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 ants that cohabit with the termite.
2 Social Behavior
2.1 Fortress defense
The nests of N. corniger are dark brown on the surface and have small
bumps over their exterior. When small (less than 20 cm in
diameter) they tend to be spherical but as they grow they become more
elliptical. 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 8 cm wide and 1 cm 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
68 cm by 46 cm by 34 cm.
Fertile individuals of N. corniger have black wings, dark bodies, and
ocelli which are located relatively far from the eyes.
Termite colonies are examples of eusocial insects.
are animals that develop large, multigenerational cooperative
societies that assist each other in the rearing of young, often at the
cost of an individual’s life or reproductive ability. Such altruism
is explained in that eusocial insects benefit from giving up
reproductive ability of many individuals to improve the overall
fitness of closely related offspring. Hamilton’s rule is the key to
explaining this phenomenon, where altruism is justified evolutionarily
when the benefit to the individual receiving the help, weighted by the
relatedness to said individual, outweighs the cost to the organism
being altruistic. In most cases, termites included, individuals
specialize to fill different needs that the overall colony may have.
These are called castes. In
Nasutitermes as well as most other termite
species, there are three main castes: reproductive alates, workers,
The benefits to being altruistic come in two ecological modes: “life
insurers” and “fortress defenders”. Most Hymenoptera, the
large majority of social insects, are life insurers, where eusociality
is adapted as a safeguard from decreased life expectancy of
offspring. Termites, as fortress defenders, benefit from working
together to best exploit a valuable ecological resource, in the case
Nasutitermes corniger a vast wood gallery. Fortress defense is
sufficient to evolve eusociality when three criteria are met: food
coinciding with shelter, selection for defense against intruders and
predators, and the ability to defend such a habitat. Termite
colonies are generally large enclosed nests or mounds that house large
supplies of wood for the termites to exploit, fulfilling the first
criteria for fortress defense. In N. corniger, the soldier caste has
had their heads modified to spew a noxious, sticky liquid when under
Tamandua anteaters. The secretion contains pinene,
limonene and other high molecular weight compounds that deter the
anteater from returning. The termites then remain on guard near the
breach for several minutes. This adaptive morphology and defense of
the habitat are sufficient for satisfying the second two criteria for
fortress defense. The fortress defense strategy necessitated the
evolution of soldiers first, which has resulted in the unique
specialization of the nasute termites.
Nasutitermes corniger exhibits a large amount of aggression to rival
conspecific colonies. This implies that there is a method of kin
recognition among N. corniger that allow it to distinguish between its
colony and the next. While specific studies have not been done in N.
corniger, similar species in Microcerotermes and many other termites
show that they are able to detect scent on each other. It has been
shown that some separate colonies display relatively low aggression to
each other and oftentimes result in colony fusion. It is of note that
these colonies will show massive aggression towards other colonies,
indicating it is not a loss in aggressive behavior but a failure in
recognition. It has been shown that colonies that exhibit this
nonaggressive behavior have a relatively low average within-colony
relatedness of .35, whereas colonies that retained mutually aggressive
behavior had a higher relatedness average of .55. The nonaggressive
colonies often had polygamous reproductive individuals, and may have a
broader template of acceptable odor clues, leading to recognition of
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 alates.
In some years large colonies do not produce a fertile brood. Alate
nymphs develop through five instars 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.
N. corniger have been found in Mexico, Guatemala, Honduras, Costa
Rica, Panama, Venezuela, Trinidad,
Tobago and Bolivia and more
recently in Florida.
Numerous species of ants 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 tracers have
shown that when cohabiting nutrients 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
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 cuticles 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
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 trogons, puffbirds and parakeets 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.
The entire gut flora of a termite very closely related to N. corniger
has been analysed using metagenomics to determine the function of
different microbes in their gut. Typical to all wood-feeding
higher termites, bacterial gut microbiota in the guts of N. corniger
are dominated by insect-specific members of TG3_(candidate_phylum),
Fibrobacteres, and Spirochaetes. It has also been shown that the
same bacterial lineages are preferentially enriched in the
cellulolytic bacterial community that is associated with wood
particles in the gut. In addition to a role in fibre digestion,
Symbiotic bacteria in N. corniger have also been shown 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
^ a b c d Thorne, Barbara L. (1980). "Differences in Nest Architecture
Between the Neotropical
Nasutitermes corniger and
Nasutitermes ephratae(Isoptera: Termitidae)". Psyche. 87: 235–244.
Kin selection and social insects". 48: 165–175.
^ Queller, DC. "The evolution of eusociality: Reproductive head starts
of workers". Proc Natl Acad Sci U S A. 86: 3224–6.
doi:10.1073/pnas.86.9.3224. PMC 287102 .
^ "Three conditions for the evolution of eusociality: Are they
sufficient?". Insectes Sociaux. 41: 395–400.
^ "Defenses of
Nasutitermes Termites (Isoptera, Termitidae) Against
Tamandua Anteaters (Edentata, Myrmecophagidae)".
^ "Relatedness, recognition errors, and colony fusion in the termite
Nasutitermes corniger". Behavioral Ecology and Sociobiology. 61:
Alate production and sex ratio in colonies of the Neotropical
Nasutitermes corniger (Isoptera; Termitidae)". Oecologia. 58:
^ "Polygyny in the Neotropical termite
Nasutitermes corniger: life
history consequences of queen mutualism". Behavioral Ecology and
Sociobiology. 14: 117–136. doi:10.1007/BF00291903.
Nasutitermes corniger, a dangerous termite, arrives in South
Florida". Archived from the original on 2012-05-12. Retrieved
^ a b c "Behavioural Interactions Between Crematogaster brevispinosa
rochai Forel (Hymenoptera: Formicidae) and Two
(Isoptera: Termitidae)". Journal of
Insect Behavior. 18: 1–17.
^ "Mating system of a Neotropical roost-making bat: the
white-throated, round-eared bat,
Lophostoma silvicolum (Chiroptera:
Phyllostomidae)". Behavioral Ecology and Sociobiology. 58: 316–325.
University of Ulm
University of Ulm Institute of Experimental Ecology
^ Lubin, Yael D.; Montgomery, G. Gene (1981). "Defenses of
Nasutitermes Termites (Isoptera, Termitidae) Against Tamandua
Anteaters (Edentata, Myrmecophagidae)". Biotropica. 13 (1): 66–76.
doi:10.2307/2387872. JSTOR 2387872.
^ Roost Structure, Modification, and Availability in the
White-throated Round-eared Bat,
Lophostoma silvicolum (Phyllostomidae)
Living in Active
^ Warnecke, Falk; et al. (2007). "Metagenomic and functional analysis
of hindgut microbiota of a wood-feeding higher termite" (PDF). Nature.
450: 560–565. doi:10.1038/nature06269. CS1 maint: Explicit use
of et al. (link)
^ Mikaelyan, A.; Dietrich, C.; Köhler, T.; Poulsen, M.;
Sillam-Dussès, D.; Brune, A. (2015). "Diet is the primary determinant
of bacterial community structure in the guts of higher termites".
Molecular Ecology. 24 (20): 5824–5895. doi:10.1111/mec.13376.
^ Mikaelyan, A.; Strassert, J.; Tokuda, G.; Brune, A. (2014). "The
fibre-associated cellulolytic bacterial community in the hindgut of
wood-feeding higher termites (
Nasutitermes spp.)". Environmental
Microbiology. 16 (9): 2711–2722. doi:10.1111/1462-2920.12425.
Nitrogen fixation by intact colonies of the termite Nasutitermes