Termitidae

Genera in this family are referred to as the higher termites. This division is based on their lack ofdependence on symbiotic protozoa for cellulose digestion. Over 80% of the genera and 74% of all the species of termites are placed in this family. Itincludes genera and species in all the zoogeographic regions with a wide range of habits, including species that are economically important.

Amitermes

This is a large genus with species occurring in mostparts of the tropics and subtropics, butalso in some arid areas. The soldiers have rounded or pear-shaped heads with curved mandibles (Fig. 10.1a), which have a median tooth. Antennae have 13-17 segments. The alates are about 12 mm with the wings, and the body is dark brown to nearly black. They usually attack wood where it contacts the soil and often limit their feeding to wood buried in the ground. Amitermes spp. girdle and kill young citrus trees. Galleries are lined with brown fecal plaster, even in such objects as dry cattle dung (cow-chips). Swarming usually occurs during the day; some emerge at midday and others in the afternoon; in arid regions the flights may be at dusk and following rainfall. Many are mound-building termites, and some species are well-known for their nests. A. meridionalis constructs the so-called compass mounds in North Australia. These are wedge-shaped earthen mounds that are about 3.5 m high, and have their long axis always aligned north to south. Species in arid environments include A. desertorum in Algeria, A. vilis in Saudi Arabia, A. capito in Western Australia, and A. wheeleri in southwestern USA and Mexico.

Amitermes coachellae Alates are about 10 mm long with the wings, and the body is dark brown to nearly black. In the soldier, the mandibles are slender, and the tooth is near the base; the right and left mandibles are nearly identical. Swarming is during the day and usually following rainfall. This species is known from California, Arizona, and Nevada.

Amitermes emersoni Alates are about 12 mm long with the wings, and the body is dark brown to nearly black. The soldier is yellowish brown, the mandibles are long and slender, and the tooth is near the tip. Swarming occurs during the day in late January or early February. This species is known only from the Coachella valley in California, where it is one of the most common termites.

Florida dark-winged, subterranean termite, Amitermes floridensis Alates are 8.4-9.7 mm long with the wings; the wings are about 7.4 mm long; the body is about 4.3 mm long and dark brown to nearly black; the antennae and legs are pale to yellowish brown. Wings are brown, and the anterior veins dark brown. The soldier is yellowish brown; the head is longer than wide and light brown; the mandibles are blackish brown, long, slender, and curved. Workers are characterized by dark gut contents, which are visible through the abdominal wall. There is a single tooth on the inside of the mandibles. Swarming flights occur during the day, from 07:30-19:00 h, and from June to early August. Flights frequently occur after rainfall, and adults may begin to fly before the rain ceases. Nests are below ground, and above-ground foraging tubes are not constructed. This species occurs in and around the city of St. Petersburg, Pinellas county, Florida.

Amitermes herbertensis Alates are produced in the colony from March to July. Natural nest sites include rotten logs and tree stumps in rain forest habitats. In the urban environment, it damages structural timber, the foundation piles of houses, fence posts, electrical transmission poles, and other timber in contact with the soil. The interior of round logs, trees, or structural lumber may be destroyed and replaced with earthen material, which contains galleries. House support beams and posts and the stems of woody shrubs may be covered with an earthen sheath. This species occurs primarily in northern Queensland, Australia.

Amitermes minimus Alates are 7.5-8.5 mm long with the wings, and the body is dark brown to nearly black. In the soldier, the mandibles are slender, and the tooth is near the base. The rightand leftmandibles are notidentical, and the tooth on one may be large. Swarming is during the day. This species occurs in the desert region of southwestern USA, ranging from Texas and Arizona to Nevada and California. Damage is caused to poles and fence posts; it penetrates wood extensively and may construct hard, sand-covered galleries over the buried portion of the infested wood.

Amitermes snyderi Alates are about 9.5 mm long with the wings, and the body is dark brown. In the soldier, the mandibles are long and broad at the base, and the tooth is near the base; the right and left mandibles are nearly identical. Swarming is during the day. This species occurs in southwestern California in the Mojave and Colorado deserts. Damage is caused to redwood posts, pine boards partly buried in the soil, and the roots of desert trees.

Amitermes wheeleri Alates are 10-11.5 mm long with the wings, and the body is dark brown. The fontanelle is slit-like. The soldier is yellowish brown; the mandibles are strongly recurved and have a cone-like tooth on the basal half. Mandibles are abouthalfas long as the width of the head. Swarming occurs in Arizona in late afternoon and at night in July, and they may be attracted to lights. In Texas, winged adults have been found in May to June and October. Flights occur in Arizona and Texas after rainfall in the afternoon. These termites are strong flyers. This is one of the most widespread and common Amitermes in the USA. Damage is caused to woodwork in buildings, windmill towers, fence posts, and utility poles. Itis distributed from Mexico, southwestern Texas and Arizona to Nevada and California. It nests in dead trees and tree stumps, and sometimes in cactus and Agave, and in dry cattle dung (cow-chips).

Other Amitermes There are several other species of this genus in southwestern and western USA. A. silvestrianus,

A. pallidus, and A. parvulus have soldiers in which the mandibular tooth is located about the midpoint of the curved mandible, and the alates are about 9.5 mm long with the wings. The alates of these three species are dark brown. A. silvestrianus and A. parvulus occur in California, and A. parvulus occurs in Arizona and Texas. The alates of A. pallidus are yellowish brown. This species is known from southern Arizona, and winged adults were found at lights from June to August. A. dentosus occurs in Western Australia through to Queensland, Australia. It has small colonies in underground galleries, and it damages wood close to or contacting the ground.

Anoplotermes

Alates in this genus have a broad head, which is usually covered with fine setae, and they have small eyes and small ocelli. Antennae have 15 segments. Colonies of these species have no soldiers, and species identification is difficult. In the alates the mandibles are not much longer than broad. No soldier has been associated with colonies of this genus. Workers are dark gray and have an elongate, bag-like abdomen.

Anoplotermes fumosus Alates are about 7.5 mm long with the wings. The head is dark brown; the pronotum and abdominal segments are brown; legs are pale yellow. Wings are gray, and about twice as long as the body. No fontanelle is visible, and the ocelli are large. Antennae are as long as the head, and there are more than 14 segments; segments 3 and 4 are short. The body is covered with dense setae; setae on the head are dark brown, on the pronotum grayish, and on the abdomen yellowish brown. Colonies are in the ground in burrows, under buried logs, and in and under dry cattle dung (cow-chips). Swarming occurs in August. This species occurs in southwestern USA, in southwestern Texas, New Mexico, and northern Mexico.

Globitermes

The few known species in this genus are found in southeastern Asia. Nests are small mounds, with parts above and below ground. They feed primarily on decayed wood. As with other species that occur primarily in forested or undisturbed areas, their pest status is limited to occasional infestations of rural houses and other isolated buildings.

Globitermes sulphureus Alates are present in the colony during the rainy season, and swarming flights occur in January and May. Nests are common in forested areas and colonies are usually large. The above-ground portion of the nest is hardened soil and may be 80 cm high and 60 cm diameter at the base; the covering is about 1 cm thick and apparently made of a mix of soil and saliva. Internally, the nest is divided into small chambers with thick walls. The central space is filled with a mass of carton, which holds the primary reproductives, the eggs, and developing larvae. The soldiers are active and readily attack other termites and ants. Intruders are killed with the large mandibles of the soldiers and covered with a yellowish-white salivary secretion. This species is distributed in Thailand, Myanmar, Vietnam, and Malaysia. It is a structural pest of houses in rural Thailand.

Gnathamitermes

The long, and relatively straight mandibles of the soldier caste distinguish this genus. Mandibles are almost as long as the head is wide, and curve inward only at the tip. Alates are 13-16 mm long with the wings, and the body is dark brown. Members of this genus are primarily soil-dwelling, but they build extensive galleries above ground. Dry cattle dung (cow-chips), weathered vegetation, range grasses, posts, and other wooden structures are damaged. Most of the damage is the result of their galleries above ground, rather than actual consumption ofmaterials.

Gnathamitermesperplexus Alates are about 16 mm long, with the wings, and the body is brown to dark brown. Soldiers are variable and have considerable variability in the teeth on the mandibles; the teeth project from the face of the mandible and are located at about the midpoint. Often the teeth are not equal in size. Workers have a yellowish-brown head, and are slightly larger than the apterous nymphs. Alates develop from egg to adult in about 8 months. In California, soil tubes were built on the lower trunk of young trees, and the bark and wood were damaged. Swarming in Texas occurs in May to August, and it is usually in early morning after rain the previous night. In Arizona, flights take place in late June and July in the afternoon or evening during rainfall.

Gnathamitermes tubiformans Alates are about 9 mm long with the wings; the body is brown and the legs are yellowish brown. Wings are nearly twice as long as the body, and brownish yellow, costal veins are dark brown. Antennae are longer than the head and 14-segmented. The body and legs have dense, short setae. Soldiers have mandibles that are not as long as the width of the head; the length of the head plus mandibles is about 2.4 mm. The abdomen is gray, the legs are yellowish white, and the mandibles are reddish brown; antennae are brown at the base and yellowish brown from the middle to the apex. The mandibular tooth is located on the apical portion of the mandible, and does not project from the mandible, but appears to be cut away from the bottom. Colonies are in the ground. Workers actively forage at night, usually after rainfall; earthen galleries may be in low vegetation and grass. Soil tubes are built 10-15 cm above ground and are usually attached to grass or bushes. Beneath the tubes are cells in the soil. Colonies may be very large and located deep in soil. Supplementary reproductive forms, with short wing pads and dark coloring, may head colonies. Swarming occurs during the day after rainfall in spring and summer. Damage is done to grass in grazing land, and truck crops. This species occurs in western Texas and possibly New Mexico.

Macrotermes

This genus is widespread in Africa and southern and southeastern Asia. The soldiers and queens of several species are very large. These termites are mound builders and fungus growers. Their peststatus in the urban environmentis limited; they damage trees and young plants, but some are destructive to woodwork near the ground. M. barneyi occurs in central and southern China. The subterranean nest is compact and consists ofa system ofinterconnecting chambers, which contain the fungus combs. This species is a structural pest in some regions. M. gilvus is a large species which is the most common mound-building termite ofsoutheastern Asia, from Myanmar and Indonesia to the Philippines. The swarming of alates of this species in the Philippines occurs in the evening from May to September. The swarms are very large and composed mostly ofmales. This species, along with M. pakistanicus and M. anadi, occasionally infests houses in rural Thailand.

Microcerotermes

This genus has more than 100 species and it occurs in most zoogeographical regions. Species are widely distributed in the tropics, neotropics, and in the dry steppe and desert fringes. They have subterranean or small-mound nests and the colonies are moderate-sized. Species in this genus build compact nests of hard carton with a honeycomb structure. Workers forage in below- and above-ground galleries made of tough carton material; they also forage within plant stems. Dead wood from trees such as Acacia, Zizyphus, and Tamarix is used, as well as plants growing beyond the limits of tree growth. The soldiers of this genus have long rectangular heads (Fig. io.ih), which are dark brown. Their long mandibles have inward-curved tips, and the inner margins are serrated; the antennae have i2-i5 segments.

Most species in this genus eat decayed or weathered wood. Some attack structural wood and have pest status. In Australia pest species include Microcerotermes distinctus and M. implicadus, which are widespread in the country. M. nervosus and M. boreus are restricted to Western Australia and the Northern Territory, and damage farm buildings. M. biori (Fig. io.ih) is common on a large number of the islands of Melanesia and Central Polynesia. In the dry steppe and desert fringes, the common Microcerotermes species include M. palearcticus in Morocco and Algeria, M. eugnathus in Tunisia, M. palestinensis in Israel south of Tel Aviv, M. diversus from southern Iraq and adjacent Iran to the Arabian peninsula, and M. gabrielis in northern Iraq, Iran, and Afghanistan. Several species damage structural wood in southern Israel and southern Iraq, including M. diversus. M. exiguus and M. arboreus are structural pests in urban areas of Brazil and the West Indies. M. strunckii is a structural pest and damages paper products in northeastern Argentina. M. crassus is a very common pest of houses in rural areas of Thailand.

Microcerotermes turneri Alates are present in the nest from September to January. Soldiers are about 5.2 mm long. Nests are usually built on the ground, on the trunks of living or dead trees, on the tops of stumps or fence posts, or occasionally on the timber framing of buildings. The arboreal nests are rounded, and are 22-46 cm in diameter and 5 m above the ground. The arboreal nests are light brown, and not black. This species occurs north of PortMacquarie on the central coast of New South Wales and into coastal Queensland, Australia.

Nasutitermes

Soldiers in this genus are usually smaller than the workers, and they have a large rounded head withouta constriction, and there is a pointed snout (nasute) (Fig. io.ii). The mandibles are small, the antennae are short, and the ocelli are small. Most species feed on grass, some infest decaying wood, and some attack structural timbers. The mound-building habit is pronounced in some species, but others build arboreal nests, and some are subterranean. Species thattypically build arboreal nests may also infest structures, including boats. Nasutitermes is a widely distributed genus, with many species occurring in Australia, and Central and South America. N.grandinasus occurs in southeastern China, and it damages woodwork in houses that are within the foraging range of nests.

Nasutitermes corniger The alate body is about 7 mm long. The head and body are reddish brown; the mouthparts are yellowish brown. The head is broader than long and the eyes are small; the ocelli are rounded. The soldier head is i.5 mm and dark brown. The thorax and dorsum of the abdomen are yellowish brown, while the legs are pale yellow. The worker head is dark brown, and the right mandible has three teeth. Nests may be arboreal and composed of brownish-black carton material; they are usually elongate and rounded. This species occurs in Central and South America, including Argentina and Brazil. In the USA, it is known from Texas and Florida.

Nasutitermes costalis Alates are present in the nest in spring (May), and swarming flights occur during the day, and sometimes during rainfall. Natural nest sites include dead wood in living trees. In the urban environment, nests may be in fences, and in the structural wood of buildings, and they nest around the base of residential trees. Ground nests may be mounds about 1 m diameter and are sometimes referred to as comejeneras. In Trinidad, this is the mostabundanttermite species. Itis widespread in the Neotropics, occurring from Cuba to Bolivia, Puerto Rico and northern Brazil. It has been introduced into North America, and may be established as a structural pest in the region of Dania Beach, Florida.

Nasutitermes exitiosus Soldiers are about 4.2 mm long. This is a mound-building species; the mounds are 30-70 cm high and about 1 m diameter. In dry areas this species nests in tree stumps or below ground and often the soil is bare above the nest site. Colonies are usually established alongside a log, tree stump, or post, and it may take 20 years for a colony to reach mature size. Colonies usually have a single queen, and when mature the queen is often very large and capable of laying 2500-3000 eggs per day. Swarming occurs at night from September to January, but mostly in October and November. Colonies can produce 50 000 alates in a single season; they are attracted to lightatnight. Damage may be to fence posts, poles, structural timbers in bridges and buildings, and subflooring of houses when a nest beneath the house is undetected. It is not usually a pest in urban areas, because the nest mound is conspicuous. In rural areas, structures may be attacked, and sometimes froma nestthatis located 45 maway. Wood attacked by this termite has large galleries and a rough surface. Galleries in wood are often filled with blackish-brown woody carton, and packed with soil. This species occurs in most areas of southern Australia, generally south of the New South Wales and Queensland border.

Nasutitermes fumigatus, N. dixoni Soldiers ofN.fumigatus are about 3.5 mm long; soldiers of N. dixoni are about 3.8 mm long. These are subterranean species, and they attack decaying wood that is in contact with the ground. House infestations occur when there is an existing problem with decayed wood in flooring timbers, usually due to inadequate ventilation. These termites are secondary invaders in the decayed wood, and they do not infest sound wood. These two species occur along coastal Australia, from central Queensland to the Victoria and South Australia border. A related species, N. occasus,has similar habits; it occurs in southwestern Australia.

Nasutitermes longipennis Soldiers are about 4 mm long, and they have a reddish-brown head. This species constructs mounds similar to those of N. exitiosus, or the nests may be subterranean. It damages wood in contact with the soil, but it does not feed on sound, dry wood. This species occurs along coastal Queensland to Northern Territory, Australia.

Nasutitermes walkeri Soldiers are about 6 mm long, and these are among the largest in the genus. Colonies are usually started in the base or root crown of trees where there is some decay or basal fire damage. Years after a bush fire, the nests of this termite may be abundant in a comparatively small area. When colonies are well established at the base of the tree, an arboreal nestmade of soil and carton material is constructed higher up in the tree. Damage is usually to fences, poles, and wood in the ground; attack of buildings is not common and usually is associated with decaying wood and high moisture. This species occurs in the coastal bushland of Sydney, Australia and further north. A related species, N. graveolus, has similar habits; the soldiers are about 4.3 mm long. It occurs in coastal northern Australia, from Townsville to Darwin.

Other Nasutitermes There are several species in this large genus that occur in urban areas and damage structures. N. aquilinus and N. ehrhardti are have been introduced into Brazil and Argentina, and often cause structural damage. N. bivalens constructs subterranean nests and is a pest in urban areas of eastern Brazil. N. nigriceps is widely distributed in Central and South America, Cayman Islands, and Jamaica, but it is known to infest boats and to be transported to other regions. N. mexi-canus is common along the west coast ofMexico (Colima); nests are in the ground, and posts and other structures are attacked.

Odontotermes

This large genus contains some of the most common termite species in Asia. Many of the species are mound-builders; they forage above ground in protected passageways. Soldiers in this genus have a marginal tooth on the left mandible (Fig. 10.2b), and sometimes on the right. The subterranean nests of some species, such as Odontotermes bogoriensis, are 3-4 m diameter and about 50 cm below ground. Above-ground nests may be 1 m high and capped with rounded domes. The pest status of the species in the urban environmentis based oninfestations of structural wood and the damage to banks of mountain ponds and reservoirs. People of the north Karnatak region of India have used soil from the mounds of O. assmuthi for plastering walls and making household cooking ovens. Analysis of the soil shows the presence of binding elements such as silica, iron, aluminum, calcium, and magnesium.

odontotermes feae Alates are present in nests and swarming occurs in June, and September to November. Flights are usually after rainfall and begin after sunset, and they may last several hours or through the night. Alates emerge singly through small circular holes in the ground; workers are present around the holes, which they close when swarming concludes. Nests are large underground structures, consisting of interconnecting chambers that may cover an area of 1.5-2.5 m diameter. The fungus garden is usually in the center of the nest, and may be 75 cm below the surface. Foraging workers usually travel in rows in earthen tunnels and above-ground tubes. This species attacks dead wood, and structural timber in buildings. Itis one of the most common termites damaging buildings in India, and frequently infests houses in rural Thailand. Itis distributed throughoutIndia, Sri Lanka, Myanmar, Thailand, and Vietnam.

odontotermes redemanni Alates are present in nests and swarming occurs in September in eastern India, and in November and December in Sri Lanka. It builds a very large earthen mound that may be 2 m above ground, and 2-3 m diameter; the mound takes 10-12 years to develop fully. Inside and well below ground, there are large chambers that contain fungus combs. Nests usually have a large central chamber that contains a fungus and serves as an egg deposition site. Colonies are large and often contain several functional queens and kings. Foraging workers make covered passageways above ground. This species is common in Sri Lanka and damages woodwork in buildings close to nest sites. Distribution includes Sri Lanka and India.

Other odontotermes Many of the subterranean species of this genus attack dead wood and infest buildings that are close to nest sites. O. ceylonicus is a structural pest in Sri Lanka. It often occurs in the nests of other termites, including O. horni, O. rede-manni, and especially Hypotermes obscuriceps. The subterranean species, O. formosanus, attacks living trees and structural timber. It forages above ground under earthen passageways. This large species occurs in southeastern Asia, including eastern India, Myanmar, Thailand, and Vietnam. It is also a pest in southern China and Taiwan. O. proformosanus and O. longignathus infesthouses in rural areas ofThailand. O. wallonensis builds low mounds which often have 15-25-cm-high chimneys extending above the surface. It is widespread in India, where it attacks structural wood in buildings.

Schedorhinotermes

Species in this genus have a dimorphic soldier caste, and the minor (small) soldiers have mandibles with distinct teeth (Fig. io.2d). Species occur in Africa, Asia, and Australia. All of the Australian members of the genus infest wood, and form small- to moderate-sized colonies in wood in the ground; none are mound builders. Most species produce swarming flights at night and the alates are attracted to lights. Several species are pests of structural wood. S. intermedius has several forms, and it may be a species complex.

schedorhinotermes actuosus Alates are present in colonies from November through May. Swarming is usually atnightand there may be several flights from a colony during one season; flights are usually before or after rainfall. Natural nest sites include logs, trunks of trees, and hollow sections of eucalyptus, and in abandoned nests of other termite species. In the urban environment, this species is a pest of structural wood and buildings. This species occurs nearly throughout Australia, except Victoria.

schedorhinotermes intermedius Alates emerge from nests in the ground in November and December. Major soldiers are about 5.6 mm long, and minor soldiers are about 3.6 mm long. Colonies are usually large, and when only minor soldiers are found, it is not mature. Natural nest sites include trees, rotten logs, and tree stumps; in the urban environment, nests may be around building foundations and under patios. This species is distributed from New South Wales into southern Queensland, Australia. It is a pest of wooden buildings in the Sydney metropolitan area.

Other schedorhinotermes Species in this genus are usually structural pests wherever they occur. S. breinli, S. seclusus, and S. reticulatus occur in Australia and the winged forms are present from January through May. They are found in trees, both hardwood and softwoods, butalso attack structural timber in buildings. S. malaccensis occurs in Malaysia and parts of southern

China. It nests in tree stumps and in the wounds of trees, including palms, but it also attacks woodwork in buildings. S. medioobscurus occasionally infests houses in urban and rural areas ofThailand.

Tenuirostritermes

These termites are referred to as ant-like because of their rapid movement, dark color, and general behavior when the colony is disturbed. Soldiers in this genus have the front of their head produced into a snout (nasute). The head is longer than wide, and there is a constriction across the lower front, giving a distinct bilobed shape. The antennae are inserted ventrally on the head. The alate head is broader than long and the clypeus is large and bilobed. The colony contains small piles of vegetative material, primarily cuttings of weathered grass. Workers forage at night or on cloudy days, accompanied by an escort of soldiers.

Tenuirostritermescinereus(Fig. I0.6d) Alates are about2o mm long with the wings, and the abdomen is brown and densely covered with pale yellow setae. The head of the alate is broader than long; the eyes and the ocelli are large; ocelli are less than half their diameter from the eyes. The head of the soldier is 1.3-1.4 mm long, primarily black; the snout is reddish brown. The antennae are i2-segmented. It is found in and under dry cattle dung (cow-chips) or under stones. Colonies are deep in the ground, and include workers, soldiers, and reproductives. Workers forage at night in columns and attack plants. Swarming flights are at night during rainfall in late summer (September). Damage has been to cedar posts. This species occurs in central and southwestern Texas, primarily in irrigated areas.

Tenuirostritermes tenuirostris (Fig. I0.2g, h) Alates are 20-23 mm long with the wings; the abdomen is brown and densely covered with pale yellow setae. The head of the alate is broader than long; the eyes and the ocelli are large; ocelli are about half their diameter from the eyes. Wings are about three times longer than the abdomen, and often twisted at the tip; the wing membrane has minute, yellowish-brown setae. The head of the soldier is reddish brown, while the snout is blackish brown. The antennae are i3-segmented. Its habits are similar to those of T. cinereus. Swarming flights in Arizona occur from dusk to night during rainfall in June and July. This species occurs in Mexico, Texas, and in the mountains of Arizona.

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