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PicturesTop of page
|Copyright||Georg Goergen/IITA Insect Museum, Cotonou, Benin|
|Adult||Georg Goergen/IITA Insect Museum, Cotonou, Benin|
|Title||Adult - line drawing|
|Caption||C. formosanus - adult worker.|
|Adult - line drawing||C. formosanus - adult worker.||IRRI|
IdentityTop of page
Preferred Scientific Name
Preferred Common Name
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Arthropoda
- Subphylum: Uniramia
- Class: Insecta
- Order: Isoptera
- Family: Rhinotermitidae
- Genus: Coptotermes
Notes on Taxonomy and NomenclatureTop of page
The following species of Coptotermes are found in South-East Asia: C. betongensis (Malaysia), C. boetonensis (Boeton Islands), C. borneensis (Borneo), C. curvignathus (Myanmar, Vietnam, Cambodia, Peninsular Malaysia, Sumatra, Java, Borneo, Sulawesi, Philippines, Papua New Guinea, Thailand), C. ceylonicus (Vietnam), C. elisae (New Guinea), C. havilandi (Borneo, Java, Thailand), C. flavicephalus (Philippines), C. formosanus (China, Hong Kong, Japan), C. gestroi (Malaysia, Sulawesi), C. kalshoveni (Java, Sabah, Thailand, Sumatra), C. menadoensis (Sulawesi), C. minutissimus (Koena Islands, Sulawesi), C. oshimae (Sulawesi), C. peregrinator (Sulawesi), C. premrasmii (Thailand), C. sepangensis (Malaysia, Sabah), C. sinbangensis (Sumatra), C. travians (Malaysia, Sumatra, Java, Borneo), C. vastator (Philippines). Thirty more species have been identified in China (Li, 1994; Li et al., 1994); Xia and He (1986) developed a key for 24 of these species. The taxonomic status of Coptotermes needs revision.
The most important recent work on the taxonomy of South-East Asian species of Coptotermes has been the morphometric studies of Kirton (1995) who investigated the species known to occur in peninsular Malaysia, several of which are widely distributed throughout the region. Kirton made several changes in nomenclature, and it should be noted that the well known South-East Asian species C. curvignathus is considered a synonym of C. elisae. As such, C. elisae has a geographic distribution stretching from Papua New Guinea, through Sulawesi, Philippines, Java, Borneo, and Sumatra, to Peninsular Malaysia, Thailand, Cambodia and Vietnam. In addition, Kirton supports Tho's (1992) view that C. betongensis is not a valid species but a synonym of C. sepangensis.
DescriptionTop of page
The genus Coptotermes is characterized by the presence of a pear-shaped head, narrow at front with a pointed labrum in the soldier caste (Pearce et al., 1993). Mandibles are slender, sharply pointed and slightly incurved without marginal teeth. Most distinctive in the soldier caste is the large fontanelle (opening) at the front of the head which exudes a white defence secretion when the insect is disturbed. Coptotermes have been shown to possess, as for other members of the Rhinotermitidae, sunken pores on their legs which may produce a defensive secretion against predators (Bacchus, 1979).
The genus Coptotermes, despite its economic importance, has never been revised on a world-wide basis, and species identification in several parts of the world remains problematic. Twenty-two species of Coptotermes are known from the Indo-Malayan region (Tho, 1992), but it is uncertain how many of these species are the same as the 30 species which are reported to occur in China (Li et al., 1994). Roonwal and Chhotani (1962) revised the Indian species, and taxonomic accounts of the genus in Peninsular Malaysia and Sabah have been provided by Tho (1992) and Thapa (1981), respectively. However, recent investigations into the cuticular hydrocarbons of some Australian Coptotermes point to the existence of species complexes among morphologically indistinguishable populations (Brown et al., 1990). This is a relatively new area of taxonomic research, and similar studies on Coptotermes from other regions of the world have been very limited to date (Haverty et al., 1991) but may eventually reveal similar problems.
Six species of Coptotermes are now recognised from peninsular Malaysia, and Kirton (1995) provides an identification key which uses characteristics of the soldier caste to separate species. Coptotermes elisae, C. kalshoveni, C. travians, C. sepangensis and the C. gestroi group can be separated on the basis of the degree of mandible curvature and the width of head at the base of the mandibles. The C. gestroi group comprises two species, C. gestroi (Wasmann) and C. sp. nr gestroi, that can only be distinguished on the basis of differences in the alates. Kirton's (1995) study should be consulted when South-East Asian species are to be identified.
DistributionTop of page
Coptotermes has a broad pan-tropical distribution; in China over 30 species of this genus have been described.
Distribution TableTop of page
|Country||Distribution||Last Reported||Origin||First Reported||Invasive||References||Notes|
|China||Present||Li, 1991; Li et al., 1994|
|-Hong Kong||Present||Gao & Lam, 1985|
|-Sumatra||Present||Marian et al., 1992|
|Japan||Present||Tokora et al., 1989|
|Malaysia||Present||Tho et al., 1992|
|-Peninsular Malaysia||Present||Tho et al., 1992|
|Taiwan||Present||Wu et al., 1991|
|Vietnam||Present||Lõm-Binh-Loi & Durand, 1971|
|Africa South of Sahara||Present||Jones DT- Natural History Museum UK personal commu|
|Zimbabwe||Present||Coaton & Sheasby, 1976|
|-Hawaii||Present||Leong et al., 1983|
|Papua New Guinea||Present||Rokova & Konabe, 1990|
Growth StagesTop of page
Flowering stage, Fruiting stage, Vegetative growing stage
SymptomsTop of page
Damage by Coptotermes most frequently occurs in mature trees, although it can occur at earlier stages of growth. Coptotermes usually invade trees via the soil and bore into the tree through the roots. Sometimes the attack is secondary, after the tree has been damaged by fire or fungal infection (Cowie et al., 1989). Once a tree is infested, Coptotermes often hollow out or 'pipe' the heartwood of the trunk; although in most cases this may not be fatal to the tree, it does greatly reduce the value of the timber (Chan, 1983; Harris, 1971; Greaves et al., 1967). The most severely damaged trees can be so weakened that they are prone to be blown over by strong winds (Dhanarajan, 1969).
Attack can be above ground or at ground level. A common place for termite damage to trees is about 23 cm below ground at the fork of the tap root. On contact with a root, termites tunnel through it and eat into cambium and sapwood and then into the stem. In dry conditions they may be attracted by plant sap for moisture. Trees are easily blown down where roots have been destroyed. On oil palm and coconut, termites can feed just under the bark or under leaf bases. Large cavities can be also eaten out of trees.
Symptoms ListTop of page
Biology and EcologyTop of page
Coptotermes occur in all tropical biogeographical regions of the world. At the latitudinal limits of its range, Coptotermes is present in some temperate regions such as southern Japan and New Zealand. Coptotermes colonies can produce functional replacement reproductives (called neotenics) and thus rapidly exploit available food resources by establishing satellite colonies (Gay, 1955; Lenz and Barrett, 1982). This ability also allows fragmented colonies to thrive in the absence of the primary founding queen, and this is seen as possibly the most important factor in the success of Coptotermes in colonization of new areas when introduced by the activities of man (Lenz et al., 1988). Species of Coptotermes have been introduced into many parts of the world, including continental USA, New Zealand and numerous sites in the Pacific Ocean including Fiji, Hawaii, Guam and the Marshall Islands (Gay, 1967). In particular, C. formosanus is now established in many countries and has become a serious pest.
Coptotermes are wood-feeding termites that can attack both living and dead wood. The genus is notorious for its habit of colonizing living trees and hollowing out the heartwood to the extent that the trunk can be 'piped' and replaced with nest material and soil, without the tree showing external signs of their presence. The Oriental species produce subterranean nests but have the ability to construct covered runways and can forage away from their central nesting site (Kirton, 1995). When colonizing dead wood, Coptotermes show a significant preference for tree stumps and logs rather than smaller items of dead wood (Kirton, 1995).
In Peninsular Malaysia, species of Coptotermes are widespread in lowland forests, up to an altitude of approximately 1350 m (Tho, 1992). The studies of Kirton (1995) in Peninsular Malaysia revealed that Coptotermes generally reach their maximum reproductive success and colony growth in coastal regions, rather than inland forests. In particular, Coptotermes thrives in peat swamp forest, and is frequently found attacking Casuarina equisetifolia in beach strand forest, and is closely associated with Rhizophora and Bruguiera in mangroves (Kirton, 1995).
Swarming of alates (reproductives) is often at dusk when temperatures are lower and humidity is high. Flights often occur after rains; however, there may be up to six of these flights per year. After flying, the alates pair off and burrow into holes/cracks in wood. Eggs are produced within 5-10 days (Huang and Chen, 1984). C. formosanus can, once a colony is established, produce 1000 eggs per day (King and Spink, 1974). If conditions are ideal (e.g. optimum temperature for hatching is 30°C; Huang and Jung, 1980) eggs hatch after a month, the first brood being composed of one to two dozen individuals.
C. curvignathus has six larval stages. Workers and pre-soldiers are differentiated at the fourth instar. First and second instars are reared by adults and immature workers. A colony can survive for 50 years. Coptotermes can form supplementary reproductives (neotenics) so that isolation of a colony can give rise to a new colony. Lenz et al. (1986) in Australia found that three out of five colonies which had lost their reproductives could re-establish from neotenics in 1 year. Reproductives may also migrate with a colony if the colony moves (Miller, 1994).
C. formosanus can forage to a distance of 50 m from the main nest, and produce satellite nests at 30 m. During hot periods the termites move deeper into the soil or wood. In Java, C. curvignathus favours damper areas than C. havilandi.
Notes on Natural EnemiesTop of page
Fungal pathogens have been examined as a means of control. Sajap and Kaur (1990) looked at the histopathology of Metarhizium anisopliae on C. curvignathus in Malaysia. Metarhizium treatment has received much interest for the contol of termites (and white grubs) in Australia. An isolate of Beauveria bassiana from Isoptera was shown to be effective against C. formosanus (Wells et al., 1995). Pearce (1987) suggested that a fungal pathogen, Antennopsis gayi, tended to keep Coptotermes numbers down in Dumoga Bone National Park, Sulawesi, Indonesia. Nematodes also have been put forward as a possibility for biological control (Wu et al., 1991).
ImpactTop of page
The greatest losses to the value of timber usually occur in plantation forestry systems where fast-growing exotic tree species are planted. It is these exotic species which prove to be most vulnerable to severe termite attack. Generally, indigenous trees in natural forests are rarely badly damaged by termite attack, presumably because they have evolved defence mechanisms against indigenous termite species (Harris, 1955; Lee and Wood, 1971). There are some exceptions to this rule, the most relevant being the attack by Coptotermes acinaciformis and C. frenchi on Eucalyptus trees in Australia, which are responsible for up to 92% of the pre-harvest damage to trees in virgin forest, and 64% in younger managed forests (Greaves et al., 1967). In both natural and managed forests, it is usually those trees which are stressed (for example by water deficit, fire damage, or attack by other pests) that are most susceptible to attack from termites.
An overview of the impact of termites on the forestry industry, and methods of control in the Indo-Malaysian and African regions, is provided by Cowie et al. (1989). The most serious damage to mature forestry trees is from Coptotermes, especially in South-East Asia and Australia. The economic importance of Coptotermes in Peninsular Malaysia has been reviewed by Tho and Kirton (1990). In some exotic plantations of Araucaria and Pinus in Malaysia, up to 100% of trees can be attacked by Coptotermes species (especially C. elisae) and can become a major limitation to re-afforestation schemes (Dhanarajan, 1969; Tho, 1974). In urban areas and rural settlements, Coptotermes are the main cause of damage to wood-based building materials in Malaysia (Kirton, 1995).
Under natural conditions in the Malaysian rain forest Coptotermes may be rare. On clearing trees, other species of termite are killed and waste vegetation becomes an ideal food for Coptotermes. Newly planted crop or tree seedlings are not attacked until this food supply runs out. Termite attack can often reduce the value of timber not only by the direct effect of the termites themselves, but also by allowing the entry of other pests and pathogens.
Although Coptotermes mainly attack trees, they sometimes damage crops as well. For example, C. formosanus has been reported as damaging groundnuts and other food crops in China and Japan (Sands, 1973). Seasonal changes in foraging groups can affect the amount of damage that occurs. Attacks on healthy young rubber trees in Malaysia by C. curvignathus can occur within 3-4 weeks.
On some trees (e.g. oil palm) termites can feed just under the bark or under leaf bases, as in coconut. Large cavities can be also eaten out of trees. In Indonesia, C. curvignathus enters wounds and damage tends to be greater in older plantations. In Malaysia, Coptotermes spp. are more abundant in outlying habitats, paticularly Avicinia mangrove swamps. These places act as reservoirs for re-infestation (Salik and Tho, 1984).
Detection and InspectionTop of page
The presence of termites may be indicated by earth-covered runways or tubes found on the external surfaces of trees. In young plants, chlorosis or wilting on hot days may indicate damage. Young plants should be uprooted and the roots examined for damage. Examining the bark and trunk of trees for damage is also important. A knife can be pressed against the trunk to see if tunnelling has occurred. Coptotermes runways are often soil-lined and flattened in appearance, running with the grain of the wood.
Similarities to Other Species/ConditionsTop of page
Descriptions to distinguish between two of the major pest species, C. curvignathus and C. formosanus, are given by Coaton and Sheasby (1976). Thapa (1981) has separated soldiers of four species, C. curvignathus, C. sepangensis, C. borneensis and C. kalshoveni, by using head measurements.
Prevention and ControlTop of page
A good, up-to-date summary of insecticides used for termite control is given by Wiseman and Eggleton (1994), while Logan et al. (1990) provide a thorough review of non-chemical methods of control.
It has been suggested by Cowie et al. (1989) that there are only two effective and economically viable methods of controlling Coptotermes in the forestry industry. The first is by insecticide injection into nests within affected trunks (for details see Greaves et al., 1967; Hadlington, 1987). However, this method requires a skilled labour force to ensure proper insecticide application. The second method has been tested in Papua New Guinea, where most Coptotermes nests are located in stumps or logs. This technique involves a combination of nest destruction with explosives prior to the establishment of the plantation, followed by the destruction of queens in subsequently located nests in order to reduce reinfestation (Gray and Butcher, 1969).
Current termite control methods rely largely on the use of persistent organochlorine (cyclodine) insecticides. These are increasingly less readily available, and severe restrictions are being placed on their use, so the requirement for alternative strategies is becoming acute. At present, control of Coptotermes in mature trees is rarely economical or practical, but control in the nursery (which can also protect for the first few years after planting out) can be effective and affordable (Cowie et al., 1989).
The use of wood ash heaped around the base of tree trunks, or mixed into seedling bedding soil, is reputed to reduce termite attack. It is a common practice in some parts of the world and deserves scientific evaluation (Logan et al., 1990).
Grace and Yamamoto (1994) showed that sodium borate was effective in the laboratory but had poor penetration in the field for Douglas fir. It can also be phytotoxic to some plants.
Baiting with a chitin inhibitor, hexaflumuron, (Su, 1994) is used for treatment of houses against Coptotermes in Florida. Some success has also been found using this bait in citrus plantations. Grace et al. (1992) showed that another insect growth inhibitor, silafluofen, was effective against C. formosanus. Fenitrothion in baits or sprays of microcapsules (Iwata et al., 1989) is also a recent candidate for C. formosanus control; the insecticide is circulated throughout the colony by the characteristic grooming behaviour of the termites.
Where termites are well-established, insecticide treatment may be only a temporary measure (Mariau et al., 1992). Organochlorine emulsions less than 0.1% strong could prevent infection for a short period (Jayarathnam, 1968). Granules of chlorpyrifos and isofenfos are also commonly used (Tshuma, 1988). In China chlorpyrifos, deltamethrin and cypermethrin have gained popularity. Chlorpyrifos and phoxim showed very high contact toxicity (Tsunoda, 1991; Akhtar and Saleem, 1993). Also permethrin, fenvalerate and carbaryl performed well inside wood at low concentrations. Copper chrome arsenic is still used in some countries (Said et al., 1982); arsenic trioxide is used as a dust applied at the gallery exits (Li et al., 1994).
A recent but more costly method is to use slow-release insecticides. This is especially useful for seedlings. Chlorpyrifos in a thermoplastic matrix has been shown to be useful for tree protection (O'Hanlon, 1986). Mitchell (1989) examined the use of non-persistent slow-release granules in the forests of Zimbabwe.
Grace and Yates (1992) tested a formulation of neem which showed that C. formosanus fed less and avoided long-term contact with treated timber.
Alternative non-chemical methods of control have been shown significantly to reduce termite damage, although they can never eliminate termite attack completely. Most of these recommended silvicultural practices (reviewed by Wardell, 1987) concern the use of tree species appropriate to the local climatic and environmental conditions; the use of healthy and vigorous planting stock; adequate watering of nursery stock immediately prior to planting out; and the scheduling of planting out to avoid subjecting newly transplanted seedlings to drought (see Harris, 1971; Sen-Sarma, 1986; Cowie et al., 1989).
A review of non-chemical methods of control is given by Logan et al. (1990). The fungal pathogens Metarhizium anisopliae, Beauveria bassiana and Antennopsis gayi, and a number of nematode species, have been examined as means of biological control (see Natural Enemies). Logan et al. (1989) suggest that methods of biological control of termites show little promise of success because of the termites' social structure and behaviour. Although predators of termites can remove large numbers of individuals, these losses are unlikely to reduce the overall termite pest population to economically acceptable levels. Similarly, pathogens or parasites are not likely to be effective due to the termites' behaviour of isolating dead or infected colony members in walled-off blind chambers in the nest (Wood and Sands, 1978).
Nematodes have been advocated as a possible means of preventing subterranean termite damage to buildings (Weidner, 1983) and marketed as such in the USA. However, it has also been argued strongly that rigorous field trials have failed to demonstrate convincingly the efficacy of nematodes in termite control (Mix, 1985, 1986).
Tokoro et al. (1989, 1992) identified a trail pheromone and precursors for C.formosanus. Zhong and Kuang (1979) have synthesized 4-phenyl-cis-3-buten-1-ol from Coptotermes trail pheromone which showed good field activity against C. formosanus in China.
In New Guinea the turning off of electric lights at swarming times to discourage alate attraction has been used. Coptotermes is known to be attracted to a wavelength of 400-420 nm.
Physical barriers (e.g. basalt particles, sand, coral, etc.) have been used with some success in buildings but have only limited use in forestry and agriculture (Tamashiro et al., 1987). The particle size has to be too large and heavy for the termites to carry away, yet small enough to stop them making continuous passages in them.
Gray and Buchter (1969) found the most effective means of nest destruction without harmful environmental effects was by using explosives and killing the queens.
C. curvignathus: heavy hardwoods resistant to C. curvignathus include Neobalanocarpus heimii and Vatica sp. (Said et al., 1982), Intsia palembanica, Itomalium foetidum, Dracontemelon dao (Rokova and Konabe, 1990), Eusideroxylon zwageri, Intsia bijuga, Castaopsis argentea, Dalbergia latifolia, Hopea ferruginea, Lagerstroemia speciosa, Tectona grandis (Supriana, 1988) and Albizia procera (Supriana and Howse, 1982). Resistant compounds have been isolated from kayawood (Torreya nucifera) (Ikeda et al., 1978).
C. formosanus: resistant trees include Chinamadhuca hainanensis, Litchi chinensis, Vatica astrotricha, Tectona grandis, Robinia pseudacaria, Alseodaphne hamanensis, Homalium hainanense, Hopea hainanensis (Dai et al., 1985), Melia azedarach and Adina racemosa (Yaga, 1978).
ReferencesTop of page
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- = Present, no further details
- = Evidence of pathogen
- = Widespread
- = Last reported
- = Localised
- = Presence unconfirmed
- = Confined and subject to quarantine
- = See regional map for distribution within the country
- = Occasional or few reports