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Termites: An Overview

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A description of termite biology, distribution and diversity, economic importance, and sustainable management is presented. Liquid termiticide injection to soil, to establish a toxic or repellent chemical barrier against termites, is a traditional method applied for control. Baiting programs have been used successfully to eliminate subterranean termite colonies. Biological approaches along with entomophagy are also effective to manage termite population.
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1© Springer International Publishing AG 2018
M.A. Khan, W. Ahmad (eds.), Termites and Sustainable Management,
Sustainability in Plant and Crop Protection,
https://doi.org/10.1007/978-3-319-72110-1_1
Chapter 1
Termites: AnOverview
Md.AslamKhan andWasimAhmad
Contents
1.1 Introduction 2
1.2 Biology and Behavior 3
1.3 Systematics, Distribution, and Diversity 6
1.4 Invasive Termites 8
1.5 Termite-Gut Microbiota 9
1.6 Feeding Groups 10
1.7 Biotic and Abiotic Factors 10
1.8 Economic Importance 11
1.9 Prevention and Control 13
1.10 Conclusion 16
References 16
Abstract A description of termite biology, distribution and diversity, economic
importance, and sustainable management is presented. Liquid termiticide injection
to soil, to establish a toxic or repellent chemical barrier against termites, is a tradi-
tional method applied for control. Baiting programs have been used successfully to
eliminate subterranean termite colonies. Biological approaches along with ento-
mophagy are also effective to manage termite population.
Keywords Termites • Economic importance • Management • Control
M.A. Khan (*)
Department of Biology, Faculty of Science, Jazan University, Jazan, Saudi Arabia
e-mail: mdaslam30@gmail.com
W. Ahmad
Department of Zoology, Section of Nematology, Aligarh Muslim University, Aligarh, India
mdaslam30@gmail.com
2
1.1 Introduction
Termites are dominant invertebrate decomposers of dead organic matter in tropical
and subtropical regions (Bignell and Eggleton 2000). Their ecological success is
often attributed to the combination of a sophisticated social organization with
unique ability to feed on recalcitrant plant matters such as wood (Bignell et al.
2011). The phylogenetic position of termites has been long debated. They constitute
an ecologically and evolutionary diversied group of social insects that share a
common ancestry with cockroaches (Inward etal. 2007a). Termites play an impor-
tant role in ecosystems, with a major inuence on soil chemical and physical struc-
ture, plant decomposition, nitrogen and carbon cycling, and microbial activity (Holt
and Lepage 2000). While in temperate zones termites play a minor ecological role,
in the tropics they are the most important invertebrate decomposer (Bignell and
Eggleton 2000; Bignell etal. 2011). In tropical ecosystems, termites often make up
over 10% of the total animal biomass and up to 95% of soil insect biomass (Jones
and Eggleton 2000) and are considered to enhance ecosystem productivity
(Bourguignon et al. 2016). They may reach enormous population density in the
tropics, sometimes up to 1000 individuals per square meter (Eggleton etal. 1996).
These insects are key species in ecosystems as they recycle a large amount of nutri-
ents, but they are also pests, exerting major economic impacts. Among the eusocial
taxa, ants and bees are by far the most studied, whereas termites have received much
less attention in spite of their comparable abundance. Similarly, a few termite clades
attract the attention of most researchers, while others are almost entirely neglected
(Bourguignon etal. 2016). Our global view of the termite world is thus strongly
biased toward a few economically important genera that make up approximately
12% of the described termite species (Krishna etal. 2013a) while overlooking other
ecologically important and diverse taxa.
Soil is one of the most complex and species-rich habitats, hosting a wide range of
life forms. Termites form eusocial societies and live in colonies, creating nest sys-
tems that may be underground, epigeous, or arboreal. Based on habitat, termites can
be grouped into three general categories: subterranean, dry-wood, and damp- wood
termites (Paul and Rueben 2005). Subterranean termites live in soil and in wood that
is in contact with soil (Fig.1.1). The name subterranean comes from the strong need
of moisture in their environment that is satised by nesting inside or in close contact
with the soil (Thorne 1998). Subterranean termites are major structural pests causing
tremendous amounts of damage (Su and Scheffrahn 1998) and are reported as
responsible alone for at least 80% of losses caused by termites (Su and Scheffrahn
1990). Dry-wood termites live entirely in the wood, both nesting and feeding there.
Since they have the ability to thrive in wood with low moisture content, they may
attack all kinds of dead and dry wood, such as structural timbers, furniture, ooring,
and other wooden articles (Myles etal. 2007). Damp-wood termites, however, live
inside the wood of varying levels of decay and moisture content.
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1.2 Biology andBehavior
A termite colony is usually founded by a pair of alates (winged), the primary repro-
ductives, which produce all the nestmates. In some species, secondary reproductives
appear to either replace the primaries or supplement colony reproduction (Haig
etal. 2016). Inside termites’ complex society, the individuals are morphologically,
physiologically, and behaviorally specialized into distinct castes (Figs.1.2a, 1.2b,
and 1.2c). The castes work together to accomplish specic and complementary
tasks within a colony. Division of labor among castes is the key to efcient colony
development, survival, and reproduction. It may take 4–6 years for an incipient
colony of Coptotermes formosanus Shiraki to reach maturity and produce alates
(Chouvenc and Su 2014). As social insects, mature Coptotermes colonies can reach
more than a million individuals (Su and Scheffrahn 1988) with caste polymorphism
and polyethism (Chouvenc and Su 2014). They have underground foraging galleries
reaching up to 100m, making detection and control difcult (Su and Scheffrahn
1998). Colonies of the desert subterranean termite, Heterotermes aureus (Snyder),
have been estimated to include as many as 300,000 individuals around structures in
urban environments (Baker and Haverty 2007). Seasonal variation in caste distribu-
tion of foraging populations of the subterranean termite, Reticulitermes avipes
(Kollar), was recorded. Workers were most abundant in the spring and summer
months, and soldiers were most abundant immediately preceding alate ights
(Howard and Haverty 1981). In termites, foraging is usually performed by blind
castes, and the communication within individuals is mediated mainly by phero-
mones (Costa-Leonardo and Haig 2014).
Observation of the behavioral repertoire of some more derived termite species
with large colony size and extended nesting type (Abe 1987) remains challenging
Fig. 1.1 A live colony of subterranean termite
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Fig. 1.2b Castes (reproductive, soldiers, and pseudergates– immature reproductives) of the West
Indian dry-wood termite, Cryptotermes brevis (Walker) (Photo courtesy: Scheffrahn RH,
University of Florida)
Fig. 1.2a Life cycle of the Formosan subterranean termite, Coptotermes formosanus Shiraki
(Source: Su NY; University of Florida, Publication No. EENY121)
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because of the difculty to maintain live colonies in the laboratory. The problem is to
provide a nesting environment that is not overly articial (possibly resulting in
behavioral artifacts), while having a visual on all individuals of the colony, at all
times. Researchers and termite control practitioners can only observe termite activity
through one or more “windows.” This limited view of the diffuse network of tunnels
and feeding sites occupied by a termite population is at the heart of the problem of
determining the population parameters. An ethogram is the description of an ani-
mal’s behavior repertoire that forms the basis of ethological studies. Ethograms have
been constructed for many animals, including insects (Dinesh and Venkatesha 2013),
particularly social insects such as bees (Seeley 1982), ants (Jayasuriya and Traniello
1985), and termites (Rosengaus and Traniello 1991). In social insects, ethograms are
particularly relevant in understanding division of labor among individuals in a col-
ony, where caste polyethism and age polyethism can result in optimized growth and
tness for the colony (Seeley 1982). The thorough descriptions of ethograms in ter-
mites are rare, owing to their cryptic lifestyle in a closed nest system. There is, there-
fore, an inherent difculty in observing the range of behaviors of an entire colony
with all castes. Behavioral observations of termites have typically focused on a few
fragmentary behaviors, such as feeding (Indrayani etal. 2007) and foraging (Li and
Su 2008). According to the origin, relatedness, and number of active reproductives,
termite colonies are classied as simple families, extended families, and mixed fami-
lies (reviewed in Vargo and Husseneder 2011). Simple families are colonies headed
by a regular monogamous pair, the royal couple, whereas extended and mixed
Fig. 1.2c Life cycle of the Cryptotermes brevis (Walker) (Source: Scheffrahn RH; University of
Florida, Publication No. EENY079)
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families present multiple reproductives. In both extended and mixed families, the
multiple females may be accompanied by multiple males, because, in termites, poly-
andry is often associated with polygyny (Roisin and Pasteels 1985). Mixed families
can also result from colony fusion (DeHeer and Vargo 2008).
Many species build nests, but it is common to nd more than one type of termite
living in the same active or abandoned nest (Bandeira 1983). Members of
Inquilinitermes are obligatory inhabitants found in the nests of Constrictotermes, a
termite that builds arboreal nests (Melo and Bandeira 2004). Chiu et al. (2015)
speculated that the various food sources and their distributions are likely the main
selection force for the gallery structures of soil-feeding termites. Studies of the tun-
neling behavior of marked C. formosanus determined that a small number of spe-
cic individuals performed most of the work while most of the individuals remained
inactive (Cornelius 2012). Study by Cornelius and Gallatin (2015) provided a
detailed analysis of the tunneling behavior of workers of the subterranean termite,
C. formosanus. Chemical communication certainly represents the dominant mean
of information exchange in social insects (Richard and Hunt 2013). Chemical medi-
ators of intraspecic communication, the pheromones, are secreted from exocrine
glands (Billen 2011) and are perceived by specialized chemoreceptors, located pre-
dominantly on the antennae (Wyatt 2003).
Termites’ evolutionary success has been linked to their defense mechanisms.
They have developed a multitude of active and passive defensive traits. The active
defenses comprise morphological (Deligne etal. 1981), chemical (Prestwich 1984;
Sobotnik etal. 2010), and behavioral (Sobotnik etal. 2012) adaptations present pre-
dominantly in soldiers, while passive defense include cryptic way of life and nest
fortication preventing attacks from nonspecialist predators (Noirot and Darlington
2000). Soldiers are recruited for defense in a disturbed area. It was noticed that the
disturbed soldiers did not escape in proportion to the workers; rather, there was a
modest increase in soldier number (Gautam and Henderson 2012).
1.3 Systematics, Distribution, andDiversity
Termites are traditionally ranked as an insect order (Isoptera), representing a sub-
group within Blattodea, with Cryptocercus being their sister taxon (Lo etal. 2000;
Inward etal. 2007b; Djernaes etal. 2015). Termite systematics is traditionally based
on the external morphology of soldiers and alate imagoes1 (e.g., Holmgren 1912;
Emerson 1925; Snyder 1926). However, species with overlapping geographic
ranges are notoriously difcult to distinguish morphologically. Fortunately, the
worker morphology also allows species identication, especially in soil-feeding
taxa, whose digestive tract is highly modied and morphologically distinct among
species (Noirot 2001). Despite the wide distribution of Coptotermes in the world,
and the large body of associated scientic literature for population management, the
1 Imago: the last stage attained by an insect at the issue of metamorphosis.
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taxonomy of Coptotermes remains unsettled, and many species names may be syn-
onyms of others (Chouvenc etal. 2015). Krishna etal. (2013b) listed 110 species
names within Coptotermes that conformed to the rules of the International Code of
Zoological Nomenclature (ICZN). Among them, 69 were regarded as valid in the
taxonomic literature, and 42 were listed as subjective synonyms. Moreover, several
species currently included in genus Ruptitermes Mathews 1977 were initially clas-
sied in the genera Anoplotermes Fr. Muller 1873 and Speculitermes Wasmann
1902 (Acioli and Constantino 2015). Termite experts have also made attempts to
solve taxonomic cold cases (http://entomologytoday.org/2015/12/22/termite-
experts-attempt-to-solve-taxonomic-cold-cases). The American Museum of Natural
History (AMNH) collection of termites is, without question, the largest and most
comprehensive in the world for these insects and fully global in scope. The collec-
tion consists of over a million specimens belonging to about 80% of the world’s
species (excluding the plethora of species recently described from China). A refer-
ence library on termites contains the most comprehensive archive of original publi-
cations on the systematics of Isoptera (http://www.amnh.org/our-research/
invertebrate-zoology/collections/isoptera).
Although the presence of the soldier caste is a synapomorphy2 of all termites
(Roisin 2000), the soldier to worker ratio decreases in soil-feeding termites com-
pared with their wood-feeding relatives (Haverty 1977). Soldierless termites
(Termitidae, Apicotermitinae) constitute about one third of the termite diversity in
African and South American rainforests (Eggleton 2000). Because they lack the
soldier caste, species identication must be based on alate imagoes, when collected,
and on workers and requires the dissection and close examination of their digestive
tube (Noirot 2001). An excellent review on poorly known and ecologically domi-
nant soldierless Apicotermitinae is presented by Bourguignon etal. (2016).
Although human transport of termite-infested material is the primary method of
expansion to new regions, natural dispersal occurs slowly through the annual nup-
tial ights of alates. Alates of C. formosanus generally swarm from April through
June to give rise to new colonies (Henderson 1996). Average recorded alate ight
was 621m, and the longest ight was 1.3km distant from the parent colony (Mullins
etal. 2015). Therefore, alate dispersal plays an important role in the spread of C.
formosanus in areas where it has become established and in the reinvasion of areas
where colonies have been eliminated through treatments. C. formosanus is endemic
to China and Taiwan and has spread to many temperate and subtropical regions
(Evans etal. 2013). It is now found throughout the southeastern United States and
is responsible for more than $1billion of structural damage each year in this country
alone (Corn and Johnson 2013). Coptotermes gestroi (Wasmann) is native to
Southeast Asia and has spread in many tropical regions, being potentially the most
ubiquitous and destructive subterranean termite pest in the world (Evans etal.
2013). Both species have distinct ecological requirements (Grace 2014), but there
are now established populations in many non-native areas due to human activity
2 Synapomorphy, presence of a shared derived character that characterizes a clade from other
lineages.
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(Hochmair and Scheffrahn 2010). C. formosanus probably became established in
Florida in the early 1970s, remained undetected until the rst reported record
(Koehler 1980) in Hallandale (Broward County), and has since been found in most
urban localities throughout Florida (Scheffrahn 2013).
Reticulitermes Holmgren, 1913, is a Holarctic genus of subterranean termites
(Isoptera: Rhinotermitidae) that is widespread and abundant in temperate regions
where their biomass can approach that of many termite taxa living in tropical regions
(Bignell and Eggleton 2000). Human commercial activities unwittingly transport
termites to their nonendemic areas. This is evidenced by the appearance of
Reticulitermes infestations in Hallein (Austria), Hamburg (Germany), Devon
(England), and Toronto (Canada) (Gay 1969; Su and Tamashiro 1987; Jenkins etal.
2001). Thus subterranean termites will continue to be a worldwide problem for
urban and suburban property owners (Forschler and Jenkins 2000). There is also
mounting evidence that warming environments resulting from climate change can
be an important factor for altering the species distribution (Chunco 2014).
Termite faunal structure is determined by climatic and vegetational characteris-
tics. Termite species richnesses, abundances, diversity, and trophic structures, how-
ever, differ between the two kinds of ecosystems. Greater species richness (Eggleton
etal. 1997; Vasconcellos etal. 2010) and density (Bignell and Eggleton 2000) have
been observed in humid forests than in arid or semiarid environments. Differences
related to feeding groups are also observed, with humus-consuming species being
more abundant and vulnerable to environmental alterations in humid forests
(Eggleton etal. 1997; Vasconcellos 2010). Xylophagic species constitute the most
abundant and vulnerable group in dry tropical forests (Vasconcellos etal. 2010).
The structure of the termite fauna can vary considerably between areas in the same
ecosystem, after exposure to different degrees of anthropogenic alterations
(Vasconcellos etal. 2010). The modication of natural areas to form agroecosys-
tems, for example, can result in signicant loss of species richness, abundance, and
diversity (Bandeira etal. 2003).
1.4 Invasive Termites
The invasion of a new habitat by an introduced species may depend on a number of
factors such as the suitability of the abiotic environment (Blackburn and Duncan
2001), the ability of the species to adapt to the novel environment (Sax and Brown
2000), and the interaction between the invader and the recipient community (Holway
1998). Coptotermes is a termite genus that is ecologically successful. Two species,
the Formosan subterranean termite, C. formosanus, and the Asian subterranean ter-
mite, C. gestroi, are particularly invasive (Chouvenc etal. 2015). They have spread
far beyond their native range with the help of human maritime activities (Scheffrahn
and Crowe 2011; Rust and Su 2012). These two species contribute in large part to
the annual $40billion cost associated with termite damage and control around the
world (Rust and Su 2012). Whereas C. formosanus has a warm temperate/
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subtropical distribution, C. gestroi has a tropical distribution (Cao and Su 2016). In
the New World, C. formosanus has invaded most of the southeastern United States,
whereas C. gestroi has invaded areas of Brazil, most of the Caribbean, and, more
recently, parts of south Florida (Su etal. 1997; Scheffrahn etal. 2015). One of the
reasons for the success of termite species comes from their ability to adapt to dis-
turbed environments and display a high behavioral plasticity at the colony level,
with efcient task division (Du etal. 2016). Chouvenc etal. (2016) reported that the
risk for structures in metropolitan southeastern Florida with known Coptotermes
infestations increased from 0.49% to 7.3% (from year 2000 to 2015), with some
species distributional overlap. In addition, several localities that had Coptotermes
records before 2000 have registered an increased density of termite infestation and
swarming activity. It is expected that the distribution and structural infestations by
Coptotermes will continue to increase in the years to come, with an estimated 50%
of all structures in southeastern Florida at risk by 2040.
1.5 Termite-Gut Microbiota
Gut-associated microbes of insects are postulated to provide a variety of nutri-
tional functions. The diet of termites is diverse, with cellulose as the main food
resource exploited (Moore 1969; Lima and Costa-Leonardo 2007). However, ter-
mites are decient in enzymes that decompose cellulose and lignin, which provide
them with extra carbohydrates (Williams 1965). For this reason, they require the
aid of symbiotic microorganisms in their feeding channel to digest these com-
pounds. Termite- gut microbiota is very diverse and comprises many phylogenetic
lineages that have been extensively documented in recent decades (Ohkuma and
Brune 2011). The gut of these insects is a specialized habitat for bacteria, archaea,
and protists which make them highly efcient decomposers (Eggleton 2011).
Based on feeding ecology, termites can be grouped as the higher termites and the
lower termites. Lower termites (all families except Termitidae) harbor in their gut
a dense and diverse population of prokaryotes and agellated protists (Ohkuma
2003). Protozoan symbionts residing in lower termites are responsible for ligno-
cellulose digestion in this group. Digestion of cellulose and hemicellulose is
attributed to a consortium of termite, bacteria, and protist-derived cellulases that
ultimately liberate carbon in plant tissues (Scharf etal. 2011). Higher termites
(Termitidae) lack agellates and harbor only prokaryotes in their highly struc-
tured guts. However, recently, a low- abundant ciliate has been detected in the guts
of higher termite species (Rahman et al. 2015). Lower termites predominantly
feed on dead wood (with few exceptions). The diversication of feeding habits is
high in Termitidae. The symbionts are not transmitted vertically (from mother to
offspring) but become established by a gradual process allowing the offspring to
have access to the bulk of the microbiota prior to the emergence of workers and,
therefore, presumably through social exchanges with nursing workers
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(Diouf etal. 2015). The acquisition of genetic data in termites and their gut micro-
bial community has been of recent interest to the scientic community. This is
mostly due to the development and accessibility of new sequencing technologies
such as 454 pyrosequencing and Illumina sequencing (Scharf 2015).
1.6 Feeding Groups
Termites have also been classied into several functional feeding groups: soil feed-
ers, soil/wood interface feeders, wood feeders, litter foragers, epiphyte feeder, grass
feeders, and some other minor feeding groups (Collins 1984). Although several
classications into feeding groups have been proposed (Inward etal. 2007a), the
fundamental differential trait lies in the distinction between wood- and soil-feeding
groups (Bourguignon etal. 2009). Wood-feeding termites feed on sound dead wood
and true soil-feeding termites feed on soil organic matter in mineral soil, with no
visible plant remains. Soil-feeding termites were found in the surface horizon of soil
(Inoue etal. 2001), in the intermediate organic matter between wood and topsoil
(Souza and Brown 1994), and in the mounds of other termite species (Eggleton and
Bignell 1997). These termites increase the polysaccharide content of soil (Garnier-
Sillam and Harry 1995) and facilitate its humication process (Brauman 2000).
Most soil-feeding termites build subterranean and diffuse gallery systems that are
difcult to observe. The high diversity of soil-feeding termites indicates soil feeders
as a successful feeding guild of termites (Brauman etal. 2000).
1.7 Biotic andAbiotic Factors
Biotic and abiotic factors play a role in the successful establishment of invasive spe-
cies. Among the abiotic factors, moisture and temperature play a vital role in deter-
mining which areas are the most suitable for establishment. Given the importance of
these factors, a uctuation in either moisture or temperature will impact the overall
termite consumption and survival. Humidity directly affects temperature and vegeta-
tion structure, which strongly inuence the termite assemblage structure. The workers
and soldiers, which comprise a large proportion of the colony’s population, have a soft
integument that makes them extremely vulnerable to desiccation (Moore 1969).
Surrounding moisture is of utmost importance for the survival of subterranean ter-
mites, which are highly susceptible to desiccation, making moisture a critical factor
for survival. Many studies have examined the inuence of moisture on the tunneling
and feeding behavior of subterranean termites (McManamy etal. 2008; Gautam and
Henderson 2011) which have a relatively soft cuticle that readily desiccates (Moore
1969). As a survival strategy, subterranean termites always associate with moist and
humid environments. Moisture can be obtained from many sources, including meta-
bolic breakdown of sugars (food source) and wet food materials (Pearce 1997).
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Barriers of dry soil affect the ability of termites. Cornelius and Osbrink (2011)
observed high signicant effect on the ability of termites to colonize food located in
dry sand. They reported that only one feeding station located in dry sand was colo-
nized by termites, compared with 11 feeding stations located in moist sand. Delaplane
and LaFage (1989) reported that Coptotermes formosanus Shiraki preferred wet wood
blocks over dry blocks. Behr etal. (1972) also showed a positive correlation between
wood moisture level and the feeding by Reticulitermes avipes (Kollar).
Temperature is another important factor that determines the geographic distribu-
tions, feeding, and survival of species. It was believed that soil temperature has
more effect on activity of subterranean termites than air temperature (Ettershank
etal. 1980). Sen-Sarma and Mishra (1968) studied the seasonal activity variation of
Microcerotermes beesoni Snyder in North India and indicated that soil temperatures
determined termite activity levels in different seasons. Evans and Gleeson (2001)
documented similar observations from the study of another subterranean termite
species, Coptotermes lacteus (Froggatt), in Australia. Haverty etal. (1974) noted
that foraging intensity of Heterotermes aureus (Snyder) increased in spring and fall
but decreased during the winter months in desert grassland. Reticulitermes sp. pre-
ferred signicantly lower temperatures than Coptotermes sp. (Cao and Su 2016).
The highest survival (Fei and Henderson 2002) and the highest feeding rate
(Nakayama etal. 2004) for C. formosanus were reported at 30°C.
Fire strongly affects habitat resources used by termites, such as plant biomass and
dead wood (Haslem etal. 2011), and therefore may indirectly modify their commu-
nities. However, termites appear to be resistant to the effects of re at multiple spa-
tial scales (Avitabile etal. 2015). Termites are common in re-prone landscapes,
including savannas worldwide (Davies etal. 2010) and arid and semiarid woodlands
(Abensperg-Traun etal. 1996). In general, altitude and species richness correlate
negatively for several organisms (McCain 2009). However, Diehl etal. (2015) found
no signicant correlation between termite species richness and altitude.
1.8 Economic Importance
The economic importance of termites is twofold, extremely benecial and extremely
injurious to man. These small creatures are a part of the natural ecosystem and con-
tribute signicantly to most of the world ecosystems. The signicance of termites
for ecosystem functioning is widely acknowledged and receives considerable atten-
tion from the scientic community, with much effort applied to disentangling their
specic contributions to ecosystem functioning (Davies etal. 2014b). Termites are
important in both dry and humid tropical forests, where they are consumers of the
plant necromass, helping in the processes of nutrient cycling and soil formation
(Lee and Wood 1971; Vasconcellos and Moura 2010). A great role in the cycles of
biogenic elements in tropical forest ecosystems belongs to termites that consume up
to 50% of the leaf litter (Brauman 2000). They are often referred to as ecosystem
engineers because they shape the environment through their action. Bonachela etal.
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(2015) reported that in many arid ecosystems, termite nests impart substrate hetero-
geneity by altering soil properties, thereby enhancing plant growth. Furthermore,
they noticed that mound-eld landscapes are more robust to aridity, suggesting that
termites may help stabilize ecosystems under global change. Termite mounds shape
many environmental properties, as their soils differ from surrounding “matrix” soils
in physical and chemical composition, which enhance vegetation growth (Sileshi
etal. 2010), creating “islands of fertility” (Sileshi etal. 2010; Davies etal. 2014a).
The increased soil fertility and moisture found near termite mounds can have pro-
nounced effects on vegetation communities and their productivity (Sileshi et al.
2010). Previous studies have found that woody vegetation growing on termite
mounds increased density (Moe etal. 2009), tree height (Levick etal. 2010), species
richness (Traore etal. 2008), functional diversity (Joseph etal. 2014), and reproduc-
tive output (Brody etal. 2010).
Termites have long been studied because of their uncommon diet and complex
hindgut microbiota. Researchers have discovered that enzymes found in a termite’s
digestive system could aid in biofuel production from woody biomass (see Chap. 5
for more details). The lignocellulolytic system in wood-feeding termites has some
unique system advantages and can potentially serve as a model system to improve
our current biomass bioconversion technology for fuels and chemicals (BenGuerrero
etal. 2015). The termitaria are formed from materials burrowed from deep-seated
environments upward by termites and are residual in character. The use of termite
mound samples is an appropriate media in the search for concealed mineralization
in complex regolith environments (Arhin etal. 2015). Affam and Arhin (2006) rec-
ognized termite mounds as a good geochemical sample media for gold exploration,
and its validation has been conrmed by Arhin and Nude (2010) in northern Ghana.
Termite species, however, gain pest status when they damage building materials
or agronomic and forestry commodities. As the principal food of some of the termite
castes is cellulose, they cause economic losses by directly injuring and destroying
both living and dead vegetation, buildings, bridges, dams, etc. Many subterranean
termite species are considered “urban pests” due to their tendency to attack man-
made structures (Rust and Su 2012), and some are now invasive throughout the
world, increasingly causing structural damage (Evans etal. 2013). Subterranean ter-
mites, particularly members of the genera Coptotermes and Reticulitermes, represent
the most widespread and economically important structural insect pests in the urban
environment (Gay 1969; Su and Scheffrahn 1990). Twenty-three species in the genus
Coptotermes are among the most signicant termite pests worldwide for man-made
structures. C. formosanus and C. gestroi are of particular economic importance (Rust
and Su 2012) due to their ecological success and invasive ability (Evans etal. 2013).
Termites cause tree damage in public areas, thus threatening people safety. Infested
living trees can ultimately lead to their felling and death. Coptotermes formosanus
attacks structural wood as well as living trees (Henderson 2001).
Once a colony of C. formosanus is established in an area, it soon invades nearby
areas while searching for food and gradually spreads to new locations (Fig.1.3).
This termite lives a cryptic lifestyle where workers and soldiers forage through tun-
nels and galleries originating from their nests. Unlike subterranean Reticulitermes
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13
sp., C. formosanus has a large colony size and aggressive feeding behavior
(Tamashiro et al. 1980). In Japan and the United States, C. formosanus and in
Australia C. acinaciformis (Froggatt), as well as Mastotermes darwiniensis Froggatt,
are economically important species (Alexander etal. 2014). However, subterranean
termite R. avipes (Kollar) is the most common and widely distributed termite pest
species in the United States (Scheffrahn etal. 1988; Wang etal. 2009). Termites are
selective in which types of wood they feed on. Wood species vary in chemical
makeup and structure providing a number of possible cues that termites could use in
food selection. Natural compounds such as sugars, amino acids, urea (Castillo etal.
2013), and phosphates (Botch etal. 2010) were previously found to increase termite
feeding. It has also been suggested that wood ber density also plays an important
role in food selection.
Economic losses associated with termite damage in the United States and Japan
are around 1000 and 800million US$ a year, respectively (Verma etal. 2009), and
Japan may be the third largest user of pesticides for structural pest control in the
world. In Europe, the losses caused by termites are estimated at 313million US$ per
year (Eggleton 2000). Economic losses due to termite in India have been estimated
around 35.12million US$ (Joshi etal. 2005). However, in Malaysia 8–10million
US$ is spent toward termite treatment every year (Lee 2002). The global economic
impact of termite pests is estimated to be at least $40billion (Rust and Su 2012).
1.9 Prevention andControl
For decades, soil treatment with liquid termiticides has been the dominant method
used in subterranean termite control programs. The traditional method is by injec-
tion of liquid termiticide to the soil to establish a toxic or repellent chemical barrier
Fig. 1.3 A single colony of the Formosan subterranean termite Coptotermes formosanus Shiraki
may contain several million individuals that forage up to 100m in soil (Source: Su NY; University
of Florida, Publication No. EENY121)
1 Termites: AnOverview
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14
against termites. Barrier treatments rst were developed in the 1940s and have
changed very little since then (Lewis 1997). These treatments are labor-intensive
and require a relatively large amount of insecticide to achieve the required concen-
tration levels in the soil. Liquid termiticides are either neurotoxins or inhibitors of
mitochondrial respiration. There are six main insecticide classes, i.e., organophos-
phate, carbamate, pyrethroid, neonicotinoid, phenylpyrazole, and avermectin, of
termiticides used currently in the eld (Chen etal. 2015).
The use of proper application method of termiticides is important to reduce their
negative impact to the environment. Though treatments result in varying degrees of
success depending on the skills of the applicator, type and dosage of chemical used,
and degree of infestation, it is extremely difcult to obtain a continuous and uniform
distribution of insecticide at the correct soil concentration, around infested structure
(Forschler and Lewis 1997).
Termite control exploits their eusociality to deliver the insecticide. When ter-
mites forage in termiticide-treated areas, they acquire the active ingredient and inad-
vertently share it with unexposed nestmates, a process known as horizontal transfer.
Subsequently, horizontal transfer results in secondary mortality in situations where
a lethal dose of the active ingredient is transferred from the exposed donor termites
to the unexposed recipient termites. Transfer of pesticides among termites (mainly
through grooming) has been reported for various products, e.g., imidacloprid,
indoxacarb, ivermectin, and chlorpyrifos (Valles and Woodson 2002; Shelton and
Grace 2003; Hu etal. 2005).
It is established that, unlike repellent soil termiticides, nonrepellent, delayed
action termiticides have impacts beyond the treated area. Nonrepellent termiticides
are slow-acting insecticides that are not detected by termites when they forage
through treated soil. One important advantage of a nonrepellent termiticide is the
potential greater “coverage” as termites do not detect the chemical presence and do
not die too quickly after walking across the treatment (Thorne and Breisch 2001).
This opens the door for possible alternative treatment methods to incorporate into
integrated pest management strategies that reduce the amount of chemicals applied.
Presently, nonrepellent and relatively slow-acting liquid termiticides represented by
imidacloprid (Premise®), pronil (Termidor®), chlorfenapyr (Phantom®), indoxa-
carb (Aperion™), and chlorantraniliprole (Altriset™) are soil termiticides widely
used for the prevention and treatment of structural infestations of subterranean ter-
mites (Potter and Hillery 2002; Remmen and Su 2005; Gautam etal. 2014).
Together with prevention and control strategies, early detection of termite infes-
tations would enable the determination of infestation extent and the delineation of
areas for future treatment procedures (Su and Scheffrahn 2000). Visual inspection is
the normal procedure for dry-wood termite detection, although it is not 100% effec-
tive, so it should be accomplished with other sophisticated methods, such as mois-
ture meters, electronic odor devices, acoustic emission detectors, or infrared heat
detectors, for enhancing the reliability of termite detection (Evans 2002; Oliver-
Villanueva and Abian-Perez 2012). Throughout the world, chemical termiticides are
going to be replaced by baits, microwave, and sensor technology. Termite detection
radar, moisture meter, and remote thermal sensor with laser are available throughout
M.A. Khan and W. Ahmad
mdaslam30@gmail.com
15
the world. These can detect termites underground and use fewer chemicals than
traditional methods (Manzoor 2013). Therefore, nondestructive detection methods
together with traditional visual inspections are advisable, since the integrity of
wooden elements is maintained.
Popular control practices involve the use of nonrepellent termiticides and baiting
systems (Henderson 2001). Subterranean termite colonies have extensive under-
ground gallery systems, and it is difcult to eliminate entire colonies using soil
insecticides. Baiting programs have been used successfully to eliminate subterra-
nean termite colonies (Su etal. 1995; Eger etal. 2012). The goal of a baiting system
is to eliminate the entire termite colony from an area with the least possible cost and
harm to nontarget organisms in the environment (Su and Scheffrahn 1998). Baiting
systems depend on the exploitation of foraging behavior of subterranean termites,
where a subset of individuals from a colony feed on the cellulosic food material
impregnated with slow-acting toxicants and introduce the toxicants to the colony
(Su and Scheffrahn 1998). The success of baiting system is more variable than that
of the liquid soil termiticides (Lewis 1997). Environmental factors such as tempera-
ture, humidity, soil type, and soil moisture affect termite activity at bait stations
(Messenger and Su 2005; Ruan etal. 2015). Seasonal variations in caste distribution
of foraging populations could also inuence feeding and foraging behavior of ter-
mites on baits. Although baiting is the most environmental friendly way of control-
ling termites, about two thirds of the treatments by pest control companies rely on
the use of liquid insecticides in soil (Curl 2004). Although insecticide resistance is
extremely rare among social insects, it is nonetheless important to search for new
alternatives of conventional insecticides used in termite control.
Biological alternatives for termite control include botanicals (essential oil, seed,
bark, leaf, fruit, root, wood, resin), as well as fungal, bacterial, and nematode
approaches (Verma etal. 2009). The active component from biomass can be extracted
to prepare efcacious and potent biocidal formulations. Phytophagous insects use
plant volatiles to recognize their host plants. Therefore, the use of essential oils as a
nonhost volatile emission to repel insect pests is a viable alternative for control
(Mauchline etal. 2005). Numerous studies have documented the natural resistance of
certain wood species to termite attack. Cornelius and Osbrink (2015) reported that
toxic chemical components of teak hold the most promise as wood preservatives.
Termites are frequently preyed upon by ants in tropical forests, and most termite
species are likely to be affected by ant predators (Goncalves etal. 2005). They
exhibit several adaptations to avoid predation, including chemical defense, mandi-
ble snapping, and ghting with large, smashing mandibles (Prestwich 1984;
Legendre et al. 2008). Some species of ant, including those from the genera
Centromyrmex (Bolton and Fisher 2008), Megaponera (Dejean et al. 1999),
Anochetus (Schatz et al. 1999), Tetramorium (Longhurst et al. 1979), and
Paltothyreus (Dejean etal. 1993), specialize on particular termite taxa, while spe-
cies from a wide range of genera are known to predate termites opportunistically, to
a greater or lesser extent (Dejean et al. 1999). Other species, such as Dorylus
(Anomma) driver ants, only feed on alates during swarming (Schoning and Moffett
2007). Furthermore, there is substantial (correlational) evidence that the nest den-
1 Termites: AnOverview
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16
sity of termites is limited by the abundance of both dominant (Pequeno and Pantoja
2012) and non-dominant ant species (Ellwood etal. 2002). Several spider species
have specialized to feed on prey that is highly aggregated, including termites and
ants (Haddada etal. 2016). Wesolowska and Haddad (2002) reported Heliophanus
(Heliocapensis) termitophagus n. sp., a jumping spider in or on the termitaria of
Trinervitermes trinervoides (Sjostedt) that fed mostly on workers of T. trinervoides.
Petrakova et al. (2015) reported that the spider Ammoxenus amphalodes is a
monophagous prey specialist, specically adapted to feed on harvester termites,
Hodotermes mossambicus (Hagen). The spiders attacked the lateral side of the tho-
rax of termites and immobilized them within 1min. The paralysis efciency was
independent of the predator/prey size ratio. However, its role as a biocontrol agent
against termites is limited, due to an insufcient numerical response.
Entomophagy, the practice of using insects as a part of the human diet, has played
an important role in the history of human nutrition in Africa, Asia, and Latin America
(Srivastava etal. 2009). However the use of insects by human in medicine is known
as entomotherapy. People from different parts of the world use termites as food (for
humans and livestock) and as a source material for popular medicine (Figueiredo
et al. 2015). Kinyuru et al. (2013) reported that different edible termites from
Western Kenya contain about 45% fat and 35% dry matter. Research continues to
focus on suitable termite control measures that are both effective and environmen-
tally benign. Ultimately there is a dire need to develop strategies for sustainable
termite management locally that would save money and protect the environment.
1.10 Conclusion
The cryptic nature and social organization of termites represent a primary reason
why termite infestations can be difcult to study and control. Once a subterranean
colony is established in an area, it soon invades nearby areas while searching for
food and gradually spreads to new locations. Labor-intensive soil treatment with
liquid termiticides has been the dominant method used in subterranean termite con-
trol programs. There is a dire need to develop strategies for sustainable termite
management, focusing on goals of least possible cost and harm to nontarget organ-
isms, in the environment.
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1 Termites: AnOverview
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... The termites (Insecta: Isoptera) comprise a group of eusocial insects composed of individuals that form different castes that carry out various ecological roles within their colonies (e.g., defense, feeding, reproduction, etc.) (Roisin, 2000;Eggleton, 2011;Khan and Ahmad, 2018). Currently, there are around 2600 described species worldwide, located mainly in tropical to subtropical regions, although, they have also reached temperate regions of latitudes greater than 45°in both hemispheres (Eggleton, 2000;Kambhampati and Eggleton, 2000;Jones and Eggleton, 2011). ...
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Borings filled with coprolites in silicified conifer woods from the Lower Cretaceous (Albian) of the Kachaike Formation, Santa Cruz Province, Argentina are described for the first time. Coprolites are approximately hexagonal in cross-section, like those produced by extant Kalotermitidae, and occur inside borings in the secondary xylem. The presence of this family in the mid-Cretaceous ecosystems of Patagonia indicates arid climatic conditions consistent with previous geological, palynological and paleoxylological studies for this time interval in the Austral Basin. This also adds to other worldwide records of termites from the Lower Cretaceous, supporting their ecological importance as consumers and recyclers of lignified organic matter in dry forests from this period on to today.
... Additionally, they serve as faunal refuges because of the forest-like habitats they create (Griffith, 2000). Termites are critical components of tropical soil ecosystems constituting over 90% of the insect biomass in tropical forest soils (Eggleton, 2000;Jouquet et al., 2011;Khan & Ahmad, 2018;Nonoh, 2013). They play major roles in processes such as decomposition, and nutrient and carbon cycling Evans et al., 2011;Griffiths et al., 2019). ...
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Cocoa is an important crop for Ghana's economy, contributing 25% of Gross Domestic Product (GDP). The crop, however, is mainly cultivated on forest‐derived soils and is a major cause of land‐use change. Termites are an important biological component of tropical ecosystems providing numerous ecosystem services. Previous studies have indicated that termites are sensitive to forest disturbance and decrease in richness and abundance across land‐use intensification gradients, with consequences for the essential services that they provide. Native shade trees are often used to improve cocoa cultivation and may reduce the detrimental effects of land‐use change on some aspects of biodiversity. The aim of this study was therefore to explore how termites respond to land‐use change along a shade‐tree gradient in Kakum National Park and surrounding cocoa farms in Ghana (from forest at 80% tree cover to cocoa with no shade cover, to the extreme of cultivated arable crop land). It was predicted that termite richness and abundance would decrease with decreasing shade cover, and with increasing distance from the forest edge. Thirty‐four species from 29 genera were sampled, with Ancistrotermes crucifer being found in all the locations (47% of all encounters). Species richness and abundance differed marginally across the land‐use gradient, as well as the distance from the forest edge; however, species richness did not show any significance with distance. All the same, termite communities were robust to the disturbance. Our findings suggest that though site influenced species richness and abundance, cocoa trees can play a crucial role in maintaining biodiversity and environmental quality in an agricultural landscape by providing a habitat for forest species that are not found in pastures or farm fields. However, we caution that the relatively low forest baseline of existing forest diversity may inflate the value of cocoa land, with those forests no longer representing undisturbed natural habitats: this highlights that shifting baselines may need to be accounted for when interpreting findings in the Anthropocene. Termites are an important biological component of tropical ecosystems providing numerous ecosystem services. Native shade trees are often used to improve cocoa cultivation and may reduce the detrimental effects of land‐use change on some aspects of biodiversity. Our findings suggest that though site influenced species richness and abundance, cocoa trees can play a crucial role in maintaining biodiversity and environmental quality in an agricultural landscape by providing a habitat for forest species that are not found in pastures or farm fields
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Reticulitermes flavipes, one of the most harmful subterranean termite pests, is reported for the first time from Tenerife (Canary Islands, Spain). Cytochrome oxidase II was sequenced from five specimens in order to confirm the identification. To date, this invasive species has been detected in a limited area in the northeast of the island, affecting buildings, crops and native plant species. Another colony with the identical haplotype found in the southwest, 60 km away from the main population, indicates that this invasive insect may be more widespread over the island.
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Results of previous field excavation of nest structure of Coptotermes formosanus Shiraki (Isoptera: Rhinotermitidae), the Formosan subterranean termite, indicated that the total length of the gallery system ranged up to 580 m, and its total space may occupy >34,800 cm³. Because Formosan subterranean termite does not build mounds, it has been speculated that it creates tunnel space by compacting soil. The objective of this study is to test a “modified soil-compaction hypothesis” that Formosan subterranean termite removed and compacted soil to increase space during tunnel excavation. Contrary to the hypothesis, the deposited sand was less dense than the unexcavated sand in all replications; thus, we rejected the soil-compaction hypothesis. Instead, we offered wood-consumption hypothesis that termites gain the tunnel space as a result of consuming wood. When termites were placed with wood pieces for 30 d, the decreased volume of consumed wood was significantly higher than the increased volume of carton material. The net increased space is ≈;50% of volume of consumed wood. We speculate that the space created by wood consumption could be transformed into tunnel space during soil displacement.
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This study reports the spread of 2 major invasive subterranean termite species (Isoptera: Rhinotermitidae) in metropolitan southeastern Florida: Coptotermes formosanus Shiraki and C. gestroi (Wasmann). Termite records from 1990 to 2015 were analyzed to determine the expansion of their distribution. Our results suggest that the ranges of their distribution have increased exponentially during this time frame. This observation raises concerns about potential structural damage in this urbanized area, which includes 6 million residents. The risk to structures located in an area with known Coptotermes infestation increased from 0.49% in 2000 to 7.3% in 2015, with some species distributional overlap. In addition, several localities that had Coptotermes records before 2000 have registered an increased density of termite infestation and swarming activity. We argue that the subterranean termite problem in metropolitan southeastern Florida is still in its early phase of invasion, and we predict that the distribution and structural infestations by Coptotermes will continue to increase in the years to come, with an estimated 50% of all structures in southeastern Florida at risk by 2040.
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We are entering one of the most exciting periods in the study of chemical communication since the first pheromones were identified some 40 years ago. This rapid progress is reflected in this book, the first to cover the whole animal kingdom at this level for 25 years. The importance of chemical communication is illustrated with examples from a diverse range of animals including humans, marine copepods, Drosophila, Caenorhabditis elegans, moths, snakes, goldfish, elephants and mice. It is designed to be advanced, but at the same time accessible to readers whatever their scientific background. For students of ecology, evolution and behaviour, this book gives an introduction to the rapid progress in our understanding of olfaction at the molecular and neurological level. In addition, it offers chemists, molecular and neurobiologists an insight into the ecological, evolutionary and behavioural context of olfactory communication.