Colonies and life cycles

Dispersal of winged reproductives from established colonies is common to all termites and it is an important means of establishing new colonies. Winged forms or alates are usually produced by relatively mature colonies, and, depending on the species and resources, this occurs 3-5 years after the colony founding. In some of the primitive families, small alates are released over several months, but often there is a seasonal peak. In higher termites, large numbers of alates emerge at one time. The swarming of alates from these colonies is usually restricted to certain times of the year, and specific times of the day, evening, or night. Swarming is synchronized with regional and local weather conditions; it occurs during warm months in temperate regions and with the arrival of the rainy season in the tropics. The exact time of year and day that swarming takes place is species-specific, and dependent on the condition of the colony. Soldiers and workers are involved in excavating exit holes and protecting the emerging alates and the colony against predators. The number of winged reproductives produced ranges from 1 to 43% of the total individuals in the colony.

The flight oftermites is weak and fluttering, and their flight direction and distance are often directed by winds. In general, the alates of primitive wood-dwelling termites fly further than do the highly evolved species. The latter tend to lose their wings after a short initial flight. They often lose one or more wings in flight, and they fly erratically to the ground; the process offlight weakens the wing at the basal suture and the wings are shed when they land. In the lower termites, the wings are stronger and a series offlights occur. To remove the wings the termites brace against some irregularity of the substrate and turn and twist to break the wings. Generally, alates are attracted to areas of greatest light intensity. Evening- and night-flying termites are often attracted to lights and windows. The emergence times for alates of different species in the same area are separated, and there is usually no overlap or more than one species flying at the same time. Typically, the flight of a single species occurs from many colonies over a wide area at the same time. Sporadic flights from isolated colonies are often encountered in urban areas, where the conditions suitable for flight exist only in small habitats. The emergence of alates from a colony may be interrupted before all the alates emerge. This results in a series of small and large flights from a colony during one season.

Alates are in the air for a short time, and predation by other insects and birds usually takes about 99% of the swarm. In the higher termites, the female lands first and assumes a position in which the abdomen is elevated to release a chemical sex attractant from either tergal or sternal glands. When a male approaches, and perhaps contacts the female, both sexes shed their wings and the female leads the male to a suitable nest site in moist soil. The dampwood termites seek cracks or holes in timbers above ground; subterranean termites select moist soil or decayed wood in tree stumps. Drywood termites fly to sites in buildings, or crawl up the outside to reach roofing timbers and begin excavating in cracks or regions softened by decay fungi. Mating usually does not occur until the male and female prepare and enter a small chamber and the entrance hole is sealed. This takes 1-2 days and involves displacing soil to make a cavity or chewing into a wood substrate. Establishment of a new colony by flight of alates is dependent on the survival of both reproductives until they are in a secure location and the female is inseminated. Continued development of the colony is dependent upon reinsemination of the female.

Egg-laying begins 1-2 weeks after mating. In lower termites, about 20 eggs are laid in the first batch. In higher termites, aboutioo eggs are laid by young queens. Hatching takes 30-80 days, depending on species and temperature. After hatching, larvae are fed salivary and anal secretions from the male and female, but later in their development they care for the eggs produced by the queen, begin to expand the nest, and forage for food. At this early stage, the search for food is essential to the health and fertility of the male and female, because they do notleave the nestto feed. Chambers thatare formed in wood are a ready source of food for the new workers and there is little need to leave the site, but subterranean termites must travel away from the nest site to find food. The number of workers and soldiers in the colony increases slowly. The colony enlarges with the addition of workers, and new galleries are excavated, and the queen and king are isolated in a cell ata central position in the nest. In most of the higher termites, the abdomen of the queen enlarges and, in this physogastric condition, she is nearly immobile. The king does not increase in size, and remains with the queen. Mating occurs throughout their life, which is up to 15 years. In lower termites, the king and queen are not isolated, and in some drywood termites, they move around the nest. Mating takes place atintervals and the queen's abdomen only slightly enlarges with the development of eggs. The rate of egg-laying in lower termites is several hundred to several thousand eggs per year.

The queen and king are groomed and fed by workers, which enter the royal cell through small openings in its hard protective wall. Queens lay eggs at a steady rate; workers carry the eggs to incubation chambers. The number of eggs produced by the queen varies depending on the species, and the age of the queen. In tropical regions, egg production is continuous throughout the year, although there are seasonal fluctuations. In termites living in temperate regions, egg production is often suspended during the cold months. After hatching, young larvae are taken to nursery chambers where they are fed and groomed by workers. They are moved to other chambers in the nest until their final molt into workers or soldiers. Subterranean termites use undigested and partially digested cellulose to build galleries and extend the nest. This material is mixed with excreted soil particles and moistened with saliva to make a pliable material that is formed into chamber walls.

Composition of the colony, in terms of young and mature forms, varies for different species and during different periods of the year. The cyclic presence in the colony of immature and terminal forms is more pronounced in termites in temperate regions than in those in the tropics. In north temperate regions colony activity and egg production are reduced during winter months. In mature colonies, the alates usually appear over a short time before emergence. They remain in the nest for an extended period if environmental conditions are not suitable for their release. The number ofsoldiers in the colony usually increases with the production of alates, and reaches a peak before swarming flights occur. The maximum number ofsol-diers in a colony coincides with the time the nest is opened to release alates, and when exposure to predators is highest.

Young colonies grow in relation to the rate eggs are laid during the first few years, and on the length of life of the colony members. Colony size is expanded by increased egg production and retention of workers to care for the young and forage for food. In lower termites, young colonies are distinguished by queens that feed on the available substrate and produce a small number of eggs for several years. These female alates emerge without fully developed eggs; development of eggs is completed when the young queen begins to feed. If feeding is delayed, egg-laying is also delayed. After the first group is deposited, egg production usually stops for several months; a second period of egg production may be followed by a period of rest and more feeding. As the young queen matures, a large number of eggs are laid during each egg-laying period, and the number of workers in the colony increases, along with the overall size of the colony. Mature colonies are usually defined as producing all castes and young normally produced by the species, including alates. The time required for a young colony to develop to a point where it will produce alates varies with species and local conditions, and the unique features ofindi-vidual colonies.

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