epifamily Termitoidae, of the cockroach order Blattodea...
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
Barro Colorado Island
Barro Colorado Island is located in the man-made Gatun Lake in the middle of the Panama Canal. The island was formed when the waters of the Chagres River were dammed to form the lake. When the waters rose, they covered a significant part of the existing rainforest, and the hilltops remained as...
, Panama
Panama
Panama , officially the Republic of Panama , is the southernmost country of Central America. Situated on the isthmus connecting North and South America, it is bordered by Costa Rica to the northwest, Colombia to the southeast, the Caribbean Sea to the north and the Pacific Ocean to the south. The...
. These studies and others have shown that the termite interacts with many different organisms including a bat
Bat
Bats are mammals of the order Chiroptera "hand" and pteron "wing") whose forelimbs form webbed wings, making them the only mammals naturally capable of true and sustained flight. By contrast, other mammals said to fly, such as flying squirrels, gliding possums, and colugos, glide rather than fly,...
that roosts in its nest and various species of ant
Ant
Ants are social insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. More than...
s that cohabit with the termite.
Description
The nests of N. corniger are dark brown on the surface and have small bumps over their exterior. When small (less than 20cm in diameter) they tend to be sphericalSphere
A sphere is a perfectly round geometrical object in three-dimensional space, such as the shape of a round ball. Like a circle in two dimensions, a perfect sphere is completely symmetrical around its center, with all points on the surface lying the same distance r from the center point...
but as they grow they become more ellipsical
Ellipse
In geometry, an ellipse is a plane curve that results from the intersection of a cone by a plane in a way that produces a closed curve. Circles are special cases of ellipses, obtained when the cutting plane is orthogonal to the cone's axis...
. There may also be localised lobes on the surface of the nest. The queen lives in a chamber located in the centre of the nest, (often near the tree trunk or branch to which the nest is attached), that is up to 8cm wide and 1cm high and heavily reinforced. The thickness of the walls in the nest decreases away from the queen and towards the exterior although if the nest is attacked by predators then the walls will be reinforced. In one study of their nests the heaviest nest identified weighed 28 kilograms and measured 68cm by 46cm by 34cm.
Fertile individuals
Alate
An alate is a winged reproductive of a social insect . Alate females are typically those destined to become queens , whereas alate males are occasionally referred to as "drones"...
of N. corniger have black wings, dark bodies, and ocelli which are located relatively far from the eyes.
Reproduction
The number of fertile individuals produced by colonies of N. corniger varies widely. Mature colonies with between 50,000 and 400,000 infertile workers generally produce between 5,000 and 25,000 alateAlate
An alate is a winged reproductive of a social insect . Alate females are typically those destined to become queens , whereas alate males are occasionally referred to as "drones"...
s. In some years large colonies do not produce a fertile brood. Alate nymph
Nymph
A nymph in Greek mythology is a female minor nature deity typically associated with a particular location or landform. Different from gods, nymphs are generally regarded as divine spirits who animate nature, and are usually depicted as beautiful, young nubile maidens who love to dance and sing;...
s develop through five instar
Instar
An instar is a developmental stage of arthropods, such as insects, between each molt , until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or assume a new form. Differences between instars can often be seen in altered body proportions, colors, patterns, or...
s and spend between 5 and 8 months within the colony before leaving to mate. When the alates are mature they typically account for 35% of the colony's biomass
Biomass
Biomass, as a renewable energy source, is biological material from living, or recently living organisms. As an energy source, biomass can either be used directly, or converted into other energy products such as biofuel....
. More males than females are produced from each colony but because females are heavier (by between 20 and 40%) the energy investment in each sex is similar. Newly formed colonies tend to have multiple queens and kings all living in the same royal chamber. Slightly older colonies tend to consist of multiple queens (up to 33) but only one king, in these cases the species can be considered polygynous. Over several years the species turns to being monogamous, having only one queen and one king. Being polygynous in the early stages of the colony is advantageous as it allows the colony to produce many workers in a short period of time and allows the production of female alates more quickly than if they were monogamous from the start.
Range
N. corniger have been found in MexicoMexico
The United Mexican States , commonly known as Mexico , is a federal constitutional republic in North America. It is bordered on the north by the United States; on the south and west by the Pacific Ocean; on the southeast by Guatemala, Belize, and the Caribbean Sea; and on the east by the Gulf of...
, Guatemala
Guatemala
Guatemala is a country in Central America bordered by Mexico to the north and west, the Pacific Ocean to the southwest, Belize to the northeast, the Caribbean to the east, and Honduras and El Salvador to the southeast...
, Honduras
Honduras
Honduras is a republic in Central America. It was previously known as Spanish Honduras to differentiate it from British Honduras, which became the modern-day state of Belize...
, Costa Rica
Costa Rica
Costa Rica , officially the Republic of Costa Rica is a multilingual, multiethnic and multicultural country in Central America, bordered by Nicaragua to the north, Panama to the southeast, the Pacific Ocean to the west and the Caribbean Sea to the east....
, Panama
Panama
Panama , officially the Republic of Panama , is the southernmost country of Central America. Situated on the isthmus connecting North and South America, it is bordered by Costa Rica to the northwest, Colombia to the southeast, the Caribbean Sea to the north and the Pacific Ocean to the south. The...
, Venezuela
Venezuela
Venezuela , officially called the Bolivarian Republic of Venezuela , is a tropical country on the northern coast of South America. It borders Colombia to the west, Guyana to the east, and Brazil to the south...
, Trinidad
Trinidad
Trinidad is the larger and more populous of the two major islands and numerous landforms which make up the island nation of Trinidad and Tobago. It is the southernmost island in the Caribbean and lies just off the northeastern coast of Venezuela. With an area of it is also the fifth largest in...
, Tobago
Tobago
Tobago is the smaller of the two main islands that make up the Republic of Trinidad and Tobago. It is located in the southern Caribbean, northeast of the island of Trinidad and southeast of Grenada. The island lies outside the hurricane belt...
and Bolivia
Bolivia
Bolivia officially known as Plurinational State of Bolivia , is a landlocked country in central South America. It is the poorest country in South America...
.
Associations
Numerous species of antAnt
Ants are social insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. More than...
s cohabit the nests of N. corniger or colonise them once the termites have abandoned them. Some species prey on the termites but others do not. Studies with radioactive tracer
Radioactive tracer
A radioactive tracer, also called a radioactive label, is a substance containing a radioisotope that is used to measure the speed of chemical processes and to track the movement of a substance through a natural system such as a cell or tissue...
s have shown that when cohabiting nutrient
Nutrient
A nutrient is a chemical that an organism needs to live and grow or a substance used in an organism's metabolism which must be taken in from its environment. They are used to build and repair tissues, regulate body processes and are converted to and used as energy...
s flow both ways between the ants and the termites. Monacis bispinosa, also known as Dolichoderus bispinosus is one of the most common ant species to cohabit with the termites but is susceptible to their chemical defences and cannot prey on live termites. Camponotus abdominalis
Camponotus abdominalis
Camponotus atriceps, previously referred as C. abdominalis, is a species of carpenter ant, endemic to the Americas.A variety of parasites have been identified from the subspecies, Camponotus abdominalis floridanus...
associates with termites less often but is an aggressive predator of the termites. Camponotus species and Dolichoderus diversus have been found to inhabit N. corniger nests that have been abandoned.
Crematogaster brevispinosa rochai is one subspecies of ant whose interaction with N. corniger has been studied. C. b. rochai lives in areas of caatinga
Caatinga
Caatinga is a type of vegetation, and an ecoregion characterized by this vegetation in the northeastern part of Brazil. The name "Caatinga" is a Tupi word meaning "white forest" or "white vegetation"...
in Brazil
Brazil
Brazil , officially the Federative Republic of Brazil , is the largest country in South America. It is the world's fifth largest country, both by geographical area and by population with over 192 million people...
. No queens of the ant have been found in the nests but their larvae of all castes and sexes have been. Nests that contain C. b. rochai do not have a termite queen in either. It can therefore be concluded that both the ants and termites are members of polydomous colonies that each have multiple nest sites. The ants and termites are segregated within the nest and do not normally come into contact with each other. C. b. rochai plug channels at the boundary of the areas they occupy to cause this segregation. On occasions when the ants and termites do come into contact with each other (e.g. if the nest is broken into) they are rarely aggressive and tend to avoid each other instead. It has been hypothesised that the hydrocarbon
Hydrocarbon
In organic chemistry, a hydrocarbon is an organic compound consisting entirely of hydrogen and carbon. Hydrocarbons from which one hydrogen atom has been removed are functional groups, called hydrocarbyls....
content of their cuticle
Cuticle
A cuticle , or cuticula, is a term used for 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 "cuticles" are non-homologous; differing in their origin, structure, function, and chemical composition...
s may have changed to allow them to live together relatively peacefully.
The termites are thought to benefit from the association as the ants leave debris in the nest containing nitrogen
Nitrogen
Nitrogen is a chemical element that has the symbol N, atomic number of 7 and atomic mass 14.00674 u. Elemental nitrogen is a colorless, odorless, tasteless, and mostly inert diatomic gas at standard conditions, constituting 78.08% by volume of Earth's atmosphere...
and that this increases the availability of this important nutrient in an environment where it is scarce. They may also benefit from the ants protecting the nest from predators. The ants benefit as the termite nests provide an ideal location to raise broods, particularly of reproductive castes. The climate is suitable for this and the nests are easily defended against predators. Other Nasutitermes species have been found to produce anti-fungal compounds and these would also be beneficial to the ants although it is not known if N. corniger do produce such compounds.
The White-throated Round-eared Bat, Lophostoma silvicolum, roosts inside the nests of N.corniger. Males excavate the roost themselves, expending considerable energy whilst doing so. They consequently gain reproductive success as a harem
Harem
Harem refers to the sphere of women in what is usually a polygynous household and their enclosed quarters which are forbidden to men...
of females will join them in the roost. The termite nest is an ideal temperature for raising young and provides protection from predators of the bats. N. corniger repair the damage made to the nest by the bat meaning that the males have to constantly maintain the roost. Once the bats leave the cavity is filled by the termites within a few weeks. Scientists are currently investigating how the bats are able to create the roosts without being attacked by the termites.
Several bird species including trogon
Trogon
The trogons and quetzals are birds in the order Trogoniformes which contains only one family, the Trogonidae. The family contains 39 species in eight genera. The fossil record of the trogons dates back 49 million years to the mid-Eocene. They might constitute a member of the basal radiation of...
s, puffbird
Puffbird
The puffbirds and their relatives in the near passerine family Bucconidae are tropical birds breeding from South America up to Mexico.They are related to the jacamars, but lack the iridescent colours of that family. They are mainly brown, rufous or grey, with large heads and flattened bills with a...
s and parakeet
Parakeet
Parakeet is a term for any one of a large number of unrelated small to medium sized species of parrot, that generally have long tail feathers...
s also form nests in termite nests. These can be distinguished from those made by bats as they have a horizontal entrance whereas those made by bats have a vertical entrance at the base of the nest.
Symbioses
The entire gut floraGut flora
Gut flora consists of microorganisms that live in the digestive tracts of animals and is the largest reservoir of human flora. In this context, gut is synonymous with intestinal, and flora with microbiota and microflora....
of a termite very closely related to N. corniger has been analysed using metagenomics
Metagenomics
Metagenomics is the study of metagenomes, genetic material recovered directly from environmental samples. The broad field may also be referred to as environmental genomics, ecogenomics or community genomics. Traditional microbiology and microbial genome sequencing rely upon cultivated clonal cultures...
to determine the function of different microbes in their gut.
Symbiotic bacteria
Bacteria
Bacteria are a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria have a wide range of shapes, ranging from spheres to rods and spirals...
in their gut are able to fix nitrogen
Nitrogen fixation
Nitrogen fixation is the natural process, either biological or abiotic, by which nitrogen in the atmosphere is converted into ammonia . This process is essential for life because fixed nitrogen is required to biosynthesize the basic building blocks of life, e.g., nucleotides for DNA and RNA and...
at a rate of 0.25-1.0 mg N per colony per hour. This suggests a nitrogen doubling time of 200-500 days making it possible for the whole population of the colony to be replaced once or twice each year.
Defenses
Soldiers of N. corniger squirt a strong-smelling, sticky secretion at predators from a large glandGland
A gland is an organ in an animal's body that synthesizes a substance for release of substances such as hormones or breast milk, often into the bloodstream or into cavities inside the body or its outer surface .- Types :...
that fills most of their head. The secretion contains pinene
Pinene
Pinene is a bicyclic monoterpene chemical compound. There are two structural isomers of pinene found in nature: α-pinene and β-pinene. As the name suggests, both forms are important constituents of pine resin; they are also found in the resins of many other conifers, as well as in non-coniferous...
, limonene
Limonene
Limonene is a colourless liquid hydrocarbon classified as a cyclic terpene. The more common D isomer possesses a strong smell of oranges. It is used in chemical synthesis as a precursor to carvone and as a renewably-based solvent in cleaning products....
and other high molecular weight non-volatile compounds. The secretion is effective in repelling an anteater
Anteater
Anteaters, also known as antbear, are the four mammal species of the suborder Vermilingua commonly known for eating ants and termites. Together with the sloths, they compose the order Pilosa...
predator, of the Tamandua
Tamandua
Tamandua is a genus of anteaters. It has two members: the Southern Tamandua and the Northern Tamandua . They live in forests and grasslands, are semi-arboreal, and possess partially prehensile tails. They mainly eat ants and termites, but they occasionally eat bees, beetles, and insect larvae...
genus. When the anteater pierces the nest soldiers swarm around the break for several minutes and later just guard the break.