How To Get Rid Of Termites

Oplan Termites

Oplan Termites

You Might Start Missing Your Termites After Kickin'em Out. After All, They Have Been Your Roommates For Quite A While. Enraged With How The Termites Have Eaten Up Your Antique Furniture? Can't Wait To Have Them Exterminated Completely From The Face Of The Earth? Fret Not. We Will Tell You How To Get Rid Of Them From Your House At Least. If Not From The Face The Earth.

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Termite Extermination Information

Termites create great damage to your home, which is why you should identify and eliminate them as quickly as they appear. This eBook Oplan Termites teaches you how to solve your termite problem once and for all. Learn how to identify termites, find out if your house is really infested, and eradicate them. Discover Some Of The Most Effective And Time-Proven Methods To Get Rid Of Termites! Learn Some Mean Ways To Really Get Rid Of These Pests From Every Nook And Corner Of Your Home.

Termite Extermination Information Summary


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The author presents a well detailed summery of the major headings. As a professional in this field, I must say that the points shared in this manual are precise.

As a whole, this manual contains everything you need to know about this subject. I would recommend it as a guide for beginners as well as experts and everyone in between.

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Net methane efflux from termites

Termites (worker caste) isolated in the laboratory have been reported to produce CH4 at rates ranging from undetectable levels to 1.6pmol g-1 h-1 (Brauman et al, 1992 Bignell et al, 1997 Nunes et al, 1997 Sugimoto et al, 1998a, 1998b). Zimmerman et al (1982) first drew attention to the potentially large quantities of CH4 emitted by termites. Their estimates were 75-310Tg yr-1, equivalent to 13-56 per cent of global sources. However, subsequent estimates of annual CH4 emissions from termites have been rather lower, cf. 10-90Tg (Rasmussen and Khalil, 1983), 10-30Tg (Collins and Wood, 1984), 2-5Tg (Seiler et al, 1984) and 6-42Tg (Fraser et al, 1986). More controversially, while acknowledging the uncertainties Zimmerman et al (1982) predicted that under certain scenarios of land use change, the quantities released could equal those of all other natural sources. Since then the subject has been revisited by numerous authors and almost all of the assumptions made by Zimmerman et al (1982)...

The Termite Gut Microecosystem

The symbiotic digestion of lignocellulose by termites is a complex series of events involving both the host and its intestinal microbiota (Brune 2003, 2005). While the digestive activities in the foregut and midgut seem to be mainly caused by the host, the processes in the hindgut are largely controlled by the symbionts. The recalcitrance of the lignocellulosic diet and the dynamics of physicochemical gut conditions contribute to the heterogeneity of ecological niches and are reflected in a diverse community of prokaryotic and eukaryotic microorganisms.

Masonry Walls with Termite Caps Solid Blocks and Filled Block Tops

Builders may construct a foundation wall with solid, filled, or sealed block tops for several reasons, including termite-proofing, energy conservation, distribution of weight of the structure, and radon resistance. The NCMA28 recommends that a solid or grouted top course be installed to distribute the loads of joists and beams. Some building codes require solid tops to block hidden termite entry. In spite of this, the block tops in many residences are left open except at anchor points. Houses have been observed in which block tops were generally sealed, but cores were left unsealed at access doors to crawlspaces, around ash pit doors, and other openings. Sealing hollow cores at or near their tops can prevent soil gas from entering the basement, but more importantly might make the building easier to mitigate in the event that it has elevated radon. Sealing the bottom course might prevent air beneath the slab from entering the block wall, but if the wall cores are used as part of a...


Each termite produces, on average, about 0.5 mg CH4 day - a seemingly insignificant amount. However, when this is multiplied by the global population of termites, CH4 emissions from this source are estimated to be about 20 Tg year. There are more than 2000 different species of termites and the amounts of CH4 produced vary considerably between species, with some producing no CH4 at all. CH4 is produced in termite guts, by symbiotic bacteria and protozoa, during food digestion. This CH4 does not always end up going straight into the atmosphere. Many species are subterranean or live in aboveground earth mounds where much of the CH4 can be used up by soil methanotrophs before it gets out into the atmosphere - soil-mediated

Soil structure and nutrients

Another feature of arid and semi-arid lands is the importance of macro-organisms in decomposition. Shachak et al. (1976) found that processing of surface soii by isopods in the Negev had a significant positive effect on the rate of decomposition of organic matter in the system. Termites (Wood and Sands 1978 Whkford 1991), and the presence of vertebrate burrows and nests, can have dramatic effects on the rate of decomposition (Sehaefer and Whitford 1981 Wbitford and Parker 1989) by moving material below-ground and or by moderating temperature and humidity. Thus the removal of one or more groups of species (e.g. all termites, or all burrowing rodents)

Lignocellulose degradation

There is a large body of literature on the decomposition of wood and cellulose by termite gut flagellates, which are essential for the digestion of lignocellu-lose in lower termite (Radek 1999). By contrast, the majority of prokaryotes in termite guts do not seem to contribute to polymer degradation. They appear to be involved mainly in the fermentation of soluble metabolites released into the gut, which are derived either directly from the food by the digestive enzymes or from the fermentative activity of the intestinal protozoa (Breznak 2000 Ohkuma 2003 Brune 2005). Lignocellulose is not only difficult to degrade, but it is also an extremely nitrogen-poor substrate that lacks most of the essential nutrients required by the termite, such as amino acids, vitamins, and sterols. The capacity of the intestinal prokaryotes to fix dinitrogen, to assimilate nitrate and ammonia, and to synthesize those amino acids and vitamins essential for the host makes them also an important source of...

Prokaryotic Diversity

As in most other environments, phylogenetic analysis of the 16S rRNA genes of the prokaryotes in the hindgut of several termites has revealed an enormous diversity of the intestinal microbiota (see Brune 2005 for a review). The majority of clones in bacterial clone libraries of several species in the genus Reticu-litermes bears less than 90 sequence similarity to those of known, cultivated microorganisms. Clones in the so-called Termite Group 1 (Ohkuma and Kudo 1996 Hongoh et al. 2003 Yang et al. 2005) even represent a new bacterial phylum, the 'Endomicrobia' (Stingl et al. 2005), which contains no cultivated representatives and apparently consists exclusively of endosymbionts of termite gut protozoa (see (8).4.2.2.). The fact that most clones recovered in these studies fall within lineages consisting exclusively of clones obtained from the hindgut of termites, with the closest relatives of a given clone usually stemming from the most closely related termite (Yang et al. 2005),...

Physiology and Function

The first studies on the anaerobic nature and the fermentation of cellulose by termite gut flagellates were performed by Hungate in the 1940s (reviewed by Hungate 1955). The concept is supported by the high hydrogen partial pressure and the dominance of acetate among the short-chain fatty acids in the hindgut of all lower termites investigated (reviewed by Radek 1999 Breznak 2000 Brune 2005). explanation why the well-being and survival of lower termites on a normal, lignocellulosic diet depend on their gut protozoa (Breznak 2000 Brune 2003, 2005). Despite the demonstration of host cellulases in the secretions of midgut and salivary glands of lower termites (see Watanabe and Tokuda 2001 for a review), the cellulolytic activities in the hindgut seem to be largely of protozoan origin (Ohtoko et al. 2000 Nakashima et al. 2002). However, data on the physiology of gut flagellates are scarce, and the current concept of cellulose metabolism by termite gut flagellates is largely based on...

Symbiotic Associations with Prokaryotes

It is long known that most of the gut flagellates are associated with prokaryotes (Pierantoni 1936 Kirby 1941 Ball 1969 Bloodgood and Fitzharris 1976 Dolan 2001). In view of the enormous biovolume of protozoa in the hindgut of lower termites and the resulting cell surface area available for colonization, it is not astonishing to find that the vast majority of the prokaryotes in the dilated part of the hindgut - approx. 90 in the case of Mastotermes darwinien-sis (Berchtold et al. 1999) - is associated with the protozoan fraction.

Functional and Metabolic Implications

The most intriguing question concerns the physiology and metabolic function of the symbiotic bacteria of the termite gut flagellates. Assuming that the growth rate of gut bacteria is not necessarily sufficient to compensate for the high dilution rates of the hindgut contents, it would be advantageous for prokaryotes to associate with the protozoa to prevent wash-out from this substrate-rich and favorable environment. The intimacy of many associations, especially the elaborate attachment structures, however, indicate that there is a mutual advantage.

Removal of Phosphate Using Feldspar

Feldspar, among many natural substances such as termite mount-clay, saw dust, kaolinite, and dolomite, offers significant removal ability for phosphate, sulfate, and color colloids. Optimization laboratory tests of parameters such as solution pH and flow rate, resulted in a maximum efficiency for removal of phosphate (42 ), sulfate (52 ), and color colloids (73 ), x-ray diffraction, adsorption isotherms test and recovery studies suggest that the removal process of anions occurs via ion exchange in conjunction with surface

Slow Going For A Few Million Years

Protohumans lived a wide-ranging hunting-gathering life, harvesting a wide variety of nuts and berries as they became seasonally available. Some of the shaped tools were probably used to dig out tubers and roots. Like modern chimps and some primitive people, they likely used sticks to extract termites from holes and eat them. The landscape also contained many other delicacies for those with the stomach for them bird eggs, field mice, and other fare.

The Intestinal Microbiota of Mastotermes darwiniensis

The lower wood-feeding termite Mastotermes darwiniensis Froggatt (Fig. 5.1.) is the only living member of the family Mastotermitidae. Today, this species is restricted to Northern Australia, but mastotermitid fossil specimen from the Eocene and Miocene have been found in Central America, the Caribic region, Europe, and Australia (Thorne et al. 2000). Termites are assigned to 13 families and 282 genera (Table 5.1 Myles 1999). Mastotermes darwiniensis is believed to be the most primitive existing termite species (Gay and Calaby 1970). Mastotermes darwiniensis developed a complex symbiotic hindgut flora, which consists of protozoa (formerly named Archaezoa Cleveland and Grimstone 1964 Brugerolle et al. 1994 Berchtold and K nig 1995 Fr hlich and K nig 1999a,b), Bacteria (Berchtold and K nig 1996 Berchtold et al. 1999), Archaea (Fr hlich and K nig 1999a,b) and yeast (Prillinger et al. 1996 Sch fer et al. 1996). De-faunation experiments showed that the protozoa appeared to be essential for...

Global emission estimates

Global Estimates Emissions Species

Thus, global geo-CH4 emission estimates, stemming from mud volcanoes plus other seeps, plus microseeps, plus submarine emissions, plus geothermal and volcanic emissions range from 42 to 64Tg yr-1 (mean of 53Tg yr-1), almost 10 per cent of total CH4 emissions, representing the second most important natural CH4 source after wetlands. Geo-CH4 sources would then also represent the missing source of fossil CH4 as recognized in the recent re-evaluation of the fossil CH4 budget for the atmosphere ( 30 per cent) (see Lassey et al, 2007 Etiope et al, 2008), which implies a total fossil CH4 emission much higher than that due to the fossil fuel industry. Global geo-CH4 emission estimates are on the same level or higher than other sources (such as biomass burning, termites, wild animals, oceans and wildfires) considered by the IPCC (Denman et al, 2007) (Figure 4.5).

The Gut Microenvironment

Initially assumed that the termite hindgut is an anoxic habitat (for literature, see Breznak 2000 Brune 2005). However, the maintenance of anoxia in an environment of such minute dimensions is not a trivial issue (Brune 1998). Oxygen microsensor measurements in the gut of the termite Reticulitermes flavipes have demonstrated that the steep gradient in oxygen partial pressure between the oxic gut epithelium and the anoxic gut contents drives a continuous influx of oxygen into the hindgut (Brune et al. 1995 Ebert and Brune 1997). Also, in all other termites investigated, oxygen penetrates 50-200 m into the periphery of the hindgut lumen, leaving only the central portion of the dilated compartments anoxic (for literature, see Brune 2005). The hydrogen partial pressure in the different gut regions seems to be controlled by the spatial organization of the hydrogen-producing and hydrogen-consuming populations (Tholen and Brune 2000). Radiotracer analysis of the in situ metabolism in the...

Effect Of Species Diversity On Ecosystem Function

Several species of African grasses (such as Hypharrenia rufa, Melinis minuli-flora and Panicum maximum) have becomc naturalized in the South American llanos and cerrado and in the Australian savannas. A number of important functional differences exist between African invaders and native grasses. Thus, there are many possible functional outcomes that may arise at the ecosystem level as a result of the establishment of these species in American and Australian savannas, through their effects on herbivory, hydrology, decomposition and nutrient cycling. Invaders may also initiate new successional processes due to their effects on abiotic and biotic processes. First it must be demonstrated that the invasions are promoted by absolute differences in competitive abilities and not by disruptions in nutrient cycling, which in turn result from the removal of native species by human activities. Indeed, most invasions in South America have followed such primary disturbance of the soil and...

Atmospheric Physics And Thermodynamics

Thermodynamic Atmosphere Profile

Principal sources of methane are landfills, marshes and wetlands, agriculture and livestock, natural gas, biomass burning, and life forms such as termites. It is removed by oxidation and photodissociation in the stratosphere. Individual molecules of methane have 20-30 times the global-warming potential of CO2 and they are accumulating in the atmosphere much more rapidly, meaning that methane could overtake carbon dioxide as the principal anthropogenic contribution to global warming.

Prokaryotic Epibionts

Epibiont Bacterial

Many of the termite gut flagellates are colonized by epibiotic bacteria. Some of the associations are already evident by observation with a light microscope, but only the electron microscope reveals the enormous variety and complexity of these associations (e.g., Cleveland and Grimstone 1964 Leander and Keeling 2004 Figs. 1-3). The presence of special attachment sites on the cell envelope of the flagellates (e.g., Tamm 1980 Radek et al. 1992 Dolan and Margulis 1997 Stingl et al. 2004) indicates a tight association. In some cases, the epibionts seem to be responsible for the locomotion of the host (Cleveland and Grimstone 1964 Tamm 1982 see also Chapter by H. K nig, L. Li, M. Wenzel, and J. Fr hlich). Advances in molecular ecology made it possible to elucidate the identity of the microorganisms involved in these associations and the specificity of these symbioses. Fig. 3. Transmission-electron micrographs of ultra-thin sections of Trichonympha agilis (A) and Pyrsonympha vertens (B)...

Biology And Ecology Of Soil Fauna

Other groups of arthropods that occur commonly in soil are pseudoscorpions, symphylans, pauropods, proturans, diplurans, and the immature stages of holome-tabolous insects (Dindal, 1990). Ants and termites can also be very numerous however, these macroarthropods will not be considered here (Brian, 1978).

Morphology of Mixotrixa paradoxa

Microbes Protists

Of spirochetes with the exception of the posterior ingestive zone (Cleveland and Grimstone 1964). The rod-shaped bacteria and the spirochetes are attached to regularly arranged protrusions of the cell surface. Interestingly, Cleveland and Grimstone (1964) found that the spirochetes (not the relatively small four flagella) propel the cells. It is still unknown how the flagellates and the spirochetes communicate and coordinate the direction of movement. For hydro-mechanical reasons, it seems that cilia, flagella, sperm tails and spirochetes should automatically synchronize their movement when undulating in close proximity (Machin 1963). So far, it is not possible to cultivate either Mixotricha paradoxa or its ectosymbiotic spirochetes and rod-shaped bacteria. Although spirochetes are always a dominant part of the microflora of all termites (Margulis and Hinkle 1992), only four species have been obtained in pure culture (Leadbetter et al. 1999 Dr ge 2006).

Glycolytic Activities of Mastotermes darwiniensis and its Flagellates


Up to now, it has been believed that the hindgut flagellates produce nutrients using their own cellulolytic enzymes for the benefit of their termite host. The glycolytic activities found in separated cells of Mixotricha paradoxa are compiled in Table 5.3 (Berchtold and K nig, unpubl. res.). Surprisingly, not all of these activities seemed to be produced by the flagellates themselves, but rather taken up with the gut contents. Two endoglucanases, Cel I and Cel II, with the molecular mass of approximately 48 kD, were isolated from the not yet culturable symbiotic flagellates living in the hindgut of the most primitive Australian termite Mastotermes darwiniensis (Li et al. 2003). The N-terminal sequences of these cellulases exhibited significant homo-logy to cellulases of termite origin, which belong to glycosyl hydrolase family 9. The corresponding genes were detected not in the mRNA pool of the flagellates, but in the salivary glands of Mastotermes darwiniensis. A protein with the...

Socially and Economically Valuable Trees

Deciduous Neem tree (Azadirachta indica) symbolises a body of traditional values, knowledge, and uses. Almost all parts of the tree are used in medicine and agriculture.The leaves are used in traditional health systems, in religious rituals, and as green manure in agriculture. Oil is extracted from the seeds and has both medicinal and pesticidal properties. Neem cake which is a by-product of oil extraction is used as an organic fertiliser. The wood has a high calorific value as fuelwood. Neem wood is termite resistant and used to make door and window frames. Species with multiple values may be candidates to drive region-specific restoration of these species (and others) within the broader landscape.

Characterization Of Biodiversity In Arid Lands

Certain taxa are diverse in arid lands relative to other biomcs these include predatory arthropods, tenebrionid beetles, ants and termites, snakes and lizards, and annual plants (Cloudsley-Thompson 1975 Crawford 1981 Wallwork 1982 Pianka 1986 Ludwig et al 1988). However, there is substantial variation in the richness of particular taxa among the deserts of different continental areas, suggesting that biogeography and historical factors are as important as ecological factors. For example, predatory arthropods (spiders, scorpions, solpugids) are diverse in North America but not in Australia, while for ants that pattern is reversed.

Relationships Between Biodiversity And Function In Agroecosystems

Magnitude of local effects created by the individual within the spatial and temporal patterns of the species population in the system. The functional properties of individual species are therefore determined by the body size and activity of individuals, and the density and aggregative characteristics of the population. In the case of soil organism communities, for example, eollembola or mite species, feeding on fungi and bacteria at a scale of millimetres or less, form aggregations affecting microbial processes within a volumetric decimeter or more earthworms, forming burrows of a few millimetres in diameter, have aggregative effects on hydrologic processes in patches of several metres fungal hyphae, active at the cellular level, can be a component of a single genetic individual dominating wood decomposition over many hectares with a biomass of tonnes littcr-feeding termites (Macro-terminae) determine soil physical and chemical processes from the plot to landscape scale (Anderson...

The Drilosphere As A Selforganizing System

Soil constraints indeed have pushed soil organisms to develop intense interactions along evolutionary time, mostly of a mutualistic type (Lavelle and Spain 2006). These interactions operate within the boundaries of the rhizosphere of roots, drilosphere of earthworms, and termitosphere of termites, as well as a few other such domains (Lavelle 2002) that have

Atmospheric properties and feedbacks

Arid lands are significant determinants of the earth's albino and thus of its global radiation balance. Albedo and spectral characteristics of the surface are influenced not just by total plant cover, but also by the different properties of woody plants versus grass cover, so changes in plant functional group affect this global property. Arid lands are also significant contributors of dust (Pewe 1981 Pye 1987), which influences both the radiation balance of the atmosphere and the transport of N, S, Fe and other minerals over long distances. The extent to which changes in biota influence soil vulnerability to wind movement will determine the importance of these changes to those atmospheric properties. Desert soils, serving as possible sites for carbonate formation and having low organic matter content, represent important potential sinks for atmospheric carbon. It has also been suggested that deserts contribute significant amounts of methane to the atmosphere (due to termites), and...

Particle Size Distribution

Types of compounds detected include the alkanes from motor vehicle fuel, aldehydes, esters from plasticizers, organic acids, pesticides from agricultural runoff and termite control, and PAHs and volatiles from fuels. In particular, the pesticides and PAH fluoranthene are detected in high concentrations. The concentrations of chlordane and dieldrin, two common pesticides, are both 100 times greater than the EPA soil regulatory standard (II). The concentration of the PAH fluoranthene is nearly 20 times that detected in an outdoor sample located between the two flooded homes (21). These results, combined with the comparison of indoor and outdoor sediment metal concentrations, lends further credence to the theory that in-home sediment contaminants may be present at higher loadings as a result of the particle winnowing process. The high loadings of pesticides relative to their EPA screening standard, along with the large increase in fluoranthene concentration compared to the exterior...

Scalingup calculations and global CH4 budgets

Scaling-up calculations by Martius et al (1996), Sanderson (1996), Bignell et al (1997) and Sugimoto et al (2000) used differing assumptions and principles, but all four agree that CO2 fluxes by termites are in the range 2-5 per cent of the total from all terrestrial sources. 2 per cent is a large figure for a single insect order with about 0.01 per cent of the global terrestrial species richness, but is nevertheless only a minor source in the context of the whole C budget. With CH4 emission, it now appears that the net contribution by termites is very small, despite the fact that some trophic groups produce this potent greenhouse gas in large amounts at the point scale. Independent studies using, respectively, static chambers and natural stable isotope ratios showed that mound walls and undisturbed soil between mounds have a sink capacity exceeding production at the landscape level, presumably due to the presence of methylothrophic bacteria (Sugimoto et al, 1998b MacDonald et al,...

Biochemistry and microbiology of methane production

A detailed discussion of the process of CH4 production by microorganisms is beyond the scope of this chapter (see Chapter 2). Excellent recent reviews of the relevant microbiology and microbial physiology can be found in Breznak (2006), Brune (2006), Breznak and Leadbetter (2006) and Purdy (2007). The more general phenomenon of CH4 production by terrestrial arthropods is reviewed by Hackstein et al (2006). Although the broad basis of lignocellulose digestion by termites has been known for more than 60 years (Hungate, 1946), numerous refinements continue to be added to the picture.

Harnessing The Drilosphere To Restore Ecosystem Functions In Degraded Soils

Munities generated by exceptional rates of endemism in highly sedentary organisms. In Amazonia, for example, the average ratio of local species richness to regional richness has been estimated at approximately 1 , compared to 20-30 on average for ants and termites and 80 for Sphingidae moths (Lavelle and Lapied 2004).

The Overarching Role of Spatial and Temporal Scale

Intermediate phase in fresh biogenic structures. Microbial activation triggered during gut transit or mechanical mixing of organic materials with soil culminates in fresh biogenic structures, such as fresh earthworm casts or termite fecal pellets. Activity then progressively decreases during the few days or weeks following deposition.

Connecting Soil and Sediment Biodiversity The Role of Scale and Implications for Management

A unique feature of soil and sediment systems is the range of biological activities (e.g., burrowing, tube construction, substratum translocation, and feeding processes), that sculpt and mold the substratum, generating its fine structure and heterogeneity. The diversity of ecosystem engineers is key, producing a greater variety of biogenic structures (e.g., termite mounds and galleries, earthworm casts, and burrows in soils tubes, surface deposits, and stable aggregates cf. ray and whale pits made by bioturbators in aquatic environments). Specific interactions among their populations are thus essential to the shaping of the aggregate and pore structure of the soil and sediment substrata. Structure, in turn, determines the ability of the substratum to accommodate and sustain microorganisms that perform specific chemical transformations, and to support other functions and processes. In soils, for example, the combined activities of com (roots and soil invertebrates), through their...

Water distribution and quality

Because water is the primary limiting resource for many organisms in arid systems, changes in the biota which translate into changes in water distribution or availability will be strong drivers of a change in state. One critical stage is the infiltration of water into the soil (versus its evaporation from the surface or its horizontal transfer or loss to run-off). Vegetative cover modulates the impact energy of raindrops, reducing the amount of sediment dislodged and transported during heavy storms (e.g. Rogers and Schumm 1991). The presence of rooted plants provides root channels which in turn enhance deep percolation of water into the soil profile the nature of the plant canopy influences the proportion of rainfall that is intercepted and that falls either as throughfall or as stemfiow (West and Giftbrd 1976 Tromble 1987 Navar and Bryan 1990). Stemfiow apparently redistributes precipitation to the deep roots of shrubs and trees, favoring established vegetation at the expense of...

Agricultural intensification

The key factors determining total biodiversity in the different agroecosys-tems shown in Figure 11.2 are microclimate, habitat structure and food resources. Hence forest plantations and home gardens retain many of the community characteristics of natural forest, and pasture communities are functionally similar to savannas. The pattern of species losses in the soil biota along the gradient of agricultural development approximates that of Curve IV in Figure 11.2. The point of inflection from more to less diverse communities varies, however, for different sizes and functional groups according to the type of farming system and agricultural practice. In the extreme situation of direct conversion of forest to intensive cultivation of short-rotation crops, there is rapid disappearance of surface-active macro-invertebrates (millipedes, earthworms, beetles) which use leaf litter as both a habitat and a food resource (Lavelle et al. 1994). As the mass of soil and litter organic matter pools is...

Symbiotic Interactions between Spirochetes and Flagellates

One of the first detected symbioses between flagellates (Pseudodevescovina uniflagellata) and spirochetes was described by Kirby (1936). Pseudode-vescovina uniflagellata lives in the gut of the Australian dry wood termite Neotermes insularis. Only three years earlier Sutherland (1933) published an article about Mixotricha paradoxa where the attached spirochetes were misconceived as cilia. A detailed description of the fine structure of Mixotri-cha paradoxa and the role of the ectosymbiotic bacteria in cell locomotion was provided by Cleveland and Grimstone (1964). Over the years, more and more examples of surface symbiosis between protists and prokaryotes from the termite gut appeared (Bloodgood and Fitzharris 1976 Smith et al. 1975 To et al. 1980 Dyer and Khalsa 1993), but examples of motility symbiosis in the termite gut could be rarely detected (Tamm 1982). Locomotory mechanisms of two larger flagellates from Mastotermes darwiniensis have been studied (Cleveland and Cleveland 1966...

Sources And Sinks

Termites methane has been called marsh gas. Other natural sources are generally small but not well constrained. These include termites, oceans, and lakes. Most of the current sources are anthropogenic. While these emissions are not directly from stacks and other easily identifiable icons of man-made pollution, they are a result of human activities nonetheless. These sources may be classified mostly as agricultural and from use of energy. Of these, rice agriculture, cattle, waste management, biomass burning, coal mining, and use of natural gas are the largest contributors. There are some moderate sized sources of a few teragrams year that include transportation and fossil fuel combustion. There are perhaps many small sources that together fall within the range of uncertainty of the global emission rate and are therefore not included.

Sealevel changes

From grazing animals and termites, whereas nitrogen oxides are increasing because of the increased use of fertilizers and automobiles, and chlorofluorocar-bons are increasing as a result of release from aerosols and refrigerants. Together the greenhouse gases have allowed short-wavelength incoming solar radiation to penetrate the gas in the upper atmosphere but trapped the solar radiation after it is reemitted from the Earth in a longer wavelength. The trapped radiation causes the atmosphere to heat up, leading to greenhouse warming. other factors also influence greenhouse warming and cooling, including the abundance of volcanic ash in the atmosphere and solar luminosity variations, as evidenced by sunspot variations.


Methane is an odorless, non-toxic, flammable gas consisting of one carbon and four hydrogen atoms (CH4). Mainly anaerobic or oxygen-deprived processes produce methane. It forms when the digestion of organic material by bacteria releases single carbon atoms. The main sources of methane are wetlands, the combustion of fossil fuels, animals, rice plantations, biomass burning, landfills, termites, and the oceans. Methane is removed from the atmosphere primarily by reaction with hydroxyl radicals (OH). The atmospheric concentration of methane has also been steadily increasing since the beginning of the Industrial Revolution. The atmospheric concentration of CH4 has been increasing at the rate of about 1 percent per year.


Methane is released into the atmosphere by a number of anthropogenic and natural sources, but anthropogenic emissions used to dominate present-day CH4 budgets, accounting for more than 60 of the total global budget. The anthropogenic sources include rice agriculture, livestock, landfills and waste treatment, some biomass burning, and fossil fuel combustion, while natural sources are wetlands, oceans, forests, fire, termites and geological sources. Estimates of global CH4 sources by various workers have been reflected in Table 16.1. Natural sources Wetlands Termites Oceans Hydrates

Water Pollution

The timber industry treats timber and wood products with chemical preservatives to protect the wood from degradation due to various organisms including fungi, and insects such as borers and termites. This treatment extends the range of applications and the service life of the wood. By design, the chemicals used to protect wood must be toxic to the target organisms, but they may also affect nontarget organisms and the environment 1 .


Abbadie L, Lepage M, Le Roux X (1992) Soil fauna at the forest-savanna boundary role of termite mounds in nutrient cycling. In Furley PA, Proctor J, Ratter JA (eds) Nature and Dynamics of Forest-savanna Boundaries. Chapman & Hall, London pp 473-484 Occasional Papers 9, Oxford Howse PE (1992) The chemical ecology of forest and savanna termites. In Furley PA, Proctor J, Ratter JA (eds) Nature and Dynamics of forest-savanna boundaries. Chapman & Hall, London, pp. 485-498 Hoyle AC (1934) New trees and shrubs from tropical Africa III. Kew Bull. 188, 190 Huisman J, Olff H, Fresco LFM (1993) A hierarchical set of models for species

Natural sources

Major natural sources include wetlands, termites and release from onshore and offshore geological sources. Recently, living vegetation has also been suggested as an important natural source of CH4. Of the globally significant sources of CH4 to the atmosphere, natural sources are currently outweighed by anthropogenic sources. Together they emit some 582Tg CH4 each year, with 200Tg arising from natural sources (Denman et al, 2007). Given the estimated global CH4 sink of 581Tg per year, the current increase in atmospheric CH4 concentrations should, therefore, be only 1Tg CH4 per year. But even with ongoing efforts to reduce anthropogenic emissions, and so arrest the trend of increasing CH4 concentrations in the atmosphere, future enhancement of natural CH4 emissions due to climate change threatens to negate some, or all, of these attempts at mitigation. Termites Though some termite species produce no CH4 at all and those that do rarely exceed more than half a microgram per termite day,...

The Endomicrobia

For a long time, the nature and identity of the endosymbionts of the larger gut flagellates was completely obscure. Recently, the cytoplasmic symbionts of the hypermastigid Trichonympha agilis and the oxymonad Pyrsonympha vertens in the gut of Reticulitermes santonensis were identified as members of the so-called 'Termite Group 1' (TG-1) (Stingl et al. 2005). Based on their 16S rRNA gene sequences, whose origin was verified by fluorescence in situ hybridization with clone-specific probes, and their peculiar ultrastructure (Fig. 3), the endosymbionts of the two protozoa were provisionally classified as two species in the candidate genus 'Endomicrobium' (Stingl et al. 2005). The symbionts are specific for their respective flagellate host and do not occur on the surface of the flagellates or freely suspended in the gut fluid. Up to that point, 16S rRNA gene sequences of TG-1 bacteria had been retrieved only from termites of the genus Reticulitermes (Ohkuma and Kudo 1996 Hongoh et al....

Energy flow

Termites, larvae of wood-boring beetles, girdling beetles Larvae of cicadas and beetles Unlike most other nutrients, the major sources of nitrogen to ecosystems are precipitation and biological nitrogen-fixation by free-living bacteria and cyanobacteria, by bacteria having mutualistic associations with plants, by fungi, and by gut-dwelling symbionts of termites (Prestwich et al. 1980 Prestwich and Bentley 1981). In species-poor systems, such as those growing on young tropical lava flows, invasion of a single tree species and lichenized fungi with nitrogen-fixing bacteria may dramatically increase nitrogen input to the system, productivity and ecosystem development (Vitousek et al. 1987 Vitousek and Walker 1989). However, whether the quantities of nitrogen entering and cycling with tree-species-rich tropical forests are influenced by the number of species of free-living or symbiotic nitrogen-fixing microorganisms is unknown. Atmosphere-soil Soil microorganisms, like the macroorganisms...

Hydrogen Metabolism

The symbiosis with methanogens is likely based on the interspecies transfer of molecular hydrogen. Little is known about the physiology of the protozoa, but from thermodynamic considerations, any fermentative metabolism releasing part of the reducing equivalents as H2 will become more exergonic if this product is not allowed to accumulate (Schink 1997 see also Chapter by B. Schink). Virtually all 16S rRNA gene clones of methanogens obtained from lower termites fall into the phylogenetic radiation of the genus Methanobrevibacter. Since all members of this genus, including the three species isolated from the gut of Reticulitermes flavipes, perform methano-genesis from H2 and CO2, but use other substrates less efficiently, it is reasonable to assume that also the hitherto uncultivated Methanobrevibacter strains associated with the protozoa are potential hydrogen consumers (Lee et al. 1987 Tokura et al. 2000). It is therefore possible that the bacterial symbionts of the larger protozoa...

Alan P Covich

Various saprophytic and litter feeding invertebrates (detritivores), fungi, bacteria, actinobacteria, and other microorganisms Diverse soil organisms, e.g., earthworms, termites, fungi, eaubacteria, etc. Biological nitrogen fixation by diazotroph bacteria, conversion of NH4 to NO3 by nitrifying bacteria, conversion of NO3 to N2 by denitrifying bacteria Maintaining biodiversity in soils and water is imperative to the continued and improved effectiveness of bioremediation and biotreatment Nearly half of the current economic benefit of biotechnology related to agriculture involves nitrogen-fixing bacteria, pharmaceutical industry, etc. Soil provides microhabitats for natural enemies of pests, soil organisms (e.g., mycorrhizae) that contribute to host plant resistance and plant pathogen control

Ronald G Prinn

The major sources of methane are biological (basically methanogens operating in oxygen-poor environments like natural wetlands, rice paddies, digestive systems of termites and cattle, animal waste treatment, sewage, landfills, etc.). Another (indirect) biological source is biomass burning. Many of these biological sources (rice, cattle, biomass burning, etc.) are governed largely by human activity. Also under dominant human influence is another significant CH4 source, namely escape of this gas during mining of coal and natural gas and leakages in natural gas distribution systems. The fossil sources can be differentiated from the biological sources because of the different carbon isotopic signatures of each, which show that roughly 20 percent of global total methane emissions come from fossil sources. The total methane source is estimated to be about 600 Tg y-1 with 60 percent due to human activities (Kurylo et al. 1999 Prather et al. 2001).

Widely Disputed

The global methane cycle is comprised of a wide range of sources and sinks. Major natural sources include wetlands, termites, the oceans, and release from hydrates (lattice-like structures of ice and methane, also known as clathrates, occurring in polar regions and in oceanic sediments). Recently, living vegetation has also been suggested as an important natural source of methane. Of known sources, wetlands dominate natural emissions, with between 100 and 200 million tons of methane emitted from these waterlogged soils each year. Wetland methane emissions arise from methane-producing bacteria known as methanogens, which are active in the anaerobic, carbon-rich environments common to wetland soils.

Arable Land

In intensive tilled agriculture, short rotations and the use of high-yielding crops can trigger development of soil-borne pathogens and viruses, nematode infestations, and soil-dwelling insects, causing major yield losses. Overcoming these outbreaks requires the breeding of resistant crop varieties, the frequent use of soil pesticides (Epstein & Bassein 2001), or the development of novel biological control practices (Whipps 2001). In contrast, in traditional small-holder (predominantly tropical) farms, the role of organisms is apparent and the activities of a few species of earthworms, termites, and other fauna dominate soil communities. After hand tillage (mostly on tropical farms) or abandonment of the site in shifting agriculture, soil earthworm communities usually recover rapidly (Decaens & Jimenez 2002). This is in contrast with the microbial recovery of soil communities after intensive tilled farmed systems in temperate zones (Siepel 1996 Korthals et al. 2001).


In unmanaged humid tropical forests it is rare that any specific identifiable group of organisms have a role in soil processes. Litter breakdown is accomplished by many disparate groups of organisms such as crabs, millipedes, cockroaches, and so on. Within these groups there is always some marginal overlap in feeding-niche parameters. Similarly, soil-feeding termites (which exhibit particularly high species diversity in African forests) produce significant methane fluxes and play an important role in P cycling and soil organic matter turnover. However, all these processes are also mediated by other animals


Although spirochetes are extremely abundant in the intestinal tract of most wood-feeding termites, only a few species have been characterized in pure culture (Breznak and Leadbetter 2002 Graber et al. 2004). Termite gut spiro-chetes display an enormous morphological diversity (Margulis and Hinkle 1999 Breznak and Leadbetter 2002), but several cultivation-independent analyses have documented that they belong exclusively to the Treponema branch of the Spirochaetes, forming two distinct clusters (Lilburn et al. 1999 Ohkuma et al. 1999c).

Hindgut Protozoa

Symbiotic flagellates are found exclusively in the phylogenetically lower termites and the closely related cockroaches of the genus Cryptocercus the higher termites harbor a largely prokaryotic microbiota. The first studies that revealed the beneficial nature of these peculiar symbionts - discovered in 1856 by Lespes (see Leidy 1881) and initially considered as parasites - were reported by Cleveland (1925, 1926), who demonstrated that termites do not survive for long after elimination of their gut flagellates. The importance of the symbionts for their termite host is impressively documented by their enormous abundance in the enlarged hindgut paunch it has been estimated that the fresh weight of the protozoa may account for more than one half of the fresh weight of the termite (Katzin and Kirby 1939). Phylogenetically, the gut flagellates are extremely diverse almost 450 distinct species have been described to occur within the approx. 200 termite species investigated (Yamin 1979).


Termite gut flagellates belong to three distinct taxa trichomonads, hypermas-tigids, and oxymonads (Yamin 1979). Originally, all were considered primitive, primarily amitochondriate eukaryotes, but recent molecular data proved that they represent two separate eukaryotic lineages. Phylogenetic analyses of the termite gut flagellates by various groups (reviewed by Ohkuma 2003 Gerbod et al. 2004), mainly based on 18S rRNA gene sequence analysis, confirmed the presence of two classes of the phylum Parabasalia, the Trichomonadea and Hypermastigea (Cavalier-Smith 2002), the latter representing the most basal lineage. While Hypermastigea consist exclusively of termite gut symbionts, the Oxymonadea and the Trichomonadea comprise also species occurring in other habitats, such as the intestinal tract or body cavities of other animals, including humans.


Methanogenic archaea in termite guts are easily visualized by the characteristic autofluorescence of their coenzyme F420. While the methanogens in the hindgut of Reticulitermes flavipes are mostly attached to the gut wall or to the surface of other prokaryotes (Leadbetter and Breznak 1996), they are also associated with protozoa in other termites, either as epibionts (Fig. 4) or as endosymbionts (see below) of certain smaller gut flagellates (Lee et al. 1987 Tokura et al. 2000). Most of the clones of methanoarchaea retrieved from lower termites cluster among the family Methanobacteriaceae (for references, see Ohkuma et al. 1999b Tokura et al. 2000 Shinzato et al. 2001), and are closely related to the three Methanobrevibacter species that have been isolated from the gut of Reticulitermes flavipes (Leadbetter and Breznak 1996 Lead-better et al. 1998). They are assumed to participate in the consumption of H2 formed by the flagellates (see Sect. (8.)5).

Functional Aspects

Obtained from an enrichment culture grown on a very rich medium (M.A.Cambon-Bonavita, unpubl. data). The free-living, marine spirochetes Spirochaeta isovalerica and S. litoralis consistently form a neighboring clade of the oligochaete spirochetes. These bacteria were isolated from sulfidic muddy sediments and are obligate anaerobes that ferment carbohydrates mainly to acetate, ethanol, CO2, and H2 (Hespell and Canale-Parola 1973 Harwood and Canale-Parola 1983). While fermentation is one possible metabolic pathway of the oligochaete spirochetes, they could also have a completely different metabolism, just as the spirochete symbionts in termites do not possess properties common to their closest free-living relatives within the genera Treponema. Instead, termite spirochetes were recently discovered to be chemoautotrophic, using H2 and CO2 to produce acetate (Leadbetter et al. 1999), and were also shown to be able to fix nitrogen (Lilburn et al. 2001). These types of metabolism would...


The large number of symbioses among termite gut flagellates and prokaryotes, the high level of integration evidenced by the elaborate attachment structures or the intracellular location, and the large proportion of the prokaryotic gut microbiota that is associated with the protozoa suggest a major importance of such symbioses for the hindgut metabolism of lower termites. In order to understand the hindgut metabolisms of lower termites, it will be essential to elucidate the metabolic relationship between the flagellates and their symbionts. It is reasonable to assume that the functional roles of the partners are less diverse than their phylogenetic diversity, and in view of the possible co-evolution of the partners, the symbioses between prokaryotes and gut flagellates are also excellent case studies in the microbial ecology and evolution.


The surface of many of the larger gut flagellates is densely covered with morphologically similar rod-shaped bacteria (Bloodgood and Fitzharris 1976 Tamm 1982 Dolan and Margulis 1997 Stingl et al. 2004 Fig. 1). The epibi-onts of a Staurojoenina sp. from Neotermes cubanus were recently identified as members of a new, termite-associated lineage of Bacteroidales (Stingl et al. 2004) and have been assigned to the candidate taxon 'Vestibaculum illiga-tum' (Fig. 5), and also the epibionts of Barbulanympha and Caduceia seem to be members of the Bacteroidales (see Stingl et al. 2004). Also the rod-shaped epibionts of Mixotricha paradoxa (Wenzel et al. 2003 see also Chapter by H. K nig, L. Li, M. Wenzel, and J. Fr hlich) fall into this cluster, and it is likely that the other clones in the cluster recovered from other termites (Ohkuma et al. 2002 Hongoh et al. 2003 Yang et al. 2005) represent Fig. 5. Phylogenetic relationships between the 16S rRNA gene sequences of 'Candida-tus Vestibaculum...


Unlike soil macrofauna (e.g., earthworms, termites, ants, some insect larvae), mesofauna generally do not have the ability to reshape the soil and, therefore, are forced to use existing pore spaces, cavities, or channels for locomotion within soil. Habitable pore space (voids of sufficient size and connectivity to support mesofauna) accounts for a small portion of total pore space (Hassink et al., 1993b). Microfaunal community composition becomes increasingly dominated by smaller animals as average pore volume decreases. Within the habitable pore space, microbial and mesofaunal activity is influenced by the balance between water and air. Maximum aerobic microbial activity occurs when 60 of the pore volume is filled with water (Linn and Doran, 1984). Saturation (waterlogging) and drought are detrimental to soil faunal communities because these conditions result in anaerobiosis or dehydration, respectively.