现存等翅下目名录 Extant Isoptera
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现存等翅下目（白蚁）拉丁学名—中文正名对照名录 Extant Isoptera (Termites) — Latin–Chinese Name Comparison List

拉丁学名 · 中文正名 · 双语对照 —— 13 科 · 18 亚科 · 326 属 · 3021 种 · 68 亚种

Termite Basics: Biology

← Back to the Basics index; structures are described on the morphology page.

The morphology page answers “what does it look like”; this page answers “how does it come about, where does it live, and what does it eat”. The three questions interlock: caste differentiation decides who forages, who defends and who reproduces; the nest keeps temperature, humidity and food within a narrow range; feeding habits and gut symbionts decide which kind of plant debris a colony can take apart. Entries draw on Fauna Sinica, Insecta Vol. 17, Isoptera (Huang Fusheng et al., 2000), Termitology (Cheng Dongbao & Yang Zhaofen, 2014) and Biology of Termites: A Modern Synthesis (Bignell, Roisin & Lo, 2010); where recent work has revised older accounts, the text says so, and the sources are listed at the end.

Among social insects, termites are one of the few groups in which both sexes do the work: workers and soldiers are male and female alike, unlike the Hymenoptera, where the female performs all the labour. Caste, nest and food are the three lines that grow out of that single setting.

1. Castes and development

Termites are hemimetabolous: an egg hatches into a series of larval instars, wing buds appear externally in the nymph, and the final moult produces the alate. Around reproduction and labour the colony forms three body plans — reproductive, worker and soldier. The morphological differences are listed one by one on the morphology page; here the question is how they arise.

Eggs are carried, stacked and repeatedly licked by workers; antibacterial compounds in the saliva limit mould, and incubation runs from weeks to months depending on temperature and humidity. Larvae have no wing buds and depend on workers for food and care. In nymphs the buds grow with each instar; the number of instars varies with taxon and temperature, commonly five to eight. Shortly before a moult the integument pales and activity drops, and colonies tend to gather pre-moult individuals in one corner of a chamber.

Pseudergates and true workers are the key to termite development. In the lower families (Kalotermitidae, Archotermopsidae, Heterotermitidae and others) the worker-form individual is usually a pseudergate: intermediate between larva and nymph, without wing buds, and developmentally open — it can moult into a nymph (and thence an alate), a soldier, or a supplementary reproductive. The true workers of Termitidae are fixed in form and behaviour and no longer moult into alates; in some lineages they can still give rise to adultoid supplementary reproductives. Pseudergates let a colony replace lost reproductives on the spot; true workers lock the division of labour into place and buy efficiency with it — the ecological range of Termitidae is hard to imagine without that step.

What decides a caste is one of the most studied questions in termite biology, and the current answer is several strands laid over each other:

Soldiers fall into two morphological camps, mandibulate and nasute (morphology page §5), and within a species often into two sizes, as in the major and minor soldiers of Macrotermes. Most authors hold that soldiers arose once in the common ancestor of Isoptera and were then lost independently in several lineages — some genera of Apicotermitinae (Apicotermes, Anoplotermes) have no soldiers at all and rely on chemical defence, nest walls and rapid retreat. Soldiers are normally sterile, but Stolotermitidae and Archotermopsidae include reports of soldier-form supplementary reproductives: individuals that keep soldier morphology while the gonads go on developing and eggs are laid. “Soldier” and “sterile” are the norm, not an iron law.

Reproductive castes come by two routes. Primary reproductives are the alates: they swarm, shed their wings, pair and found a colony. Supplementary reproductives appear when the primary queen dies or a colony splits, and take three forms — brachypterous, apterous and adultoid. The queen’s abdomen swells enormously with the ovaries (physogastry), the intersegmental membranes are exposed, the body may be several times a worker’s length, and the king mates with her at intervals.

Term Chinese Notes
pseudergate 拟工蚁 Worker-form individual in lower families; no wing buds, developmentally reversible
true worker 真工蚁 Fixed worker of Termitidae; no longer moults into an alate
larva 幼蚁 Instar before wing buds appear
nymph 若蚁 Instar with external wing buds; moults into the alate
neotenic / supplementary reproductive 补充繁殖蚁 Derived from pseudergates, nymphs or workers; brachypterous, apterous or adultoid
physogastry 膨胀腹部 Queen’s abdomen distended by the ovaries, intersegmental membranes exposed
soldier dimorphism 兵蚁二型 Major and minor soldiers in one species, as in Macrotermes
soldier-form supplementary reproductive 兵蚁型补充繁殖蚁 Soldier morphology retained, gonads developed, eggs laid

2. Nesting

A nest does more than house the colony: it holds temperature and humidity within a narrow band, keeps out ants, lizards and fungi, stores food, and contains the chambers where brood is raised and — in the fungus-growing lineage — the fungus comb. Nests are sorted by position and material:

Nest type Position and material Typical taxa
Wood nest Tunnels and chambers cut directly in timber, no separate structure Kalotermitidae (Cryptotermes and others), Archotermopsidae (damp wood)
Subterranean nest Chambers and galleries in soil, surface marked only by mud tubes and sheets Macrotermitinae (Odontotermes and others), other Termitidae
Mound Soil particles cemented into ridges, towers or cones above ground Macrotermes, Cubitermitinae (Cubitermes and others)
Arboreal carton nest Board-like nest of wood fragments, saliva and faeces on a trunk or branch Nasutitermes, Microcerotermes
Cryptic nest No distinct structure; colony in wall voids, tree holes or pipes Heterotermitidae (Coptotermes, Reticulitermes)

A nest usually consists of the royal cell (the main chamber with queen and king), several satellite nests, runways (shelter tubes, mud tubes) connecting them, and ventilation pores. Runways are covered galleries that keep humidity in and enemies out; walls and runways are built from different mixes of soil, wood fragments, saliva and excreta, which set hard and shed water far better than loose soil.

The fungus comb is where nesting and feeding meet. Macrotermitinae stack chewed wood and faeces into a spongy or honeycombed comb inside the nest and inoculate it with Termitomyces mycelium; workers eat the pre-digested substrate and the mycelium itself, and ageing comb is removed or re-inoculated. The comb moves part of digestion outside the body and lets the colony use more heavily lignified material. Once a comb matures, Termitomyces fruiting bodies can push through the nest — a reliable field sign for this group.

How the nest climate is held steady has several overlapping explanations. Lüscher (1961) proposed a “thermosiphon”: metabolic heat drives air up a central chimney and down through surface pores, and this model long underwrote the description of mounds as natural air conditioners. Turner’s measurements on nearly fifty South African mounds in the 1990s revised that: the mound is primarily a gas-exchange organ, its porous wall working like an alveolus, with wind pressure and diffusion carrying oxygen in and carbon dioxide out; King, Ocko and Mahadevan (2015) then showed with measurements and models that day–night temperature swings are the main driver of that exchange. The safe statement today is that different nest types use different strategies — mounds rely on thick walls and thermal inertia, arboreal cartons on thin walls and position, wood nests hardly regulate at all and depend on the timber for insulation; mound interiors commonly sit at 25–30 °C and near-saturated humidity. “Air conditioner” survives as a popular image, not as a mechanism.

Nest building is collective: workers carry soil and wood fragments, cement them with saliva and faeces, and build upwards and outwards along humidity and pheromone gradients; damage is first sealed and then repaired, and the colony may move altogether if the nest fails. A growing colony can found new nests by budding, still connected to the old one by runways, which produces networks of satellite nests.

3. Feeding, digestion and symbiosis

Feeding habits are best classified by substrate and by stage of decay, and a single family often spans several classes, so taxon and habit have to be read separately:

Feeding type What is eaten Typical taxa Gut and symbionts
Wood-feeding Dead standing and fallen timber Kalotermitidae, Archotermopsidae, some Termitidae P3 paunch packed with flagellates in lower termites
Harvesting Dry grass and leaf litter, cut and stored Hodotermitidae Large crop, obvious storage behaviour
Soil-feeding Organic-rich soil and humus Several Termitidae subfamilies Very long, compartmentalised hindgut, pH up to 10 or more
Fungus-growing Substrate pre-digested in the comb, plus mycelium Macrotermitinae Depends on the comb; does not digest raw wood
Scavenging and keratin Colony corpses, exuviae, keratin debris Many, mostly facultative Works with the gut flora

Foraging happens under cover: workers reach food along runways, cut and carry it with the mandibles, and distribute it through the colony by trophallaxis; a new food source is marked with sternal gland secretion, and the colony concentrates on it. Hodotermitidae cut grass into lengths and store it in the nest for the dry season.

The gut is divided into fore-, mid- and hindgut (names on the morphology page §6). The proventriculus of the foregut carries ridges and spines and grinds the food; the midgut has a peritrophic membrane; the hindgut is the fermentation chamber and the home of the symbionts. In lower termites the P3 paunch is full of flagellates, cellulose is broken down anaerobically, and the products pass to gut bacteria and archaea; Termitidae have lost the flagellates and rely on bacteria and their own enzymes instead, with the hindgut divided into P1–P5, a mixed segment and an enteric valve. Soil-feeders have especially long hindguts and a pH that can rise above 10, which helps desorb organic matter from humus. In some taxa gut bacteria fix atmospheric nitrogen, making up for the nitrogen poverty of wood.

Digestion is a joint operation: the termite secretes some cellulases and hemicellulases, flagellates or bacteria supply the rest of the enzyme set, and lignin is only partly broken down, so faeces still carry lignin and some cellulose — which become food for other termites or soil organisms. Actinobacteria on the cuticle and in the nest produce antibiotic compounds that suppress fungal pathogens (see §6).

Evolutionarily, wood-feeding is the ancestral condition, soil-feeding arose repeatedly within Termitidae, and fungus-growing arose once, in the Macrotermitinae, roughly 31–37 million years ago, about when that subfamily radiated in Africa. Each shift came with changes in gut form, symbiont community and social behaviour, and together they explain how Termitidae came to dominate both tropical soils and dead wood.

The ecological consequences run in three directions: faster breakdown of dead wood and litter, returning carbon and nitrogen to the soil; altered soil porosity and water retention; and methane emission. Early estimates of termite methane were high and later independent measurements revised them downward, so old figures should be cited with their date and method.

4. Life history and colony founding

Mature colonies swarm around the start of the rainy season: alates leave mostly at dusk or at night, are attracted to light, and fly anything from tens of metres to several kilometres. On landing they shed the wings along the basal suture, pair up, and find a crack, a clod or a piece of dead wood for the copularium; after mating the first eggs are laid, and once the first workers eclose and begin to forage and extend the nest the colony is truly founded. Founding success is low — most pairs are dead within months.

Growth follows an exponential-then-saturating curve: a mature colony may hold from tens of thousands to millions of individuals, the queen may live for decades, and the colony itself can last as long. Besides swarming, some taxa found new colonies by budding, and Reticulitermes and relatives have been reported to produce supplementary reproductives by parthenogenesis, letting a colony continue without a mate.

5. Social behaviour and communication

The commonest interaction in the nest is trophallaxis: stomodeal trophallaxis passes food and saliva, proctodeal trophallaxis passes hindgut contents and symbionts — newly eclosed workers acquire their gut flora this way. Grooming removes pathogens and foreign particles from the cuticle, and cannibalism and corpse removal recycle nutrients.

Alarm and recruitment run on two systems, chemical and vibrational: disturbed workers bang their heads against the nest wall or shake the body, and the vibration travels along the runways; alarm and trail pheromones come from different glands, one dispersing and the other guiding. Nestmate recognition uses blends of cuticular hydrocarbons, and an outsider is usually attacked because its profile does not match. In the division of labour, large workers tend to cut and carry while small workers tend brood and repairs; age polyethism is weakly supported in termites, and size polyethism is the clearer pattern.

6. Defence and enemies

The first line is the nest itself: thick walls, closed runways and narrow entrances keep most predators out, and soldiers of Cryptotermes can plug a hole with the head (phragmosis). The second line is the soldier: mandibulate soldiers bite, nasute soldiers eject sticky diterpene secretions through the fontanelle, and Coptotermes soldiers release a milky acidic fluid. The third line is chemical and microbial: actinobacteria on the cuticle produce antibiotics that suppress Metarhizium and Beauveria.

Soldierless taxa have another answer: workers of some Apicotermitinae rupture themselves (autothysis), throwing sticky or toxic gut contents at intruders and sealing the breach; soldiers of Serritermitidae and workers of some Termitidae do something similar.

Enemies include ants, pangolins, anteaters, birds, lizards and frogs, as well as fungi and nematodes. Against ants, colonies seal entrances, reinforce the guard and move if necessary; when a nest is breached, workers repair the passages around the royal cell first.

7. Ecosystem roles and people

Termites are classic ecosystem engineers: feeding, nesting and excretion mix dead wood, litter and soil over and over, changing porosity, water retention and nutrient distribution, and mounds and carton nests become microhabitats for other invertebrates and plants. In the tropics and subtropics they are among the main agents breaking down woody debris, and their guts also house methanogens, so the size of their methane flux and the way it is estimated are still debated.

For people, termites are both pests and a resource. Damage falls into three groups: buildings and timber (Coptotermes, Reticulitermes, Cryptotermes), trees and crops (Odontotermes, Macrotermes), and dams and buried structures (nest systems of Odontotermes can run through an embankment). Control relies on monitoring and baiting, physical and chemical barriers, and above all on cutting the colony off from food and moisture rather than spraying wherever a termite is seen. As a resource, alates are eaten, combs and Termitomyces are traditional foods, and nest architecture has inspired building ventilation — the Eastgate Centre in Harare, Zimbabwe (completed 1996) is routinely cited as a biomimetic case, its ducts and chimneys laid out on the then-current thermosiphon model. Later measurements showed the mound to be primarily a gas-exchange organ, not a temperature regulator, and the building’s savings come largely from the thermal mass of its concrete slabs and from passive design rather than from close imitation of the nest. Cited as a biomimetic success, it should carry that caveat.

Corrections of fact or terminology are welcome on the guestbook. Classification follows the site checklist: Coptotermes and Reticulitermes, long placed in Rhinotermitidae, are placed in Heterotermitidae here.

Appendix: where this page differs from older literature

This page was checked against work of the past two decades after drafting, and the text follows the later results. Readers holding a textbook or popular account from around 2000 should note the following:

References