Evolution, paleontology, and classification
Termites are related to the roaches and probably have evolved from a primitive roachlike ancestor. The most primitive living roach, the subsocial, wood-eating Cryptocercus punctulatus, which lives in rotten logs, has affinities with the termites. Cryptocercus harbours symbiotic, cellulose-digesting protozoans of the same genera as those found in the hindgut of primitive termites. The genitalia and certain internal structures of Cryptocercus have basic anatomic resemblances to those of the most primitive living termite, Mastotermes darwiniensis, from Australia. Mastotermes has further affinities with other roaches: its hind wing has a folded anal lobe, and its eggs are not laid singly as those of other termites but in clusters held together by a gelatinous material resembling the egg case of roaches.
Evidence of the relationship to primitive roaches suggests that termites evolved in the Late Permian (approximately 251,000,000 years ago), although the known fossil termites date only from the Early Cretaceous (about 130,000,000 years ago). The termite social system may be older than any other society. Ant social systems are estimated to be only 100,000,000 years old.
Classification
Termites of the order Isoptera are small to medium-sized insects that live in social groups, or colonies, and are characterized by their highly developed caste system. The mouthparts are modified for chewing. Antennae are moniliform (beadlike) or filiform (threadlike). Isopterans are very soft-bodied insects, usually light in colour. Head structures and the presence or absence of individual caste members are used to distinguish termite families.
Termites, although often called white ants, differ from hymenopterans (bees, ants, and wasps) in several ways. Termites have a hemimetabolous (gradual) metamorphosis and pass through a series of nymphal stages. Hymenopterans have the more common holometabolous metamorphosis, with distinct larval, pupal, and adult stages. Termite social castes (reproductives, sterile workers, and sterile soldiers) usually contain members of both sexes in equal numbers, and both males and females develop from fertilized eggs. In the hymenopteran colony, however, the sterile castes contain females only, with both sterile and reproductive females developing from fertilized eggs. Reproductive males develop by parthenogenesis from unfertilized eggs. The thorax in termites is joined broadly to the abdomen, without the “waist” characteristic of bees, ants, and wasps. Termites have two pairs of membranous wings, nearly equal in size, that break along a suture when shed, leaving only the wing base, or “scale,” attached to the thorax. This is probably the most distinguishing characteristic of isopterans.
Annotated classification
- Order Isoptera (termites)
- Highly developed caste system, may contain reproductives, soldiers, and workers; reproductives shed wings after mating; distribution worldwide, mostly in tropical rainforests; about 2,750 living, 60 fossil species; may inhabit moist subterranean or hot, dry locations; foods include plant cellulose, often digested by symbiotic protozoans in termite hindgut; all families (except Termitidae) known collectively as “lower termites” contain symbiotic protozoans in hindgut.
- Family Mastotermitidae
- Primitive; 1 living species (Mastotermes darwiniensis) in Australia; 13 Cenozoic fossil species worldwide.
- Family Kalotermitidae (dry-wood termites)
- Wood-dwelling, wood-eating; survive dry conditions; 292 living, 11 fossil species (some from Baltic amber).
- Family Hodotermitidae
- Thirty living, 13 fossil species (1, the earliest known termite fossil, from Lower Cretaceous, Labrador); includes rotten-wood termites and harvester termites that forage and store food in nests; Zootermopsis, largest termite in North America, found in Rocky Mountains at altitudes of 2,000 to 2,500 metres; Archotermopsis, found in Himalayas; Hodotermes species, serious pests of African grasslands.
- Family Rhinotermitidae (subterranean termites)
- Lives under damp conditions; 158 living, 13 fossil species; Reticulitermes, widely distributed in North America and other temperate and subtemperate regions and a serious pest; Coptotermes, a serious pest in tropical and subtropical regions.
- Family Serritermitidae
- One living species in South America; specialized family evolved from Rhinotermitidae.
- Family Termitidae (higher termites)
- Largest termite family (about 75 percent of all termites), 2,100 living, 3 fossil species; 4 subfamilies variable in morphology, social organization, and nesting habits.