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1. INTRODUCTION While the preceding three chapters were organized according to insect activity (defoliating, sap-feeding and boring), this chapter deviates from this pattern for reasons that become evident by looking at similarities between the two groups. 1.1. Similarities The general public often pools ants and termites by calling the latter “white ants” and their mounds “ant hills”. Although entomologically incorrect, a number of uncanny similarities may underlie the confusion. Both are social insects of similar size and abundance, and both are polymorphic , i.e., their colonies are composed of castes which perform different functions. Both become similarly agitated and belligerent when disturbed, and they inhabit similar places, i.e., underground nests, wood, arboreal nests or mounds. Although less obvious, both share an odd trait; at least one of the reproductive castes sheds both sets of wings at the end of the nuptial flight and continues life as a wingless adult. Given their overall biomass, it is not surprising that both occupy significant ecosystem niches, ants almost globally, termites in the warmer zones. They also share a number of enemies. Ants number among the worst antagonists of termites and of other ants. In addition, certain termites and ants cultivate fungi, although no fungus-cultivating ants are known from Tanzania. Last not least, both termites and ants include species suitable for human food. 1.2. Differences Despite these similarities, a number of important differences exist (Table 6-1). First, ants and termites represent two entirely different orders. Termites belong to the primitive Isoptera and thus have three stages of development (incomplete metamorphosis), while ants belong to the more advanced Hymenoptera with four stages (complete metamorphosis). While all Isoptera are termites, not all Hymenoptera are ants. In both, the fertile females are long-lived. However, while termite kings cohabit with
the queen for years, the fertile male ant dies shortly after mating. Ants have only female workers, whereas termite workers may be of either sex. Aside from the color (termites are mainly light-colored and ants darker), more subtle morphological differences include the antennae, the relative size of front and hind wings and their type of venation, as well as the presence or absence of a pair of small abdominal appendages (cerci). In the termites thorax and abdomen are broadly jointed, as opposed to the constricted waistline of ants. Ants and termites also differ in food habits and distribution. 1.3. Significance in Forestry From a forestry point of view, it is not easy to collectively classify either ants or termites as pests, or as beneficial and useful insects. Some species tend to be one or the other, while many others combine negative with positive aspects. In ecological terms, both termites and ants are mostly beneficial. However, a small fraction of termite species in the four families occurring in East Africa kill young trees, damage older ones and destroy wooden structures. In parts of the world, some species of ants are of concern as defoliators, girdlers or wood destroyers. Of the four families of ants of special interest to Tanzanian forestry, two include predators, mostly of termites. The other two include predaceous species that protect trees against browsers, defoliators, borers and termites, while at the same time tending honeydew-producing aphids and other sap-feeding insects, to the detriment of the same trees. 2. ANTS (HYMENOPTERA: FORMICIDAE) 2.1. General The total number of recorded ant species worldwide numbers almost 12,000, but actual numbers probably exceed 20,000 species. At this time, 1,686 species have
ANTS AND TERMITES
been described from the Afrotropics (H Robertson, pers. comm.). A recent study at Mkomazi in Tanzania, one of the best-collected sites for ants in Africa, revealed 232 species, at least one third undescribed (Robertson 1999, 2002). 2.1.1. Castes Mature ants are characterized by large heads, elbowed antennae, and constrictions of the second, or second and third abdominal segments forming a distinct waist. Typically a colony consists of one queen and a set of wingless, sterile female workers that tend a brood of eggs, and the maggot-like larvae and pupae (naked or contained in cocoons, depending on species) kept in chambers. In some species, workers called minors and majors differ in size. The majors of certain species have modified heads that help them function as soldiers. Some ants are blind, while others see very well. Only kings and virgin queens are winged for the duration of their nuptial flight. Mating takes place on the wing, the male dying shortly afterwards. The queen sheds her wings to retreat to a nest, where she remains producing eggs fertilized by stored sperm, possibly for years. 2.1.2. Habits Some ant species form small colonies, while other colonies number in the millions. Based on where ants nest and forage, Robertson (1999) differentiated four groups, i.e., arboreal, ground-dwelling, litter species and subterranean ants. With the exception of a few nomadic ants, all live in permanent structures. Their nests occupy existing, or excavated hollows in soil or wood. Some build arboreal carton nests or leaf nests, others inhabit termitaria, and some occupy preformed structures (domatia or myrmecodia) on certain plants. Symbiotic relationships exist not only with such “ant plants”, but also with numerous species of invertebrate ant affiliates (myrmecophiles) (Kohl 1909; Hölldobler and Wilson 1990; Kistner 1998). Most prominent among these are various rove beetles, ant nest beetles and lycaenid caterpillars that live in ant nests as detritus feeders, parasites or predators. They often mimic ant brood or the ants themselves, morphologically and/or chemically. Many other freeliving insects and spiders are also morphological ant mimics. Other symbiotic relationships exist with many small Homoptera such as aphids, scale insects and hoppers. Ants tend and defend these insects like cows and may even build stables for them. In return they milk them for honeydew. Occasionally, when a herd becomes too big, they may even slaughter some of their “cows”. 2.1.3. Significance In terms of distribution, diversity of social systems, ecological adaptations and numbers, ants are the most successful of social insects (Hölldobler and Wilson 1990). Their biomass, especially in certain tropical forests, exceeds that of all vertebrates combined by several factors. Ants’ impact on trees often involves beneficial as well as noxious elements. On the plus side, ants may be important seed distributors (myrmecochory) and soil modifiers. Many are carnivores and thus reduce populations
while most formicines have pupal cocoons. the petiole of adults appears two-segmented and the workers carry stings. Their maggots (Figure 6-1B) are typical for ants and their pupae are naked. .2. The reddish-brown and completely blind workers (Figure 6-1C) are polymorphic. comm. nomadic colonies. beneficial or otherwise useful representatives of greater interest to forestry in Tanzania. while those in the subgenus Dorylus are entirely subterranean and timid (Silow 1983). D. especially termites (Way and Khoo 1992). but A.. Arboreal ants also discourage browsers by defending their hosts with fierce stings. Four subfamilies include species with noxious. nigricans Ill.4. and D. The dorylines occur in huge. but spray ant acid (formic acid) instead. pers. were listed for Tanzania (LePelley 1959). the same ants may kill tree shoots and buds.1. Colonies of driver ants consist of major and minor workers. Trees also suffer direct growth impact from the sap-feeding Homoptera associated with them. 2. 11 occur in Africa (Bolton 1994).182 CHAPTER 6 of tree defoliators and other pests. the related Anomma nigricans and D.). Two of the four subfamilies discussed (Dorylinae and Ponerinae) are entirely carnivorous. 2. helvolus is less common and mainly subterranean. These nomadic. it is convenient to deal with subfamilies. honeydew supports sooty mold fungi that may interfere with leaves’ photosynthesis and gas exchange. Pupae of the myrmicines have no cocoons. respectively. Out of 16 worldwide. the ponerines in smaller. which includes two subgenera with different habits but a similar appearance. molestus Gerst. helvolus. Safari or Legionary Ants. their petiole appears to be one-segmented and their workers have no stings. 2. or indirectly from the action of secondary agents. including ant specialists.2. Sausageflies Dorylus (=Anomma) spp.1. as described by Forel (1911) and Skaife (1953. On the other hand. The two other subfamilies. Only this one genus of Dorylinae. helvolus L. Myrmicinae and Formicinae. more stable colonies. soldiers and reproductives. Ants themselves are important links in food chains. Siafu. 1979). molestus in Tanzania is missing. For instance. Description and Life History As detailed information concerning D. Two species. carnivorous ants are found from the Sahara to the Cape and through tropical Asia (Skaife 1953). Driver ants undergo a four-stage metamorphosis. appears to be the main surface-dwelling species in this country (H Robertson. is represented in the Afrotropics. Their worst enemies are other ants although there are also numerous predaceous vertebrates. Species in the subgenus Anomma are partly subterranean and vicious. bites and repulsive or irritating substances. Many Homoptera are also known to be vectors of virus and phytoplasma diseases. feature more diverse characteristics and include beneficial and damaging species. Classification Given the large number of ants. may serve as a guide. Dorylinae: Driver. D.
Figure 6-1. blind and wingless queens (Figure 6-1E). F from Pinhey 1974). medium brown insects with a fluffy head and thorax. On account of their large. As their abdomen is distended with eggs produced at the rate of 1-4 million per month.. Depending on species. making them the largest of all ants (Forel 1911. They have wingspans of up to 44 mm and are the only seeing and winged form of siafu. They sport massively developed armored heads with huge sickle-shaped mandibles. .ANTS AND TERMITES 183 i. are 29-51 mm long. (A-E from Forel 1911/12. Soldiers (Figure 6-1D) are much larger than workers. Male driver ants (Figure 6-1F) are up to 30 mm long. the dark reddish-brown. Hölldobler and Wilson 1990). to individuals of about 8 mm length. Driver ant Anomma nigricans (Dorylinae). sausage-shaped abdomen they are known as “sausageflies”. considered specialized workers. (A) Egg (B) larva (C) worker (D) soldier (E) female (F) male. ranging from midgets less than 2 mm long. queens walk with difficulty only. there may be four types.e.
During that time. The sheer number of driver ants. Footage featuring the driver ants was filmed at Mt. the males are captured and de-winged by workers before being carried underground to the queen. Driver ants are in fact able to overpower most forms of life that cannot get away from them. for the simple reason that most of their prey is composed of insects including termites and many crop pests. In the early 1900s. together with their reputation as “incomparably bloodthirsty” carnivores. with as many as 22 million workers (Hölldobler and Wilson 1990).184 CHAPTER 6 The degree of social organization by driver ants is only rivaled by that of certain bees and termites. transported to the nest and consumed there. appear in numbers at the mouth of the nest (Vosseler 1905c). However. in fact Africa’s “top predators” and “the baddest little brutes on the planet”. Despite this threatening demeanor. A recent documentary by National Geographic highlighted and inevitably glamorized the world’s three most dangerous “Killer Ants”. The former two ant species sting viciously and the bulldog ant carries the most powerful venom of any insect. feared and hated creatures are credited with being overall more beneficial than damaging. the film declared siafu THE most dangerous. When caught. the leaderless swarms of many thousands of individuals raid in 12 m wide fronts accompanied by an ominous rustle. Vosseler (1905c) actually considered them a “blessing for tropical nature”. the size of their colonies. siafu supposedly killed a leopard near Tanga (Vosseler 1905c). and are rumored to have killed people such as the sick and drunks. to seek new hunting grounds. Prey is cut up. however. they ponderously mimic stinging motions. The young and the queen are carried with them. compelled by “haste. including South America’s army ants (Eciton spp. myrmecophile rove beetles that live in the nest.). invite sensationalization. They are generally considered a threat to confined domestic animals. Also. by forcing the gaping mandibles of soldiers to clamp shut at half a centimeter apart along a cut.) and Africa’s driver ants. males depart for their nuptial flight and are frequently observed buzzing around lights. sets them apart from those of other social insects. Every 20-30 days. just like certain soldier termites. they are harmless impostors without a sting and even their sharp jaws are not used for biting. fight and slaughter” (Vosseler 1905c). sleeping children. the Tasmanian bulldog ants (Myrmecia sp. because 1-2 days before the exodus. The moving hordes attack everything in their path and leave as suddenly as they come. the whole colony moves in orderly fashion from temporary bivouacs deep below the surface at the base of a buried stone or tree stump. Much of the year. On dull days or at night. A move is somewhat predictable. Meru in Tanzania. driver ants remain underground in often deeply (1-4 m) excavated bivouacs and only emerge with the rains (Skaife 1953). Pregnant Dorobo women are said to avoid stepping over siafu raiding parties for fear of abortion (Schnee 1920). For mating. before severing the bodies from the heads . siafu soldiers have traditionally been used in folk medicine to suture wounds. Based on the extraordinarily powerful mandibles of driver ant soldiers. Despite their vicious image. chase. when food resources for several miles around the nest have been depleted. these much maligned.
Plate 45). control is not likely to be needed except in the neighborhood of settlements and camps. roasted and powdered driver ants served as starvation food in times of need (Silow 1983).ANTS AND TERMITES 185 (Silow 1983). A pangolin (Manis temincki) rolled into a defensive ball.2. After healing. As an interesting tidbit of natural history. The most important natural enemy of driver ants. Usangu. a mix of soap with petroleum poured into the exits will discourage them.2. Whenever they approach human habitation. . Given the overall beneficial role of driver ants. is the pangolin (Manis temincki) (Vosseler 1905c). The Shambaa supposedly employ a specialist (“mganga ya siafu”) to ward off ants that invade houses during the rainy season (Silow 1983). This is a specialized predator of ants and termites. O’Toole and Preston-Mafham 1985). Management. Figure 6-2. as well as other ants and termites. These muscids are observed hovering over a column of driver ants. siafu columns can be diverted with strips of wood ash (Morstatt 1910a). the heads or stitches are removed. but this animal has become rare (Figure 6-2. 2. driver ants are associated with their own house flies (Stomoxys ochrosoma Speiser). then dropping a clutch of about 20 eggs on them. selecting worker ants without booty. If a colony sets up a bivouac too close for comfort. For the Ngindo of Tanzania. This egg bomb is carried into the bivouac where the maggots hatch to live as scavengers (Vosseler 1905c.
2. They attack and sting termites in their nests. Since there were an average of 1. Nests are underground.4. Since they nest mostly at the base of large trees. subterranean ants occurring from West to South Africa (Skaife 1953). at least temporarily. Other ponerine ants are specialized predators of various arthropods and earthworms in parts of Africa and Tanzania (LePelley 1959.634 ha large Namakutwa-Nyamuete Forest Reserve. Pupae are contained in cocoons.4. After each worker has secured 5-6 termites. The queen.3. then dive in for more attacks. Hölldobler and Wilson 1990. . berthoudi.1. there were three daily hunting peaks and each colony utilized 2. In the Matumbi Hills of Rufiji District. Nyamasyo 1989). Because of their aggressive behavior and conspicuous communal leaf nests they are familiar to everyone working around trees in these areas. They are aggressive predators with a painful sting. Matabele ants raid as well-organized small armies of major and minor workers marching in columns 5-10 abreast to invade termitaria. Robertson 1999). longinoda occur across most of the forested Afrotropics.300 m2 each year (Le Page 1981).8/ha in a study in Kajiado. as opposed to only 3.000-2. range from India to Australia and the Solomon Islands. looks much like a major worker but with a stouter abdomen.) (=Megaponera foetens F.144 nests were estimated on the 4. the smaller ones are about 10 mm long and shinier (Sweeney 1976). Oecophylla longinoda (Latreille): African Weaver. drag them maimed and dying to the outside. the entrances a simple hole without mound. the average predation rate of a colony of Matabele ants was 932 termites per nest per day (Bayliss and Fielding 2002). Kenya (Le Page 1981). they are assumed to afford a high degree of protection for these trees from termites. both also predators of termites. these ants are obviously a major restraint on termites in Rufiji District. about 20 mm long. Given the nest density of 16/ha in this coastal forest. Their food consists almost exclusively of the larger species of termites (Morstatt 1922. In the Kenya study. Weidner 1988. O. they reassemble at the surface and carry off their prey to the nest. Ponerinae CHAPTER 6 Pachycondyla analis (Latr. Tailor. The larger workers are up to 17 mm long and dull black with fine yellow hairs.186 2.475 worker ants per colony and a total of 74. Colonies move occasionally (Plate 73). the number of termites taken there in August/September amounted to 69 x106 daily. or Majimoto Ants These ants are considered among the most successful social insects in the Old World tropics (Lever 1979). Most notable for East Africa are the African stink ants Pachycondyla (=Paltothyreus) tarsata and P.). Matabele ants are relatively large. Red Tree. Formicinae 2. Le Page 1981. smaragdina F. while the closely related O.
In Tanzania. usually reddishbrown. dark workers (Vanderplank 1960). usually in the sunnier part of the peripheral canopy. The former tend various Homoptera and are not aggressive. incorporates living host leaves tied together by silk into tight compartments.200 m and are particularly common in riverine forests (Vanderplank 1960). One colony builds hundreds of these in several trees. A colony may exist for as long as five years (Vanderplank 1960). about a foot or more across. In Tanzania. The bulk of a colony consists of major and minor workers. but when encounters with other ants. facing the intruder with gaping mandibles and cocking their abdomen to spray acid from a modified sting gland.000 individuals (Vanderplank 1960). including their own species occur. Each leaf nest.000 and 164. foraging .ANTS AND TERMITES 187 O. Workers see very well and employ no fewer than five recruitment systems comprised of differing chemical and tactile signals. major workers that forage for food and perform a variety of other tasks. longinoda is the dominant arboreal species of ant in the canopies of many softleaved species of wild and cultivated trees in tropical Africa. and minor workers that care for the eggs and younger nymphs (Hölldobler and Wilson 1990). Neighboring colonies are separated by unoccupied corridors (“no ant’s land”). The workers hold the larvae in their mandibles and use them as animate shuttles to spin together the edges of leaves held in place by other workers (Vanderplank 1960). some colonies consist of yellow individuals with a high proportion of small workers to large ones. derives from the sharp pain associated with their fierce bite. weaver ants behave most aggressively by emitting alarm pheromones from a mandibular gland. while others are composed of mostly large. leggy individuals with a pyriform head. Males and females usually live in separate leaf nests. while the latter do not appear to tend Homoptera and are extremely aggressive. they occur at altitudes up to 1. The total population of only part of a small colony of 192 leaf nests assessed in Zanzibar was estimated to be between 115. Individual nests are connected by odor trails laid with secretions from a rectal gland (Hölldobler and Wilson 1990). More conspicuous than the ants themselves are their leaf nests (Plate 74). Major workers are 7-11 mm long. More recent evidence suggests that in many colonies the worker force may exceed half a million (Hölldobler and Wilson 1990). to organize territorial defense. The leaves selected often harbor aphids and scale insects. Their Swahili name majimoto (hot water) ants. Weaver ant colonies can be sizeable. long and strong mandibles and long antennae (Figure 6-3). Their construction requires the cooperation of ant workers as stitchers with last instar larvae who act as the silk factory. The caste system of majimoto ants consists of males and three forms of adult females: an apparently single queen. Some nests may also contain myrmecophile lycaenid caterpillars that feed on the Homoptera with impunity. Workers defending the colony are usually housed in peripheral barracks.
To lay Figure 6-3. After heavy rains. However. especially termites. one queen may produce up to 930 eggs a day (Vanderplank 1960). nest accompanied by a mob of reproduced by permission of Elsevier GmbH. from November to May. CHAPTER 6 As researched in Tanzania. Minor workers attend scale insects to collect honeydew and regurgitate it to major workers. preceded by males by days or weeks. The arboreal majimoto ant. while the naked pupae take 3-12 days. winged females escape from their nests. They even subdue large insects such as rhinoceros beetles (Oryctes) (Figure 6-4). Workers live for 68-140 days. Oecophylla her eggs. Weaver ants can be a nuisance for people around trees and their encouragement of sapsucking Homoptera may have negative implications for host trees. Majors also function as stitchers and will carry minor workers to locations where their work is required. both on the ground and in tree crowns (Hölldobler and Wilson 1990). Depending on temperature.188 excursions and the exploration of new terrain (Hölldobler and Wilson 1990). The total period from egg to adult amounts to 18-50 days. (From Schmutterer 1969. Majimoto ants. The maggot-like larvae develop in 9-25 days. they compensate to a great extent for this indirect damage by preying on a wide range of pest insects. Figure 6-4. subduing a rhinoceros beetle (Oryctes monoceros). Oecophylla longinoda (Formicinae). the eggs incubate for about 1-2 weeks. Jena). she moves from nest to longinoda (Formicinae). who transport it into the brood nest for final consumption. . protective workers acting as soldiers.
Ants building carton nests often kill the leaders of the best trees in plantations of Cupressus (Plate 46). Most conspicuous and interesting are cocktail ants that build carton nests in trees and others inhabiting domatia in stipules. most often football-sized black structures (Figure 6-5) made of chewed vegetable fibers. Interference from several antagonistic species of ants constitutes a major complication in the management of Oecophylla ants (Way 1951. 2. Most are arboreal insects living under bark. hollow branches or in conspicuous carton nests of variable size attached to tree branches.ANTS AND TERMITES 189 Based on this predatory habit. perhaps prematurely. To encourage the ants and reduce damage by P. naked caterpillars and grubs and thus keep the host free of these insects. Other Formicin Ants Many Camponotus spp. 2. occur worldwide. longinoda in similar fashion for the biological control of palm pests in Tanzania. attempts to exploit O. Vanderplank 1958. Pupae and maggots make excellent bird bait (Silow1983) and the ants themselves have been used to clean small skulls for museum purposes (Loveridge 1944). The carton nests are more or less spherical.2. 1984). a failure (Brown 1955. such as the fruit-spotting bugs Amblypelta and Pseudotheraptus wayi Brown (Coreidae). including carpenter ants. because they depress weaver ants while allowing the pest coreids mentioned above to thrive and become damaging to palm crops. Unfortunately. Crematogaster spp. Most important among these are Pheidole (Myrmicinae) and Anoplolepis ants (Formicinae).5 Myrmicinae 2. Internally. in stipules. mixed with soil and secretions from maxillary glands of the worker ants.4.700 years ago in southern China (Hölldobler and Wilson 1990). hollow branches or stems of various host plants (Kohl 1909. In Tanzania only certain arboreal species with a craving for honeydew were mentioned as potential pests by LePelley (1959).5. giving them a spongy appearance. the intercropping of citrus trees with coconut has been suggested (Behrens et al. papery walls. 1960. Control of weaver ants is only justified.: Black Cocktail Ants There are many species of black cocktail ants worldwide. their activities include both beneficial and noxious aspects. the nests are composed of cells with thin. Way and Khoo 1992). At the same time. Many prey on termites. O. where they are too much of a nuisance (Lever 1979). 1993).1. smaragdina is believed to have been the first insect employed for biological control of other insects more than 1. a group of structural pests. Hölldobler and Wilson 1990). Juniperus and Pinus (Gardner 1957a). . as they favor the buildup of sap-feeding Homoptera. Grevillea. As with many other ants. these ants are fond of honeydew and thus frequently favor the buildup of scale insects and aphids to damaging levels. wayi. were considered.
August. typically at the base of the stipule. penzigi.e. mimosae Santchi. C. Serengeti. accounting for the name whistling acacia. C. Wind rushing by these hollow structures can produce a whistling sound.190 CHAPTER 6 In Tanzania. in which they house their eggs. a short tree with conspicuous galls that are 4-7 cm across (Figure 6-6. Hocking 1970). The ants build interior partitions in these gall-like structures. making this essentially an obligate antacacia association (Hocking 1970). drepanolobium. Normally only one species of ant is associated with each tree. Apart from nest sites. or from many holes. sjöstedti Mayr. Less than 1% of A. Mt. (Myrmicinae). drepanolobium. respectively. food. Meru.. nigriceps Emery. several species of cocktail ants inhabit the swollen thorn bases (stipules) of some 13 species of dry-land acacias (Sjöstedt 1910. drepanolobium are ant-free. nectar from leaf . Dark-colored galls may indicate higher elevations. The most ant-dependent species of Old World acacias is A. or rarely. to A. larvae and pupae (Sjöstedt 1910). The trees initially produce intact swellings before cocktail ants or the ponerine ant Tetraponera penzigi (Mayr) hollow them Figure 6-5. Carton nest of black cocktail ants out from at most two. Crematogaster sp. Plate 48). None of the East African cocktail ants is more strongly adapted to its host than are the “red heads”. the tree provides the Figure 6-6. Acacia drepanolobium with the pseudo-galls ants with two sources of housing Crematogaster ants. i. African acacias also have a number of true galls (Plate 47) caused by other agents. mostly insects (Monod 1968). and the remainder with T. Two thirds of these acacias are inhabited by this ant while a little less than one third is associated with “yellow tails” C.
An equally interesting adaptation is reflected in the preying mantis Sphodromantis obscura. modified stinger. vidua have thousands of tiny yellow workers barely more than 1 mm long. excluding the abdomen (Skaife 1979. drepanolobium and preferentially preys on cocktail ants. but only half her size. Hocking 1970).5. the ants keep herbivorous insects in check and deter browsers. Workers of Crematogaster are 3-6 mm long. causes a painful reaction (Skaife 1953. Workers often are found alone and eggs may be concentrated in a few swellings. Colonies of C. The queen resembles workers in shape and color but is up to 1 cm long and her abdomen is swollen. recruited by an alarm pheromone from the mandibular glands of other workers (O’Toole and Preston-Mafham 1985).e. the former correlated with higher altitudes or shade. Huis 1996). whitish liquid from glands at the tip of the abdomen which when wiped into their bites with the spatula-like. Cocktail ants derive their name from the way they “cock their tail”. The weight differential between queens and workers is over 4. Mature swellings of A. Highly agitated when disturbed. most remarkably some chrysomelid mimics of C. Any one pseudo-gall may contain a combination of stages and sexes or just one.2.ANTS AND TERMITES 191 nectaries. . They also exude a sticky. Carebara vidua Smith. Unlike ants in other subfamilies. but are more slender. malodorous. apparently an adaptation to prevailing temperatures (Hocking 1970). discovered in Tanzania and described by Beier and Hocking (1965). 1979). which the ants harvest and take to their nest (O’Toole and Preston-Mafham 1985). They are covered with pale hairs and have a yellow/orange abdomen. This insect has an uncanny resemblance to pseudo-galls on A. drepanolobium are either dull black or bleached white. pointed abdomen raised over the back when alarmed. There are also numerous associates of ant-occupied swellings. carry their flat.. as well as oils and proteins in little outgrowths on the anthers.: African Thief Ants The genus Carebara is pantropical and in Africa is represented with 11 species. Males resemble workers. The winged males are similar to the queens. Natural enemies of cocktail ants include driver ants and birds (Sjöstedt 1910. myrmicins have a two-jointed petiole. In turn. they typically swarm in numbers. The pruning of axillary buds of the host stipules by the ants is interpreted as promoting a tree’s adaptation to dry environments (Hocking 1970). mimosae (Hocking 1970). i. The giant queens are up to 25 mm long and have wingspans of 50 mm. Some peoples in southern and eastern Africa regard the queens a delicacy eaten raw or fried. Despite the common occurrence of these ants. 2. The queens (Figure 6-7) are dark-brown or black and have a bulbous thorax and large abdomen. Mating flights occur in November-December. overall evidence concerning their biology is circumstantial and there are still unsolved mysteries (Hölldobler and Wilson 1990).000 (Hölldobler and Wilson 1990). the latter with lower altitudes.
Miscellaneous Other Myrmicine Ants Myrmicine relatives of cocktail ants in Tanzania include some that notoriously contribute to the buildup of Homoptera on crop trees such as species of Myrmicaria and Solenopsis (Le Pelley 1959).900 species of termites are widely distributed. include Decamorium spp. are crop pests by tending Homoptera and impacting native ant faunas. With approximately 170 genera (77 % endemic) and at least 664 species in tropical Africa. The 286 genera and almost 2.3. Myrmicine termite predators. At that time. sheds her wings and excavates her new nest (Robertson 1995). Africa is termite territory par excellence. she eventually flies to the ground. Early January. After a short flight. Eyasi. Pseudacanthotermes and Odontotermes (Grassé 1986). Others. large numbers of workers escape from underground through emergence holes built by the workers. Fossil records reach back to at least 144165 million years ago. The huge quantity of sperm stored in her spermatheca suggests a large number of eggs. except after the first heavy summer rains when the sun is high. 2. thief ants are rarely seen. where the minute workers apparently gain access to termite eggs and nymphs and then escape with their booty through tunnels small enough to prevent soldier termites from catching up.5. mostly in the warmer regions of the world (Scheffrahn 2004). possibly to help with the founding of the new nest. including the now cosmopolitan Pheidole megacephala F. Reports of young queens carrying workers attached to dense tufts of tarsal hair during their wedding flight. other than the just discussed African thief ant. the queen settles on a branch and releases a pheromone. Because of their subterranean existence. Figure 6-7. Nuptial flights continue for about 2-3 days.192 CHAPTER 6 African thief ants are exclusively associated with the nests of large termites Macrotermes. 3. this continent has the richest termite fauna . (Robertson and Villet 1989). Carebara appear to be a myth vidua (Myrmicinae). Mating repeatedly. Mangola.. TERMITES (ISOPTERA) Termites are considered primitive insects. The queen of an African thief ant.
By 1959. This list eventually grew to 177 species for East Africa. Still other termites are harvesters of grass or soil feeders. 10 new to the territory and two species new to science. Even if attacking wood above ground. Habits All termites are plant or fungus feeders and many rely on wood for both food and habitat. Depending on species. Uys 2002). Le Pelley had listed a total of 124 species of agricultural and tree termites for East Africa. Several genera of higher termites ments underground or in trees (Figure build arboreal nests. all but 20 of them documented for then Tanganyika. Chalinze. 8 in southern Africa. including 116 species in Tanzania (Weidner 1960. . while others build carton nests made of soil. the higher ones employ bacteria. Wanyoni et al. The remaining 39 were endemic to East Africa. termite nests are either entirely confined to wood (wood dwellers). 11 were widely distributed throughout the afrotropical region. Harris (1941) numbered the then known termites in East Africa at 77 species in 19 genera including 53 species in Tanganyika. but he predicted that many more awaited discovery. Extremely few occurred in the cedar and evergreen montane forests of the Usambara Mountains. and found the coast had the greatest diversity in habitats and species. Others cultivate fungi in a network of combs composed of termite feces and partially digested wood to supplement other food including the aging fungal combs. saliva. and 1 in Ethiopia. Kambhampati and Eggleton 2000. as frequently encoun6-8). 1904. 3. fecal matter and/or woody fragFigure 6-8. subsequently listed 51 species of termites. Kemp (1955) investigated the distribution of termites in northeastern Tanzania in relation to eight vegetation zones.300 m. 18 were also found in the Congo and in West Africa. 1984).1. Of these 77 species. 1910. Twenty of the 60 species occurring there were restricted to that zone. Indeed. Some termites construct elaborate concrete-like mounds or “termitaria” (Plate 49). The lower termites utilize cellulose with the help of symbiotic intestinal protozoa. or they are located entirely or partially underground (ground dwellers). or on cultivated land above 1. tered in humid coastal climates. Morstatt (1913a) had reported only 25 termite species in German East Africa including seven new ones discovered during Sjöstedt’s expedition at Kilimanjaro/Meru. Harris (1936a) following his three-year exploration of the Tanganyika territory. 1913. Five years later.ANTS AND TERMITES 193 in the world (Sjöstedt 1900-04.
As a result. and have more favorable physical characteristics. In primitive termites. soil structure and aeration (Hesse 1955a.194 CHAPTER 6 the ground dwellers usually retain nests in the soil from which they seek access to remote sources of wood. They are furthermore a prolific source of protein-rich food for some . these insects play enormous roles in tropical ecology.2. entire landscapes can be visibly shaped by the activity of termites. effectively supplementing the action of fires.2. represent external signs of termite activity.b). Termites and Animal Associates Termites also provide wooden and earthen cave habitats for numerous organisms (Escherich 1909. Earthen runways of Odontotermes sp. These foraging tubes. unless one exposes the extensive honeycombing within or the structure affected is completely hollowed out and collapses. i. 3. 2000). They significantly contribute to nutrient cycling of dead organic matter. Ecological Significance of Termites Figure 6-9.. where distinctive non-grass flora is mostly associated with the insular termite mounds dotting a sea of grass. many termites build earthen runways over the bark of living trees (Figure 6-9. To avoid exposing themselves during these forays. Pearce 1998) including numerous “termitophiles”. New studies indicate that soil is a significant sink for termite metabolic gases. ranging from less than 1 gram to over 10 grams per m3. in more advanced forms the nests are concentrated and have distinctive architectures (Beeson 1961. Termites and Soil Since termite biomass. (Termitidae) on Delonix regia. i. Soils in the neighborhood of mounds tend to be enriched with clay and organic matter. mostly beetles and other arthropods (Wilson 1971. arboreal nests. bacteria and other decomposers (Abe et al. together with mounds. Most of their tunneling in wood may go unnoticed. none more so than the so-called termite savannas and woodlands. Morogoro. 3. commensal insects. Plate 75) or when bridging concrete basements en route to structural timbers.2.e. the nests tend to be irregular (diffuse). 3. often rivals that of terrestrial vertebrates (Scheffrahn 2004).1. Kistner 1998)..2. Noirot and Darlington 2000).e. swarming termites or in the case of drywood termites fecal pellets expelled to the outside. Grassé 1982-86.
General Characteristics As an order. Their principal predators include certain ants and Bengalia floccosa (Wulp). and often a pair of small cerci. These are the fruiting bodies of the termites’ obligate symbiotic fungi (Heim 1977). microcarpus (Berk. “mkufu” T.3. beaded to threadlike antennae.1. Description and Life History 3. where “mgunda” T. (Heim). The edible termite mushroom Termitomyces (Eutermitomyces) letestui dia of Termitomyces before embarking on growing out of a mound of Macrotermes their nuptial flight. held flat and extend beyond the tip of the abdomen. letestui (Pat.) Sands (1970) The about 10-11 weeks later. Krishna is built by the workers at the new site and Weesner (eds. Termites are polymorphic. 495-524”. “busolele” T. Fungal propagules sur. Kemp 1955). reproductives of Microtermes usambaricus Sjöst. aardwolf.. Pearce 1998) including termite specialists such as aardvark. as well as for many animals (Grassé 1986. each differing in appearance and performing different . are known to eat coni. pangolin (Figure 6-2. termites have typical chewing mouthparts. The order’s name Isoptera refers to the two pairs of equal-sized wings. 3.ANTS AND TERMITES 195 indigenous people.sp. singidensis are among the most highly prized mushrooms (Härkönen et al.3. from fungus combs in situ.2.) Heim (Plate 50) and “impora” T. with permission from Elsevier). usually four tarsal segments. Plate 45). “kimelo” T. which are delicate. & Br. one species has several castes. For instance. Termites actively disseminate their symbiotic fungi either by inoculating propagules near the mound or by ingesting them.e.3. inoculates the comb with the fungus. aurantiacus Heim. 1995).) Heim. 3. and the female association of termites and fungi. This tropical genus of termitophile agarics (Amanitaceae) is well represented in Tanzania. 1922. straight. Termites and Fungal Associates Equally as sought-after for food as the termites themselves are highly nutritious and sometimes huge mushrooms (Termitomyces spp. until a new fungal comb from “Biology of termites Vol 2.) that grow exclusively in association with termite mounds of the Macrotermitinae (Figure 6-10). (Reproduced vive in their guts. i. eurrhizus Berk. a predatory fly (Morstatt 1913a.Figure 6-10.
With the exception of some queens. Figure 6-11. (From Morstatt 1913c.196 CHAPTER 6 tasks with the indisputable logic of a computer. These include workers. termites tend to be small to medium-sized insects of less than 25 mm. (Termitidae). modified). . Harris 1940b) (Figure 6-11). showing all castes except secondary reproductives. nutrition and/or sensory stimuli. that may reach a length of over 100 mm at maturity. Factors affecting caste formation include gender. soldiers and reproductives (Morstatt 1913a. pheromones. Schematic life cycle of Macrotermes sp.
In many species. in others they may have different tasks (Pearce 1998). After some time. They are usually slightly larger than the workers and are mostly blind and white. and attract mates with the help of pheromones. In the process. undifferentiated instars and nymphs in the advanced instars of the reproductive forms. often asymmetrical mandibles. early instar larvae take over the routine chores of the growing colony.ANTS AND TERMITES 197 Depending on family and species. Once a suitable site is found. In drywood termites there are no workers at all. Like the workers. or around 100 million in a lifetime. mostly unsclerotized individuals responsible for foraging. the male follows the female in tandem (Plate 51). These royals initially resemble each other and are equipped with two pairs of wings. common termites in much of sub-Saharan Africa. they jointly excavate space to start a new colony. egg-laying queens often becomes huge (Figure 6-13). However. They are also referred to as “rainflies” because the waves of dispersing squadrons are often synchronized with the arrival of rains. In certain species. they have compound as well as two simple eyes.2. tending other castes and nest building. sclerotized heads. Soldiers of some species are equipped with huge. and some species lack soldiers altogether.000 cylindrical eggs per day. workers of either sex share chores. After a short flight. During their “engagement”. they are sterile males or females and in some species appear in two or even three distinct sizes. flights for different species of termites in eastern Tanzania were recorded in all months except July and August (Krishna and Weesner 1970). Only nymphs of the latter exhibit wing buds. repellent or toxic substances in their chemical war against attackers (Quennedy 1975). in others they are fewer. In most species. while others have either frontal glands (“fontanelle”) or nose-like projections (“nasus”) that emit sticky. Soldiers defend the colonies against intruders. soldiers or fertile adults (reproductives). for a possible total of over 10 million in a single year. Primary Reproductives are fully sclerotized kings and queens. mating takes place in the new royal cell that effectively becomes a prison the two will never leave. In some termite colonies soldiers may make up as much as one fourth of the population. only nymphs or so-called “pseudergates”. queens of wood-feeding termites lay from 12 to over 48. there may be workers of two distinct sizes (“majors” and “minors”). They are typically white. Castes Immature termites of different castes are called larvae in the lower. female reproductives shed their wings retaining only triangular stubs called “scales”.3. the abdomen of mature. the inter-segmental membranes stretch . except for their oversized. They care for their first offspring until the first. They molt at least three and as many as ten times before becoming sterile workers. 3. Workers are generally the most numerous members of a colony and include both sexes. Males also shed their wings as soon as they have found a mate (Figure 6-12). as calculated by Escherich (1909) for Macrotermes. Because of her growing ovaries. during which time they are called “alates”.
April. Olmotonyi. These queens. Gembloux. two queens share one royal chamber. the two royals remain together in their fortified chamber for years and possibly decades. brown islands in an expanse of soft. (Termitidae) attracted to a door light. Lemaire. king (front right) and soldiers of Macrotermes falciger (Termitidae) in royal cell (Photo: L. white skin thin enough to see the pumping heart. Mature queen. although up to eight have been found. frequently described as “bloated sausages” or “gross egg-laying machines”. mating intermittently. eventually become so obese that they cannot move on their own. Figure 6-13. Unlike social Hymenoptera. and rarely. by permission of Les Presses agronomiques. Shed wings of a swarm of reproductives of Odontotermes sp. A colony usually contains but one queen. . From Malaisse (1997) Se nourrir en forêt claire africaine. Belgium).198 to an extent that the abdominal sclerites are widely separated as small. CHAPTER 6 Figure 6-12. Approche écologique et nutritionelle.
as they not only rely on mandibles. One family attacks only wet. Up to 200 secondary reproductives and. as well as in much of subSaharan Africa. For instance. with emphasis on wing venation. digestive tubes. two are restricted to Australia and South America. rotten wood and. Cryptotermes dudleyi Banks has been described under at least seven generic names. keys for termite identification were mostly based on alates and soldiers (Table 6-2). 3. . The use of recently developed keys for the identification of worker castes of termite genera from the soils of Africa is more complicated.3. Identification Termite classification is to a large extent based on the appearance of reproductives or soldier castes. even tertiary reproductives may occur in a colony. Traditionally.3. rarely. These are small-eyed. also called “neotenics” or “supplemental” reproductives. weakly sclerotized. Of the seven families of termites worldwide. The four remaining families of termites are commonly found in Tanzania. antennae.ANTS AND TERMITES 199 Secondary reproductives. but on internal characteristics as well (Sands 1998). occurs in only two species restricted to South Africa. which allows application under field condition. only occur if something should happen to a single queen. Many termite genera in Africa are considered in need of taxonomic revision (Uys 2002). respectively. modified female workers that develop only non-functional wing buds. eyes and the frontal gland called fontanelle. Numerous taxonomic changes through the years account for much of the confusion with names. in Africa. mandibles.
4. Becker 1976. 1971. but this is not an absolute requirement. Wilkinson 1965. dams. dead branches and stumps or the exposed wood on living trees in nature. before renovation. In East Africa. 3. railroad ties. furniture.4. they attack several trees from an external base connected through subterranean tunnels or earthen runways. with the bulk of damage being attributable to fewer than 2 % of species (Scheffrahn 2004). altitude. Under most conditions. temperature and other climatic elements. Bigger 1966). and higher termites (Termitidae). . including species and condition of trees. Termites in Forestry CHAPTER 6 While overall termites play extremely beneficial ecological roles. as well as in Machakos District. Living Trees Certain termites also attack sound wood in living trees.. damage occurring in restricted (Kalotermitidae) and free-range (Rhinotermitidae and Termitidae) categories (Wilkinson 1965).3. but some make incursions into living sapwood and beyond. Structural pest termites are often the same species as those associated with snags. pianos and such (Figure 6-14). foremost for eucalypts but also for agricultural crops (Brown 1965. Harris 1943. termites can become forest pests in three major contexts (Morstatt 1913a. books and other documents in the Tanzanian National Archives in Dar es Salaam suffered serious damage from termites. Impact on Young Trees Many of the higher termites (Termitidae) and occasionally harvester termites (Hodotermitidae) injure and kill living seedlings and saplings by ring-barking them on or just below the surface. as very healthy trees may also suffer severely under certain circumstances.1. scars and fire damage. Becker 1976). Harris 1971. They generally remain confined to dead wood. or by destroying the roots (Morstatt 1913a. soil type. Impact on Structural Wood Structural timber in buildings. For instance. the most seriously affected regions are round Lake Victoria into Central and Western Uganda. Wilkinson 1965. into heartwood. Selander and Bubala 1983). Sangster 1956. Drywood termites are restricted to one tree. gaining access through dead branches. 3. Impact on Larger. about 10 % of species worldwide are considered pests (Pearce 1998). species of termite. Factors accounting for attacks on new plantations are very complex. telegraph poles. food alternatives.2. More specifically. Wilkinson 1965.4.200 3.4. fences. Kenya and Iringa. (Morstatt 1913a. as well as anything containing cellulose and sometimes other organic substances is likely to be attacked by various drywood (Kalotermitidae). i. frustrating researchers by eliminating portions of irreplaceable documents (Schabel 1990). 3. subterranean (Rhinotermitidae).e. This is the biggest pest problem in exotic tree nurseries and plantations in semiarid Africa during the dry season. moisture content. unhealthy trees are the most susceptible. The majority of injurious termites are free ranging. 1977). Tanzania (Wilkinson 1965).
There are no fontanelles. There are 45 species in tropical Africa. Damage by Microtermes (Termitidae) in door of imported. Kambhampati and Eggleton 2000). grass seeds stored by these termites in underground chambers are collected by locals and used for food and the brewing of beer. While under natural conditions these termites contribute to the breakdown of dead trees and branches. A.2. and there are no mounds. Structural damage by various termites to timberwork in buildings. (From Harris 1943. Damage by Coptotermes amanii (Rhinotermitidae) in untreated. In parts of Africa. Their deep subterranean nests are difficult to find. Kalotermitidae The so-called drywood and dampwood termites occur throughout the (sub)tropics (Eggleton 2000).1. Kenya Agricultural Research Institute – KARI). Soldiers have robust mandibles and antennae are composed of 23-31 antennal segments. and six genera in East Africa (Harris 1950. Major Termites of Tanzania 3.5. Hodotermitidae Two species of Old World harvester termites are widely distributed in dryland Africa and parts of Asia. reproduced by permission of Director. These termites are more sclerotized than most others.ANTS AND TERMITES 201 3.5. All castes have compound eyes but at most rudimentary ocelli. some Figure 6-14. although minor damage to young trees and shrubs has also been reported (Wilkinson 1965). They feed mostly on grass. With lengths of over 15 mm. these are fairly large termites. C.5. which makes them less susceptible to desiccation and thus allows foraging excursions outside the nest even during the day. 3. imported ceiling boards. . painted but not treated timber. Drywood termite Cryptotermes (Kalotermitidae) galleries in roof pole. B.
than for damage in structural wood. There are no special royal chambers and the queen produces only about 12 eggs a day. Cryptotermes. Major Drywood Termites of East Africa. composed of a few hundred to a few thousand larvae. hexagonal fecal pellets the size of poppy seeds (Figure 6-15) that pile up in telltale small heaps beneath the infested wood. Virgin reproductives have wings with three or more heavy longitudinal veins at the anterior margin. Records exist from Hong Kong. University of Florida). Scanning electron micrograph developed mandibles and often short. Cryptotermes brevis Walker is of neotropical origin. Reproductives of these termites have antennae with 14-16 segments and iridescent wings with the medius and radius sector meeting in the distal half. (Reproduced by permission of R Scheffrahn. plug tunnel entrances (Figure 6-16). This accounts for generally clean galleries. logs. Drywood termites live confined to galleries in dry or slightly moist (15-20 %). sound wood of seasoned timber. the normally plugged exit holes are opened to expel dry. Only the reproductives venture outside for their dispersal flight. even internationally. In other parts of the tropics. Despite intrinsically limited mobility. Canada and some Pacific Islands. the southern USA. It is considered a completely domesticated termite and one of the most destructive of structural pests (Krishna and Weesner 1970). Although this termite has not been found in Tanzania. it has been thoroughly entrenched in South Africa since 1918. no fontanelles and antennae with fewer than 21 segments. of fecal pellets by Cryptotermes brevis truncated (phragmotic) heads used to (Kalotermitidae). Periodically however. colonies build up slowly without giving much external evidence of their presence. occasional soldiers and the reproductives. Soldiers typically have strongly Figure 6-15. There is no worker caste. Colonies are small. Only one genus. some species have occasionally damaged certain tree crops by moving from dead wood into living tree parts (Weidner 1988). some of the drywood termites have expanded their range considerably. by hitching rides on infested ships and wood products (Krishna and Weesner 1970). less for deteriorating dead trees. and there have been isolated . brownish termites resemble harvester termites. Three species in East Africa are of interest. Their reproductives have simple eyes. dead branches and structures that have no direct ground contact. but anything containing cellulose. As a result. These medium-sized.202 CHAPTER 6 cause concern when they attack not only structural wood (Plate 76) and various wood products. is of major concern.
Scanning electron micrograph of structural damage in East Africa the phragmotic head of a Cryptotermes brevis including stored Podocarpus timber. this termite must be considered a potential invader of Tanzania. Sierra Leone and Zimbabwe (R Scheffrahn. Dorsal view of head and pronotum of Cryptotermes havilandi (Kalotermitidae).). (Reproduced by permission of R Scheffrahn. Cryptotermes (=Calotermes) havilandi Sjöstedt is apparently a native of Africa from Kenya to Zululand. University of Florida). The remaining genera of dry-wood termites occurring in Tanzania (Bifiditermes (=Kalotermes. pers. including Cape Verde. Zaire. In May this termite is on the wing in eastern Tanzania (Krishna and Weesner 1970). This termite (Figure 6-17) typically occurs in dead branches. Bicornitermes (=Calotermes). accounts for most Figure 6-16. . It also occurs in Uganda. but is now also common in Madagascar. Brazil and parts of Asia. the Comoro Islands. Egypt. fallen wood or stumps but rarely in houses. the latter two being pantropical. Helena. including Zanzibar (Gardner 1957a). Mauritius and Madagascar. This termite is of oriental origin and now widely spread in neotropical America. Glyptotermes and Neotermes). British Guyana. Pretoria. Epicalotermes (=Kalotermes). Nigeria. Miscellaneous Other Drywood Termites. Procryptotermes).ANTS AND TERMITES finds in 11 other African countries. (Kalotermitidae) soldier. St. Trinidad. As a result. comm. Records of this termite from buildings in Dar es Salaam and Mombassa may be based on misidentification of C. It was first reported in 1950 in stored timber at Tanga and subsequently became thoroughly entrenched along the East African coast from Somalia to the Rovuma River. Gambia. Agricultural Research Council. Madagascar. and Sri Lanka (Krishna and Weesner 1970). dudleyi. 203 Cryptotermes dudleyi Banks. Senegal. reproduced by permission of GL Prinsloo. the drywood termite. Southeast Asia. (From Uys 2002. northern Australia. There has been one report of it having “possibly” surfaced in Uganda (Campbell 1974). SA). from the Congo to West Africa. include seven mostly “wild” species found in fallen logs or Figure 6-17. Ghana.
Subterranean termites live in large colonies in fixed nests underground. They are commonly found in old tree stumps and buried timber but also excavate permission of GL Prinsloo. Soldiers have a flat pronotum and antennae with 14-22 segments. the oriental C. of Bifiditermes durbanensis (Kalotermitidae). As a result. Harris (1941. 1950). Satellite nests may be maintained in trees and buildings. . and the gallery walls are covered with a pasty-looking wood pulp. Dorsal view of head and pronotum of soldier family Termitidae.204 CHAPTER 6 dead branches on trees. They do. formosanus Shiraki has become established as a structural pest in subtropical South Africa (Harris 1966) and. reproduced by into the temperate zone. 3.5. lamellar fashion with denser wood remaining as walls. Agricultural Research pipes in the heartwood of mature trees. as it has damaged buildings and in trees may go from dead into live wood (Wilkinson 1965. or a mixture of soil with chewed wood. SA). They destroy wood in a layered. (1965) and Weidner (1988). All castes have frontal glands (fontanelles) that are conspicuous in soldiers. and attack Council. The wings of reproductives have two heavy longitudinal veins in the anterior margin. (From Rhinotermitids occur throughout the tropics and Uys 2002. Front wing stubs are longer than the pronotum. Rhinotermitidae may survive in ships’ timbers. some of them have become pests far from their place of origin. Major Subterranean Termites of East Africa. Kemp (1955). Wilkinson et al. (=Kalotermes) durbanensis (Haviland) (Figure 6-18) is of slightly greater concern than the others. When alarmed. Only Bif. or simply “subterranean termites” to differentiate them from other ground-dwelling termites. range as far as 100 m from this primary nest commuting through foraging tubes made of chewed wood and soil.3. There are 14 African species of Rhinotermitidae and two genera with two species each occur in Tanzania. soldiers exude a drop of sticky fluid from this gland. They were listed and some illustrated by Sjöstedt (1926). however. This termite has spread and now occurs from Tanzania to South Africa. Because of their need for high wood moisture. but remain connected to the primary nest by runways. floors and ceilings of a series of compartments. These termites are less than 5 mm long. especially the “higher subterranean termites” in the Figure 6-18. Pretoria. Rhinotermitidae Insects in this family are called the “lower subterranean termites”. For instance. Weidner 1988). buildings. The pantropical genus Coptotermes sports an extensive record of introductions through various ports of entry. Infested wood typically contains soil.
Of the two types of (Rhinotermitidae). The soldiers in this genus have highly developed fontanelles. It may have many subsidiary nests in neighboring lamanianus. sjöstedti occurs in western Tanzania and is most prevalent in West Africa. In 1999 it was documented as the first African endemic termite to become established in the Americas (Scheffrahn et al. and a slightly drawn out or markedly arched clypeus. 1966). as well as and pronotum of in gallery forests (Gardner 1957a). .ANTS AND TERMITES 205 in 1960. the other with an extended labrum. Head and pronotum of soldiers of (A) Coptotermes amanii and (B) Coptotermes sjöstedti (Rhinotermitidae).000 m. (From Harris 1966. Kenya and adjacent islands (Gardner 1957a. which occurs from Somalia through East Africa to Zimbabwe and now also South Africa. Two indigenous species. Harris 1966). mature trees used in agro-forestry or as ornamentals. routinely attacks mature trees in many species. putorius Sjöst. The main nest tends to be at major soldier of the base of trees. It normally inhabits tree stumps and rotting logs. S. but has also been documented damaging old seasoned timber in buildings in Tanzania. the larger ones (Figure 6-20) are less common. Figure 6-19. Schedorhinotermes (=Rhinotermes) spp. such as Samanea saman. Manihot and Persea Figure 6-20. one with large mandibles.) (Figure 6-19) prefer dead tree roots or wounds near the collar of standing trees where carton nests with a thick-walled. lamanianus (Sjöst. moist coastal climates. Coptotermes sjöstedti (Holmgren) and C. This genus is widespread not only in Tanzania but in the Afrotropics altogether (Harris 1941. amanii. but can also do serious structural damage and attack mature trees through wounds. (From Sjöstedt 1926). The reproductives have a circular head. It occurs at altitudes up to 1. Head americana in cities in warm. Jacaranda. 2004). Dalbergia. is best known to attack badly pruned. central queen cell are constructed (Harris 1941. 1971). with occasional subsidiary carton nests above Schedorhinotermes (Kemp 1955). reproduced by permission of The Royal Entomological Society of London). soldiers. A B C. amanii (Sjöst. and is particularly conspicuous in Delonix. The golden tree termite S. and S. have soldiers of two sizes with little resemblance to each other.). where it accounts for much damage to buildings and timbers (Harris 1966). was discovered on a fisheries research ship moored in several East African ports (Wilkinson 1965). trees connected by tunnels (Pearce 1998). lamanianus Sjöst. C. Its flight in eastern Tanzania is in November (Krishna and Weesner 1970).
this is the most troublesome family of termites (Atkinson 1989. Different species cause serious damage to natural regeneration. trunk wood. and arboreal nests. cambium. These termites cause damage in seven distinct patterns (Figure 6-21) (Selander and Bubala 1983). Amitermes. they have two heavy. Microtermes. especially during the drier part of the growing season. but the front wing stubs in the Termitidae are shorter than the pronotum. Macrotermes. all of which are shared by Tanzania. but some also damage agricultural crops. root collars and roots. partially subterranean concrete-like mounds. grass. Most live entirely or partially underground.e. including 601 African species (Kambhampati and Eggleton 2000). girdling at the root collar occasionally occurred. longitudinal veins at the anterior margin. Odontotermes. Certain species occur in enormous colonies of sometimes more than 1. Depending on soil. and in one subfamily there are no soldiers at all.5. The most common structural termites from this family include Macrotermes subhyalinus Rambur. and some feed on soil. There are subterranean nests made of wood carton. Mound builders start below ground before building turreted. Included are mound builders. often killing trees or leading to breakage or wind throw. Microcerotermes. In black wattle. but to retain the rigidity of the structure they are hollowing out. Ancistrotermes. Odontotermes badius (Hav. seedlings in nurseries and young. . Termitidae The huge family of so-called “higher termites” makes up about 80 % of termite species. recent transplants until crown closure occurs. but mature trees can also be affected. These act as respiratory devices to capture wind energy for ventilation (Figure 6-23). Pseudacanthotermes and Synacanthotermes. These termites tend to remove wood en masse. The pronotum of soldiers and workers is narrow and has a raised median lobe in front. outer layer with packed earth. higher termites caused very serious damage in 4-5 year old rubber tree and black wattle plantations (Morstatt 1913b). Significance to forestry. In older rubber trees that had been tapped. Most Termitidae are wood eaters. climate and other factors. During a drought in German East Africa. fungus growers and guest termites that cohabitate with other termite species. For forestry in East Africa. Workers in this family are blind.206 CHAPTER 6 3. In Zambian plantations. Wood below ground or encased in masonry tends to disappear entirely.) and Microtermes redenianus (Sjöst. Wilkinson 1965).) (Harris 1943). including Allodontotermes. i. but genera tend towards predictable architectural types (Noirot and Darlington 2000).. mound shapes vary in the same species. they fill the void spaces within the remaining thin. the major termite pests notoriously belong to 10 genera of Termitidae. Anoplotermes. damage was done to the bark. Many species have nasute soldiers. Wings of reproductives are similar to those in Rhinotermitidae.4. dung and other vegetable matter.000 individuals that may have existed for 80 years or more. buttressed castles (termitaria) sometimes extending to a towering 12 m above ground level (Bölsche 1931).500.
Pseudacanthotermes. dotting internal feeding. The taxonomy of these termites has experienced major revisions during the past decades. Ancistrotermes and Microtermes (Wood and Pearce 1991). the Macrotermitinae. Kambhampati and Eggleton 2000).. Termitinae and Nasutitermitinae. i. (From Selander and Bubala 1983). Crosshatching indicates superficial. especially Africa. The first three usually attack plants externally. Different patterns of termite damage to eucalypts caused by a variety of termite genera. Apicotermitinae. Included are the “big 5” genera. The stronghold of the fungus-farming termites is semiarid to arid land in the Old World tropics. Seven of the above-mentioned important genera belong to this subfamily of termites.e.e. Given the large numbers of genera and species in this family. Macrotermes. Subfamily Macrotermitinae. although it is still very much in flux for some groups (Kambhampati and Eggleton 2000). making it the single most important group of termites for field forestry in Africa. where 165 species occur (Eggleton 2000. it is convenient to discuss the more conspicuous or economically important higher termites which belong to four subfamilies..ANTS AND TERMITES 207 Figure 6-21. Odontotermes. . i.
Harris 1971. Selander and Bubala 1983). however. Ancistrotermes.208 CHAPTER 6 the latter two internally. This genus is widely distributed in tropical Africa and Asia and includes some of the largest termites. field damage to tree seedlings. The Prinsloo. damages crops in the miombo zone of southern Tanzania and in semiarid country routinely causes problems for seedlings in the 6-21 c. and in still others only males. They are pests in nurseries and attack young trees in agroforestry or in the field. Bigger 1966. The Macrotermitinae are not associated with symbiotic protozoa. head and pronotum of major Soldiers and workers are. Pretoria. or up to 45 mm from head to the wing tips (Morstatt Figure 6-22. Eggleton 2000). subspherical cells each containing a rosette-like fungus comb. Ancistrotermes latinotus Macrotermes soldiers are derived from female (Termitidae). They are small termites resembling Microtermes. Termes). Their soldiers sport formidable. sabre-like mandibles and have a saddle-shaped pronotum. Winged forms of Macrotermes falciger Gerst. Dorsal view of 1913c). Some are also known as structural pests in wood already subject to a degree of fungal deterioration. On semiarid sites in East Africa with a history of overgrazing. Ancistrotermes latinotus (Holmgren) (Figure 6-22) attacks wood. much smaller. toothed mandibles. In eastern Tanzania this termite flies in February and October (Krishna and Weesner 1970). but also in other parts of the Old World tropics. mandibles of soldiers are without teeth. These three genera occur not only in Africa. but cultivate fungi either underground or in their mounds (Wood and Thomas 1989). while in other genera soldiers may be sterile reproduced by permission of GL males and females. Most of the ten species in this genus are endemic to savannas in the afrotropical region (Coaton and Sheasby 1975. These termites live in subterranean nests composed of several. but their soldiers have more robust. (From Uys 2002. They have a rounded labrum with hyaline tip and mandibles with few teeth (Pearce 1998). (above) and minor soldier of measuring only about 20 and 11 mm. SA). Mature queens may be over 100 mm long. have wingspreads up to 88 mm and their bodies can measure 22 mm. nymphs. less than 10 cm long. f patterns (Kemp 1951. deforestation and over-reliance on exotic plantations. Agricultural Research Council. Macrotermes (=Bellicositermes. Reproductives have a labrum longer than broad and a chitinous. respectively. Odontotermes and Microtermes is almost predictable. . transverse band. especially by species of Macrotermes.
and Darlington 2000. The new M. as revealed by sheet. natalensis into M. falciger is considered THE most dangerous among the ten important plantation Termitidae in Zambia (Selander and Bubala 1983). The new M. subhyalinus. M. are composed of spongy grey-brown section through the vented mound of Macrotermes subhyalinus “combs” made of digested plant matter and feces.). M. but also contain underground corridors and caves (Noirot and Darlington 2000). older clove trees were attacked. cleaning up much confusion concerning a cluster of 23 species and 11 forms or varieties found in the literature. This species commonly makes bark tunnels. M. reaching as far North as Zambia and Mozambique. bellicosus (Smeath. Selander and Bubala 1983). According to this new scheme. has one isolated record from Tanzania. many of them synonyms. Next to. subhyalinus tolerates drier conditions than does M. Even the taproots of the most vigorous young trees are devoured in the 6-21d pattern. they were considered “THE major entomological problem in forestry” in that country (DW Barnett. and is well represented throughout the Zaire basin. The fungus gardens. falciger (=M. bellicosus and M. natalensis (Haviland) is mostly southern African. comm. In eastern Tanzania. On Zanzibar. He gave one former variety species status and split one species into two. goliath and occurs from Uganda and Tanzania southward into South Africa. when Malawi heavily relied on eucalypts for an ambitious fuel wood plantation program. bellicosus (Smeath. M. pers. subhyalinus (= part M. grouped around a substantial Figure 6-23. shows the royal chamber embedded These termites attack sound wood. . Their mounds are enormous. and in concert with drought. leaving 12 recognized and re-described species. natalensis (Hav. after the splitting of M. Most of these termitaria reach aboveground and are equipped with vertical chimneys (Figure 6-23). Ruelle (1970) revised this genus for the afrotropical region. like earthen tubes on trunks. Diagrammatic vertical queen cell. ukuzii Fuller is restricted to the western part of Tanzania (Ruelle 1970).ANTS AND TERMITES 209 The huge colonies of Macrotermes live in very large mounds built of sand and clay mixed with saliva. modified). killing all ages of eucalypts. goliath) are important in Tanzania. Macrotermes usually fly in (Termitidae).) replaced the former M. bellicosus (Ruelle 1970). bellicosus and is common in the northern half of Tanzania. and 50 % of some coconut seedbeds were destroyed by Macrotermes (MansfieldAders 1919/20). is M. Two species of Macrotermes. including in the core of the nest (From Noirot living but unhealthy trees. they are considered the most significant obstacle to the culture of eucalypts and other trees in the drier parts of Africa often destroying entire populations of seedlings in their first year and even larger tree crops (Mansfield-Aders 1919/20.) occurs from Senegal to Uganda.)) and M. the most widely distributed species in Africa. M.). The second common and most important species in Tanzania. falciger (Gerst. replacing the former M. The bubbles indicate fungal combs and the dark area December (Krishna and Weesner 1970). In the 1980s.
there are 42 species of these most common of the small fungus growers. These are large termites. are very small insects. M. They include M. redenianus may be the most important.).210 CHAPTER 6 Microtermes. at least eight species of Odontotermes attacked all kinds of eucalypts in the 6-21b. Among these. M. At least six species of Microtermes are known in Tanzania. The nests (Figure 6-25) are often found under buildings (Gardner 1957a). magnoscellus Sjöst. occasionally in buildings. badius and latericius. Gardner 1957a). only somewhat smaller than Macrotermes. they were among four main species of termites attacking wood (Kemp 1951). f and g patterns (Selander and Bubala 1983). Their population densities in the upper 1 m of soil may reach over 4.. Most important in East Africa. O. alluaudanus was reported to fly in eastern Tanzania from October to December (Krishna and Weesner 1970). in South Africa. O. They can also become noxious in nurseries and on young trees. as their name suggests. They are widely distributed in tropical Asia and Africa (Eggleton 2000). There are a variety of nest types. In Zambia (Selander and Bubala 1983) at least three species attacked eucalypts of any size in the 6-21b. 1926 ). as it attacks both structural wood and young planted trees (Harris 1943. (=Microcerotermes) alluaudanus Sjöst. usambaricus Sjöst. and M. subterranean nests these termites attack sound wood. vadschaggae Sjöst. In Africa alone. both serious destroyers of structural wood. Odontotermes (=Termes) make up another important group of widely distributed fungus growers (Eggleton 2000). Head of soldier of Microtermes luteus (Termitidae).. are O. (=Termes) badius (Haviland) builds subterranean nests that reveal themselves only with an inconspicuous soil elevation without ventilation shafts. M. These termites differ morphologically from Macrotermes by having only one soldier caste which is equipped with a marginal tooth on the inner edge of the left and sometimes also on the right mandible. luteus Harris (Figure 6-24). some in trees. Together with Microtermes spp. seedlings and young trees (Harris 1971). especially in intensely cultivated areas.. M. M. but they engage in little or no mound building. There are 78 species in tropical Africa and others in Asia. may be the worst of the subterranean termites in terms of destructiveness (Skaife 1979). There are workers and soldiers in two sizes (Sjöst. This is the most common termite in East Africa and. In Zambia. From their diffuse. in East Africa causing mainly Figure 6-24. East and southern Africa. they are the most abundant termites on cultivated land (Kemp 1955). redenianus (Sjöst. . mostly in conjunction with dead wood. e and g patterns. as well as other parts of Africa.000 m2 per meter. Transplanted trees are also attacked. (From Harris 1961). M. e. Soldiers have smooth mandibles. In an experiment in Shinyanga. (=Termes) latericius (Haviland) (Figure 6-26) is also very abundant in dead wood throughout Central.
Unlike O. Pretoria. Pseudacanthotermes (=Acanthotermes) is an afrotropical endemic genus with eight species widely distributed in the woodlands of Africa. (Kemp 1955). (From Uys 2002. At least another 14 species of Odontotermes occur in Tanzania. SA). Their earthen runs are commonly seen on Commiphora spp. Figure 6-27.. Infestation of a house from a nest of Odontotermes badius (Termitidae) situated extramurally. Figure 6-26. Pretoria. . Transverse section through nest of Odontotermes latericius (Termitidae). militaris Hagen and P. structural damage. but nursery and young trees can also be attacked. the shallow nests of this termite are equipped with chimneys of clay around wide airshafts (Figure 6-27). They attack mature. reproduced by permission of GL Prinsloo. often ornamental trees. At least two species. Agricultural Research Council. occur in Tanzania (Harris 1940b). showing chimneys. (=Acanthotermes) spiniger Sjöst. Agricultural Research Council. badius. as evidenced by earthen runways on the trunks through which they reach wounds and dead branches. (From Uys 2002. reproduced by permission of GL Prinsloo.ANTS AND TERMITES 211 Figure 6-25. P. (From Uys 2000). SA). Dorsal view of head and pronotum of soldier of Odontotermes latericius (Termitidae).
Macrotermitinae of Lesser Significance in Forestry. they are the most enigmatic group of termites. militaris (Figure 6-28) attacked sound wood of tapped rubber trees at Nyussi. Gardner 1957a). Dorsal view of head and pronotum of major (left) and minor soldier of Pseudacanthotermes militaris (Termitidae). g patterns (Selander and Bubala 1983).212 CHAPTER 6 The prothorax of soldiers of these middle-sized termites has two spines on the anterior margin. (From Uys 2002. In Zambia. Most live in diffuse underground gallery systems. The three species of Allodontotermes are essentially southern African but reach into East Africa. resulting in hollow roots (Selander and Bubala 1983). killing 10 % (Morstatt 1913b). In addition to the preceding “big 5”. zanzibariensis Sjöst. attacking newly planted stock to mature trees in the 6-21b. The species group “Anoplotermes” consists of termites concentrated in southern Africa (Sands 1972). Both biologically and taxonomically. they count among the ten most important genera of higher termite pests in plantations. Three species of Synacanthotermes occur in Africa. including Allodontotermes. P. reproduced by permission of GL Prinsloo. morogorensis Harris. and only a few build nests. In German East Africa. some were implicated in attacks on eucalypts of various ages and sizes in the 6-21c pattern. These termites live in stumps and clove timber and build earthen runs. Protermes. is the principal one. These termites often build sheet-like rather than tubular tunnels over tree bark. an earth-nesting termite. SA). Figure 6-28. attacks on both young and mature plantation trees in the 1d pattern are common (Selander and Bubala 1983). They form a significant part of the soil fauna of tropical Africa and two genera include species that cause damage to trees and wood. several other genera of Macrotermitinae. e. Sphaerotermes and Synacanthotermes. Subfamily Apicotermitinae. occur in Tanzania. In Zambia. attacking newly planted to mature trees in the 6-21a. These soldierless termites consist of 108 species of soilfeeding termites endemic to Africa (Kambhampati and Eggleton 2000). causes damage to various trees in East Africa and also attacked wood in Shinyanga (Kemp 1951. In Zambia they were among the 10 most important higher termite pests in plantations. e and g patterns (Selander and Bubala 1983). Agricultural Research Council. In Zambia. Reproductives of this termite are widely appreciated for food (Harris 1940a). c. They have cryptic and diffuse nesting habits. Pretoria. The majority is . of which S. A.
Noditermes and Procubitermes). others prefer wood. This is a widely distributed pantropical mission of GL Prinsloo. the Amitermitinae. parvus Haviland may be the best known of the eight species in this genus in Tanzania (Harris 1961). Most form amorphous subterranean colony centers with often strikingly high biomass densities (Eggleton 2000). Amitermes spp. These termites are found in dead. very small species. The Amitermes group. Subfamily Termitinae. Council. Their antennae have 12-15 segments. (From Uys collar or root exterior in the 6-21a. cellulose and other plant matter.ANTS AND TERMITES 213 likely to be of greater ecological significance than as forest pests. reproduced by perBubala 1983). are a common sight in many parts of sub-Saharan Africa. Because of the great diversity of their habits. and they generally occur in soil. about 35 cm high and 50 cm wide at the base. Figure 6-29.e. rectangular head capsules and mandibles with incurved tips and inner serrations (Figure 6-30). domed mounds (Figure 6-29). Their black. feed mostly on dead trees and Amitermes sp. Many build small to larger mounds. dry wood. Three other species of Amitermes are found in dead wood or litter in Tanzania. Tanzania has species in at least the genera Angulitermes. Some were previously included in a separate subfamily. c.b patterns (Selander and 2002. genus that builds carton nests made from feces containing Agricultural Research lignin. hastatus Haviland are a common. Some are soil feeders. including the wood feeding Amitermes and Microcerotermes. Amitermes (=Hamitermes) were among the ten most important higher termites damaging plantation trees. attack newly planted eucalypts to mature trees and coppice crops in the a.g patterns (Selander and Bubala 1983). This is a small mound builder widely spread in dry woodlands from West through East to South . including Tanzanian representatives (Basidentitermes. SA). Cubitermes. Their small. In Zambia. these termites do not carry a common name. Crenetermes. animal dung and broken branches. is predominantly pantropical. Some are guest termites in the nests of other Isoptera. Euchilotermes. Vertical section through nest of Microcerotermes spp.The black mound termites A. domed mounds are subterranean or concealed in other termite colonies. Pretoria. The Cubitermes group is an endemic African soil-feeding cluster of termites. Pericapritermes. sound wood and also forage humus (Skaife 1979). but also attack seedlings and young trees at the (Termitidae). litter. There are 272 species in tropical Africa (Kambhampati and Eggleton 2000). Promirotermes and Termes. Soldiers of Microcerotermes are easily recognized by their long. M. The Termes group has typically soldiers with asymmetrical mandibles.
Dorsal view of head and pronotum of soldier of Nasutitermes kempae (Termitidae). At one time or other. Wigg (1946) offered a provisional list of some 285 species of East African timbers proven or believed to resist destructive agencies. 1971) turned out to be fairly small. Structural Damage Prevention of termite damage to structures ideally envisions the use of naturally resistant wood. others build small domeshaped mounds and still others are subterranean. Olea welwitschii. These termites have soldiers with pear-shaped heads. 3.. This was one of the four main species responsible for attacking wood in an experiment in Shinyanga (Kemp 1951). Wilkinson 1965. Piptadenia africana and Pterocarpus angolensis. Some Nasutitermes.1. 1971. 3. Pretoria. . Other species were recommended on a more qualified basis. Subfamily Nasutitermitinae. Termite Management Traditionally. and there are 601 species in the Afrotropics alone (Kambhampati and Eggleton 2000). different physical. (From Sjöstedt 1926). It attacks dead trees as well as wounds in young and mature trees.214 CHAPTER 6 Africa. Grallatotermes. which is supposedly perishable in ground contact but highly resistant used in roofs (Campbell 1974). characterized by a snout-like extension (nasus) and greatly reduced mandibles (Figure 6-31). including Afzelia quanzensis. both the German and British colonial governments promoted the search for and evaluation of tree species that would hold up against attack by termites and other destructive agents.6. Heads of major soldier of Microcerotermes parvus (Termitidae). Nasutitermes and Trinervitermes) occur throughout the tropics. Diospyros spp. The so-called nasutiform termites (Coarctotermes. Many species construct spherical carton nests on tree trunks. including termites Figure 6-30. Ultimately. chemical or silvicultural methods have been employed to address termite problems in structural or field contexts (Harris 1943.6. are minor structural pests. the most species-diverse genus in the higher termites. (From: Uys 2002. Erythrophleum guineense. Agricultural Research Council. including Leucaena glauca. SA). Pearce 1998). Figure 6-31. the list of resistant commercial timbers in tropical Africa or East Africa (Harris 1961. Brachylaena hutchinsii. reproduced by permission of GL Prinsloo. including the mangrove Rhizophora mucronata. Chlorophora excelsa.
if the ends or joints of timber. relevant experience with pesticides was summarized (Parry 1959. but it worked for the recent Tendaguru dinosaur expeditionary force (Maier 2003). or it was sprinkled below and around the roots and stems. chemical control of drywood termites provides only temporary relief. pesticides began to be restricted or banned by certain governments or donors since the 1970s. calomel with sugar mixed into the soil at the time of planting was considered effective in protecting seedlings. and the establishment of shields and other physical barriers on wooden structures. 18 as “ moderately attacked” and five as “almost unattacked”. Acacia fischeri. (1989) reported other approaches. the use of non-wood building materials. Conscientious application of oil-based paint does. Alternatively. followed by a second dose before outplanting. Cowie et al. Harris 1948. are considered termite-resistant (Schnee 1920). are fully covered. For instance. this effective . (1989). Among exotic species. Experiments with DDT. Acacia rovumae and Abrus schimperi. the same pesticides were used in the field. although at different rates. however. Strychnos heterodoxa. given a number of variables. Dichrostachys glomerata. 1949. promise some protection. Both pesticides gave 5-6 years protection. Insecticidal dust was either mixed with soil in the planting holes at planting time. 3. Based on concerns for the environment. According to Cowie et al. against “undue optimism about the natural resistance of any timber”. pressure-treating wood with copper-chromium-arsenate (CCA). This involved the thorough mixing of potting soil with dust or wettable powder of aldrin or dieldrin. where termites notoriously attack. Most of these solutions have limited prospects for wide-scale use in Tanzania. Reddy 1975). Kemp (1951) listed six species of timbers as “readily attacked”. As well. As a result.6. He cautioned. 1952). wettable powder mixed with water was applied to the root region at planting. BHC. When deep planting precluded nursery treatment. is only resistant after water seasoning. Simply standing the legs of chairs and tables in water or oil may be a somewhat awkward solution to keep termites at bay. mercuric chloride and creosote gave mixed and qualified results (Morstatt 1912b. as wood is likely to be attacked again. based on experiments conducted in Shinyanga. sodium arsenite. Field Damage In German colonial days.ANTS AND TERMITES 215 and made recommendations for the use of certain species in various regions. Wilkinson 1964. Important in this procedure was establishing a complete chemical barrier around the roots and on the ground surface around the tree. however. Other recommendations for structural termite control or prevention include bait laced with pesticides or molting inhibitors. Prompted by routine complications with termites in Tanzanian tree nurseries and dryland plantations. used by the Sukuma for poles. such as the blowing up of termite nests or the use of cyclodienes and other persistent pesticides as mound poisons or root barriers against termites attacking seedlings. Tectona grandis and Camphora spp. Wigg 1946.2. as well as the digging out of queen termites (Morstatt 1913b). The latter include Ormocarpum trichocarpum.
. . i. cultural and integrated controls accelerated (Weidner 1988. are also advised. 200 g asafetida. the use of poison baits. Jacaranda.216 CHAPTER 6 chemical approach was no longer feasible. Curry 1965a. Casuarina. 100 g flour. and 80 g sugar mixed into a stiff dough and. and the search for alternatives. Cassia siamea. a concoction developed in India. since controlled release formulations of more acceptable. Maesopsis eminii and Tectona grandis for such sites. such as 25 g Paris green. A lime/sulfur mix forked into soil also supposedly discourages termites. including 100 g gum. biological. Cowie et al. As well.. Douglas (1991) outlined alternative home-made termite controls for agroforestry. reliance on using termite-resistant seedlings continues to be a better solution for dry areas of Africa.e. and to provide alternate foods for termites. Gmelina. These included the use of Gondal fluid. Another cultural approach encourages planting of about 20-30 % more seedlings than are needed. However. mulching. modern insecticides tend to be expensive. alternatively. Callitris. and fertilizing. Albizia lebbeck. 1989. in conjunction with avoidance of susceptible exotic tree species. 200 g aloes and 80 g castor cake. weeding. including biorational. Recently. Wood and Pearce 1991). Harris (1961) and Brown (1965) recommended Acacia spp. thick flour dough mixed with 50 g borax and 100 g sugar were encouraged. Supplemental measures to enhance the vigor of seedlings by carefully matching trees with proper sites in combination with post-planting care.
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