The family Kalotermitidae and the subfamily Termopsinae (family Hodotermitidae) make their nests in the wood on which they feed. These termites excavate irregular networks of galleries with no external openings, except the temporary ones created during swarming. The nest galleries have partitions made of fecal matter and are lined or coated with plaster made of fecal matter. The Kalotermitidae live in the sound wood of stumps and branches of trees. Examples are Neotermes tectonae, which lives in and attacks teak trees in Java, and Cryptotermes, which bores into trees and furniture in various parts of the world. The Termopsinae live in damp rotten logs. Although true wood dwellers never invade soil, and their nests have no soil connections, all other termites are basically subterranean, building their nests either in soil or with soil connections and exploiting food sources away from the nest.
Many species of Rhinotermitidae build nests in wood that is buried in damp soil and from which a diffused network of tunnels to food sources may radiate into the soil or above the ground in the form of covered runways. Other termites build a diffused subterranean nest with many chambers or pockets in soil and a network of galleries.
Many termites build discrete and concentrated nests. Some nests rise partly above the ground as mounds or hills, whereas others are totally underground or arboreal. Dirt, particles of fine clay, or chewed wood glued together with saliva or excreta are used to build nests. During nest construction a termite deposits fecal matter to cement particles in place.
Arboreal nests are ovoid structures built of “carton” (a mixture of fecal matter and wood fragments), which resembles cardboard or papier-mâché. Carton may be papery and fragile, or woody and very hard. The inside of an arboreal nest consists of horizontal layers of cells, with the queen occupying a special compartment near the centre. The nests always maintain connections with the ground through covered runways.
The large termite mounds, or hills, which are a prominent landscape feature in the tropics, may be domelike or conical. Some have chimneys and pinnacles. Longitudinal and horizontal chambers and galleries comprise the interior. Generally the outer wall is constructed of hard soil material, distinct from the internal central portion (or nursery), which is composed of softer carton material. In northern Australia Amitermes meridionalis builds wedge-shaped mounds, called compass or magnetic mounds, that are 3 to 4 metres (9.8 to 13.1 feet) high, 2.5 metres (8.1 feet) wide, and 1 metre (3.2 feet) thick at the base. The long axis is always directed north-south, and the broad side faces east-west, an orientation that probably functions to help regulate temperature. Spectacular mounds are built by fungus-growing termites in Indomalaya and Africa. Mounds of some African Macrotermes species reach a height of 8 to 9 metres (26.2 to 29.5 feet) and have pinnacles, chimneys, and ridges on their outer walls. Such mounds are built of fine particles of clay glued together by saliva to form an exceedingly hard substance. Inside the mounds are honeycomb-like structures on which the fungus grows.
Many termite nests harbour various other invertebrates as guests (e.g., beetles, flies, bugs, caterpillars, millipedes). Some, called termitophiles, are unable to survive independent of their termite hosts. True termitophiles actually have evolved with their hosts and are species specific. Some beetles and flies have developed glands that secrete substances sought and licked by the termites. The termite nest, because it provides shelter and warmth, may also be occupied by lizards, snakes, scorpions, and some birds.
A few termites, known as inquilinous species, live only in obligatory association with other termite species. Examples of such obligate relationships are Ahamitermes and Incolitermes species, which live only in the mound nests of certain Coptotermes species. In these, the galleries of guests and hosts are completely separate. Inquilinous species feed on the inner carton material of the host nests. Incolitermes, however, depend on the host species not only for food but also for exit holes from the nest during swarming when alates of the inquiline and the host emerge together. Such species’ tolerance is highly unusual. Normally, different species of termites are hostile to one another, and host termites may attack inquilinous guests if partitions between galleries are broken.
Form and function
Castes and their roles
The primary reproductives in a termite colony are usually one royal pair, a king and queen. They have developed from winged forms (alates) that have flown from a parent colony and shed their wings. Because they spend time outside of the colony on the mating flight, they have hardened, pigmented bodies and large compound eyes. The primary reproductives have several important functions: reproduction, dispersal, and colony formation. In addition, during the initial stages of colony formation, the primary reproductives perform tasks later taken over by the worker caste (e.g., nest construction, housekeeping, care of young).
If the king or queen dies, it is replaced by several supplementary reproductives that are slightly pigmented and have either short wing pads (brachypterous) or none (apterous) and reduced compound eyes. These secondary reproductives, which develop from nymphs and may be called neotenics, normally are not present in a colony as long as the primary reproductives remain healthy. If a primary reproductive is lost, a neotenic achieves sexual maturity without attaining a fully winged adult stage or leaving the nest.