Exotic species invasion is one among the many of the world’s serious environmental threats (Buczkowski and Bertelsmeier, 2017). Although growing human population and their injudicious actions are the critical reasons for climate change and urbanization, invasions by exotic species can be traced back to mobility of humans and movement of products to meet human needs. Huge numbers of exotic, pestiferous insects are moving from continent to continent as a result of globalization and trade, resulting in substantial economic loss (Sharma et al., 2018). In the last decade, various invasive insects are known from India and have been found to be establishing in ‘new’ habitats (Kalleshwaraswamy et al., 2015; Sharanabasappa et al., 2018; Sundararaj et al., 2020) and termites are no exception (Evans et al., 2013). Many species of termites are serious nuisance organisms, inflicting damage to the wood used in both human-made structures and agricultural and forest environments (Kalleshwaraswamy et al., 2018). They occur more abundantly in the tropical and subtropical regions of the world. We now know that the Isoptera have evolved from the subsocial Cryptocercidae (Blattodea: Blattoidea) (Inward et al., 2007; Evangelista, 2019). Hence, the termites are now classified under Isoptera, Epifamily Termitoidae within Blattodea. However, the family status of all extant termites is retained to avoid confusion arising out of this modified classification. About 3000 species of Isoptera were described from nine families in 2013 (Krishna et al., 2013) and from then onwards, a few new species have been added from different parts of the world. Among them, 1160 species are from the Oriental region. Two hundred and ninety-five species placed under 52 genera in six families are known from India. Among the 295, 188 are endemic to India (Rajamohana et al., 2019). Out of c. 3000 species known around the globe, 366 affect human dwellings in different parts of the world. Yet, we need to recognize that these taxa are one among the many critical invertebrate decomposers in arid and semi-arid environments, and they contribute to improving the physical and chemical properties of soil by their activities and especially by the construction of structures such as mounds, galleries, sheetings with varying physical and chemical qualities (Jouquet et al., 2018). On an average, the termites increase water infiltration above the natural rate by a factor of 1 to 4, depending on their activity, soil type, and rainfall intensity (Kaiser et al., 2017). |
Diverged from cockroaches (Cryptocercidae : Blattodea) c. 150 mya (late Jurassic), the termites developed advanced social nature and caste system, which their ancestors lacked. They live in colonies with a large number of workers and soldiers that are usually sterile. Colonies have fertile males, the ‘kings’, and one or more fertile females, the ‘queens’. Pheromones maintain the caste system, preventing all but a few termites from becoming reproductives and the rest will remain sterile, assuring a division of labour (Simpson et al., 2011). Primary reproductives and secondary reproductives are the two types of reproductives found in the Isoptera. Nymphs give raise to adults, which mate and reproduce the primary reproductive, hemimetabolically. In some species, such as Coptotermes, when primary reproductives are either lost or immatures are separated from their parent colony, nymphs or workers (= pseudergates) can give rise to either a king or a queen, referred to as secondary reproductives (Myles, 1999). |
With the increased trade of wood, more termites are becoming invasive in different countries; India is no exception. Therefore, anticipating this potential megaproblem strict policy measures are vitally necessary. In this article, I have made an effort to enumerate the potentially invasive termite species in India, based on wood import from different countries, especially in the recent past. Monitoring and treatment of wood material imported are critical to minimize entry of exotic Isoptera into India. Examples of success stories restricting the entry of nuisance species of Isoptera have also been highlighted in this article. The importance of both morphological and molecular taxonomy of Indian Isoptera for correct determination of taxa enabling better management is also indicated. |
Characteristics of invasive termites Although various termite species are considered nuisance organisms of different crops (Rana et al., 2021), forest (Junqueira and Florencio, 2018) and urban ecosystems (Olaniyan et al., 2015), all of these are not invasive. Most of the invasive Isoptera have three qualities in common: (i) they eat wood, (ii) construct nests in the wood, and (iii) quickly generate secondary reproductives (Evans et al., 2013; Donovan et al., 2001). Many a time, in combination of the above, the termites have an increased likelihood of producing reproductively viable offspring. These qualities are especially common in the Kalotermitidae and Rhinotermitidae, which collectively constitute 23 species in the invasive termites list, in genera such as Cryptotermes, Heterotermes, and Coptotermes (19 species) (Table 1). The Termitidae comprising c. 70% of the Isoptera, includes only one invasive species, Nasutitermes corniger. The innate characters that make this species invasive need to be studied. Wood eating In general, four types of food habits occur among the Isoptera (Donovon et al., 2001; Yamada et al., 2007; Jones and Eggleton, 2000; Kaur et al., 2013; Kaur et al., 2017). Type I and II eat undecomposed plant matter such as wood, grass, and leaf litter. Those of type III and IV eat decomposed plant matter; those of type III eat on the decomposing wood either covered with or embedded within soil (plant matter–soil interface), whereas those of type IV are exclusive soil feeders. Those of types III and IV make up c. 50% of all Isoptera. Wood eating is a common characteristic among the known invasives, that means they belong to either Type I or Type II. Nest constructing Nest construction is classified into four categories (Abe, 1987): Single-piece nesters: Isoptera of this category nest and forage in either one piece of wood or wood pieces bound tightly together and feed within: e.g., species of Cryptotermes (Kalotermitidae) and Archotermopsidae Intermediate-piece nesters: Colonize a single piece of wood, but they move out searching food material and eat pieces of wood, where they construct nests and start new nests once the originally occupied piece of wood is completely eaten: e.g., species of Coptotermes (Rhinotermitidae). Separate-piece nesters: build a nest in soil or nest separate from their food and forage away from their nest to find food: e.g., mound-building Macrotermitinae (Termitidae). Continuously mobile (no permanent nest): Do not settle in a nest permanently and are mobile continuously. Found among the Type III and IV soil-feeders that eat their way through the soil. A majority of the invasive species are either single-piece nesters (12— Archotermopsidae and Kalotermitidae) or intermediate-piece nesters (Rhinotermitidae) (Table 1). The possible reason is either presence of all the castes within the nest or the ability to produce viable secondary reproductives (explained in the following paragraph). Can generate secondary reproductive If the nymphs or workers develop as reproductives, they are collectively referred as the nymphoids and ergatoids, respectively. Here sexual maturity is achieved without attaining a fully winged adult stage, and hence they are ‘secondary reproductives’. There may be a situation when an infested wood log is transported, a small number of nymphs may be carried from their original habitat, there is a chance that secondary reproductives produced from those nymphs in a new habitat, making that species invasive. These few nymphs carried from the infested material could act as propagules. Because of this trait, many termite species are potential invaders in new habitats even though primary reproductives are not transported. Number of invasive termite species Termites were not thought to be invasive previously, despite accounts of their spread away from their native ranges. A first worldwide enumeration by Gay published in 1969 recorded 17 species of termites as invasive with the evidence of their establishment in new habitats. All the 17 determined species appear to be associated with either buildings or cultivated crops. Presently 28 species of termites are considered invasive worldwide, (Evans et al., 2013). This implies that the numbers of invasive species have increased from 17 in 1969 to 28 today and are expanding their geographic locations. In the past 50 years, 14 species have been added to the list; 10 of them have extensive distributions and four have no reported changes in distribution, and three species are no more considered invasive (Evans et al., 2013). It appears among the invasive termite species, most are pestiferous in urban areas, although six species have been found to invade natural forests (Evans et al., 2013; Chouvenc et al., 2016). Transportation of live stages of termites Most wood feeding termites (Kalotemitidae, Rhinotermitidae and Archotermopsidae) display the above-said three qualities and they can be transported from their original habitats to new habitats through the movement of wood. Many countries export wood and wooden material to different parts of the world. If the transported wood includes any developmental stage of the Isoptera, then they serve as propogules and become secondary reproductives (Evans, 2011; Myles, 1999). A recent report with dead stages of C. gestroi in the seaport of Goa (15.50° N, 73.83° E) (Venkateshan et al, 2021) from an imported wood consignment sends an alarm signal and seeks a close monitoring of the invasive Isoptera. Species-rich Termitidae cannot produce secondary reproductive, hence they lack the invasive capability. Places of origin and continents invaded Initial reports of termite invasion were mainly restricted to movement of Isoptera from continents to nearby islands (Gay, 1969). This is attributed to free trade of wood products. However, the reverse could be true, since species such as Cryptotermes brevis and C. formosanus, have reached the coast of Queensland, Australia, and Hawaii (USA) in the North Pacific Ocean and have invaded inland as well (Constantino, 2002; Austin et al., 2006; Jenkins et al., 2001). Among the 28 invasive species known to date a perusal of subcontinents or land mass from which the invasive species originated, Indo-Malayan region includes a maximum of seven species, indicating many endemic species have moved from this region to different parts of the world (Table 1). This is followed by South America (six), Australia (five), Africa (three), North America (two), the Caribbean Islands (one), East Asia (one), and Europe (one). Heterotermes perfidus and Coptotermes truncatus are of unknown origin. Similarly, a perusal of invaded subcontinents indicates that the islands of the Pacific Ocean are the most invaded that include 13 species, followed by the Caribbean islands (nine), North America (eight), Indian Ocean (six), South America (five), Australia (four), Atlantic Ocean islands (four), East Asia (three), Africa (three), South and Southeast Asia (two), and Europe (one) (Evans et al., 2013). Coastal regions are more prone to invasion by the Isoptera and inflict damage to built-structures consisting of wood and crop ecosystems (Ferreira et al., 2013; Szalanski., 2004). The probable reason for this pattern is that the wood is mostly imported through sea-routes and the invasive Termitidae first establish themselves in coastal areas, and then propagate inwards. Buczkowski and Bertelsmeier (2017) employed two alternative representative concentration pathways (RCPs) to predict climate scenarios: RCP 4.5 and RCP 8.5, and two projection years (2050 and 2070 AD) to offer the first worldwide risk assessment for 13 of the world’s most invasive termites. Representative concentration pathways (RCPs) are widely used to describe different climate future depending on the volume of greenhouse gases (GHG) emitted in the years to come and thereby predict population dynamics. Buczkowski and Bertelsmeier’s findings reveal that, regardless of the climate or year, most of the species will have a substantial rise in their global spread by 2050 AD. The most appropriate places for invasion are the tropics and subtropics. All the continents have large land spaces and natural conditions suitable for more than four species to coexist. Climate change, growing urbanization, and accelerating economic globalization, working singly or in concert, are anticipated to exacerbate the enormous economic and ecological damage caused by invasive termites. Invasive termites: an Indian perspective India, being one of the largest importers of wood and wood-related material, is vulnerable to Isoptera invasives. Its tropical and subtropical climates also support rapid multiplication of individuals after they arrive. Earlier Cryptotermes dudleyi (Kalotermitidae), C. havilandi (Kalotermitidae), and C. gestroi (Rhinotermitidae) were the invasives known in India. Among these, the taxonomic authenticity of C. dudleyi is confirmed presently. The incidence of C. havilandi is questionable, in spite of its reported spread from India to Bangladesh (Maiti, 1983, Bose 1984). As per Krishna et al. (2013), a world authority on the Isoptera, it is distributed in West Africa (from Sierra Leone to Nigeria), Brazil, Trinidad, the Guianas, Barbados and other west-Indian Islands. Similarly, C. gestroi’s original locality is Myanmar (Wasmann, 1896) and its report in Assam (Roonwal and Chhotani, 1965) and in north-eastern Puducherry (Harit et al., 2014) needs validation by further collection and molecular determination. Not many serious efforts have been made to determine the taxonomic identity, distribution, and damage of these species in the Indian landscape. There is no NCBI sequence for this species submitted for specimens collected from India, which may offer a valid confirmation of its presence in India. Originally described from Myanmar, C. gestroi is considered to occur in north-eastern India and Thailand. This species inflicts serious economic losses by feeding on wood in built structures in other Asian countries, such as Malaysia, Taiwan, Indonesia and in Brazil, the Caribbean islands, and in the peninsular Florida, USA. Coptotermes gestroi is the predominant termite species attacking buildings in urban area of Taiwan (Sornnuwat, 1996). Up to 27% of the trees in the city of São Paulo, Brazil, found infesting contributed by four subterranean species of termites but C. gestroi (Wasmann) being the dominant species (Zorzenon and Campos, 2014). However, such a serious damage has not been reported from India and hence intensive surveys and data collection on damage by the Isoptera may shed light on its presence and its impact on the economy. Venkateshan et al. (2021) offer substantial evidences of both morphological and molecular characters of C. gestroi collected from the wooden packing material of a consignment received in Goa coming from Harrisonburg, Virginia, USA. In another report made in December 2018, a few Isoptera were received for identification from Visakapatnam Plant Quarantine Station collected from the timber of Maclora tinctoria (Moraceae) imported from Guyana (South America). In July and September 2019, I received different specimens of Isoptera for determination intercepted from the wood of a species of Gmelina (Verbenaceae) in Tuticorin port (Tamil Nadu) imported from Columbia, M. tinctoria and a species of Erythrophleum (Fabaceae) wood logs from Suriname. Using morphological and molecular characters, the intercepted specimens were identified as C. testaceus and C. sjöstedti. Coptotermes testaceus’s 16S rRNA gene sequences were deposited with NCBI GenBank with accession numbers MK559590, MK559591, MK559592, and MK559593. Whereas C. sjöestedti 16S rRNA gene got NCBI GenBank accession number MN540914 (Nagaraju et al., 2020). Such clarity is needed in taxonomy for identifying and reporting of any termites intercepted in India. Potential invasive termites in India The spatial spread of Termitidae is increasing due to trade, urbanization, and climate change (Buczkowski and Bertelsmeier, 2017). Extrapolating this scenario, India could be invaded by another 8-10 species by 2050 AD as predicted in the RCP 4.5 scenario, indicating a greater risk and potential damage to urban and agricultural ecosystems. India imports timber from South America, Africa, southeast Asia and from the Caribbean Island nations as evident in the data of noncompliance report of pest interceptions (Directorate of Plant Protection, Quarantine & Storage website, updated monthly http://plantquarantineindia.nic.in/PQISPub/html/ncrep.htm (accessed on 10 December 2021). There is a need for a thorough scrutiny of every wooden log imported into India. The countries mentioned have a high diversity of invasive Isoptera, which have the potential to enter and cause serious ecological and economic damage in India (Table 2). A list of potential invasive species from where wooden logs were imported into India in 2019 is presented in Table 2. This list is not comprehensive but will serve as a base for an early detection of species and to look for the possible entry of potentially damaging species from those countries. Scope of integrative taxonomy Because of their wide distribution and intraspecific character variations, taxonomic validity of many species of the Isoptera remains unclear. For example, Coptotermes, currently includes 69 named species, and among them, only 21 are considered valid (Chouvenc et al., 2016). According to Chouvenc et al. (2016), most of the described Chinese species of Coptotermes are invalid and need revision. Species identification in the Termitoidae is principally based on the measurement of different body parts of soldiers and hence intraspecific variation is highly possible. Hence adapting molecular taxonomy using more than one mitochondrial gene, such as mtCo1, 12S rRNA, and 16S rRNA and nuclear genes such as 28S rRNA, 18S rRNA may provide better clarity on valid species. There is a dire need to barcode all the morphologically identified Isoptera. Currently, barcode sequences are available only for approximately 70 Indian taxa, out of more than 300 species known. In this article, efforts have been to list the available NCBI submissions so far made from India. Apparently, the genes targeted were 12s rRNA and 16s rRNA, that are not robust for species delimitation. A few submissions are with Mt COI and Mt COII used universally for molecular identification of insects. Moreover, there is a need of high-throughput mtCOI or mt COII DNA extraction procedure so as to determine valid species of the Isoptera. Vidyashree et al. (2018) successfully sequenced 12 Isoptera species from the Western Ghats and developed a DNA extraction procedure. However, they reported for 12S rRNA and hence there is a need of standardization of preservation technique of collected termite specimens and storage for successful DNA extraction. The following example may clarify the imperative need for molecular data in resolving taxonomic confusions in the Indian species of Coptotermes. A total of seven species are known from India (Rajamohana et al., 2019): C. beckeri, C. ceylonicus, C. gauri, C. gestroi, C. heimi, C. kishori, and C. premarasmii. Among these, the presence and distribution of C. gestroi and C. travians are doubtful. Among the remaining five, taxonomic validity of C. beckeri, C. ceylonicus and C. kishori is uncertain, because they are possibly junior synonoms of C. amanii, C. brunneus and C. kalshoveni (Chouvenc et al., 2016). Among these, C. amanii is of African origin, C. brunneus is of Australian origin and C. kalshoveni is a serious nuisance organism in Indonesia and Malaysia. This example may throw light on the need for an integrated taxonomy of the Indian Isoptera. There is a greater need for extensive collection, morphological and molecular characterization of Indian Isoptera. Threats to biodiversity The ecological and economic importance of the Isoptera is extensive. The most positive environmental relevance is their role as soil engineers. The most important negative relevance is their activity as nuisance organisms since they damage built structures and agricultural crops, a majority of them belonging to the Termitidae, Rhinotermitidae, and Kalotermitidae (Jouquet et al., 2018). In recent decades the tendency of expansion of the range of harmful Isoptera has led to an increase in economic damage. Invasive termites spread with infested timbers and invade human environments, before spreading into natural ecosystems. A recent study utilized occurrence data and climate modeling to predict the potential habitats of C. formosanus and C. gestroi in Florida and demonstrated that future distribution projections for both species are influenced by urban development and climate change (Buczkowski and Bertelsmeier, 2017). Another negative outcome of increased Isoptera invasions is a potential increase in pesticide use in urban and natural landscapes. Although chemicals are hazardous to the environment, farmers and structural engineers all over the world use them extensively for the management of agricultural pests and in structures. Termites are eusocial, live in nests constructed with well protected soil or many inches below the soil surface and move in galleries which themselves protected from outside threats hence management of termites with termiticides is a difficult task. However, some termiticides such as imidacloprid, chlorpyrifos, fipronil, spinosad, chlorfenapyre, bifenthrin, cypermethrin, permethrin, disodium octaborate tetrahydrate, calcium arsenate, lindane, endosulfan, and chlorantraniliprole have been used worldwide for the management of termites (Ahmad et al., 2019). Leaching of termiticide is toxic to non-target organisms, due to indirect accumulation, which simultaneously affects human population through food chains (Arias-Estévez et al., 2008). The known 28 invasive species have the potential to expand in the range of distribution, just as 10 of the 17 known invasive species have expanded between 1969 and today (Table 1). The spatial spread of the invasive Isoptera is a consequence of a combination of intrinsic and extrinsic factors that shape the population dynamics of the involved species. Intrinsic factors include dispersal, growth, survival, and reproductive constraints dictated by the species’ physiological capabilities. Extrinsic factors include factors such as the spatial and temporal availability of suitable habitat for survival, growth, and reproduction. Human-induced environmental changes, most notably climate change and urbanization, are likely to affect both intrinsic and extrinsic factors. For example, invasive termites have been shown to adapt their reproductive phenology in response to climate change. In parts of Florida, the dispersal flight season of C. formosanus and C. gestroi has begun to overlap due to changes in local climate. Mating pairs of heterospecific individuals were observed in the field with C. gestroi males preferentially engaging in mating with C. formosanus females rather than females of C. gestroi. This leads to hybridization between the two species and the potential evolution of highly destructive “super-termites” due to hybrid vigour (Chouvenc et al., 2016). Management of invasive termites In most cases of invasive Isoptera, attempts were made to eradicate them applying synthetic pyrethroids and organochlorine insecticides, and by baiting. Baiting has been a successful tactic using area-wide management strategy. However, many attempts to eradicate the invasive Isoptera have proved futile (Bravey and Verkrek, 2010). Thus, only instances of successful management refer to those of C. formosanus in South Africa and C. frenchi in New Zealand (Chouvenc et al., 2016). Therefore, monitoring their entry and successful management are the viable options presently. At the time of shipment, log fumigation must reduce the risk of introductions. It is possible to use a less environmentally harmful fumigant like sulfuryl fluoride. A strict policy for the importing country and strict monitoring of imported material could minimize the intrusion into a new geographical region, reducing both economic and ecological risks. Imported wood should be fumigated with methyl bromide @ 48g/ m3 for 24 h to destroy infestation to avoid their entry (https://plantquarantineindia.nic.in/ accessed on 10 December 2021). The treatment is as per the Plant Quarantine (Regulation of Import into India) Order, 2003 which regulates import and prohibition of import of plant and plant products into India and amended recently (S.O. 3686 (E), dated 9 September 2021). The treatment should be endorsed on Phytosanitary Certificate issued at the country of origin or re-export (https://plantquarantineindia.nic.in/ accessed on 10 December 2021). Top Conclusions Tropical, subtropical conditions of India are highly congenial for the rapid multiplication of termites. If any species is introduced accidentally, there is every chance of successful establishment. Hence monitoring, observation, reporting their identity and intervention to avoid the establishment is the need of the hour. Training and recruiting taxonomists with a centralized diagnostic centre is one possible solution to strengthen the activity of the quarantine stations. |
Top Tables | Table 1.: Invasive Isoptera, their country or bioregion of origin, and the country or continent invaded
| | Name of the species | Family | Native country or bioregion | Country or continent invaded and (probable year of invasion) | References | | Mastotermes darwiniensis | Mastotermitidae | Northern Australia | Papua New Guinea (before 1959) | Gray, 1968; Thistleton et al., 2007 | | Zootermopsis angusticollis | Archotermopsidae | Western North America | Hawaii (1999) | Haverty et al., 2000; Grace et al., 2002 | | Zootermopsis nevadensis | Archotermopsidae | Western North America | Kawanishi, Japan (2000) | Kiritani and Morimoto, 2004 | | Porotermes adamsoni | Stolotermitidae | Southeast Australia | New Zealand 1941 | Bain and Jenkins, 1983; Phillip et al., 2008 | | Incisitermes immigrans | Kalotermitidae | Pacific coast of Panama to Peru | Pacific Islands, Polynesia, Hawaii, Japan (1995) | Gay, 1969; Constantino, 1998; Grace et al., 2002; Kiritani and Morimoto, 2004 | | Incisitermes minor | Kalotermitidae | Southwestern United States and northern Mexico | Japan (1975), Eastern United States (1995); Toronto Canada (1989), Ninghai, Zhejiang Province, China (1937), Hawaii (1999) | Grace et al., 1991; Scheffrahn et al., 2001; Xie et al., 2001; Grace et al., 2002 | | Kalotermes banksiae | Kalotermitidae | Southeast coast of Australia | New Zealand (1942) | Bain and Jenkins, 1983 | | Glyptotermes breviconis | Kalotermitidae | Southeast coast of Australia | New Zealand (Pre 1983), Fiji (pre 1942) | Gay, 1969; Bain and Jenkins, 1983; Evenhuis, 2007 | | Cryptotermes brevis | Kalotermitidae | Coastal deserts of Peru and Chile | Egypt, Queensland, Australia (≃ 1941), Azores (2002), Canary Island, Lisbon, Portugal (87) | Peters, 1990; Borges et al, 2007; Scheffrahn et al., 2009; Nunes et al., 2010; Ferreira et al., 2013 | | Cryptotermes cynocephalus | Kalotermitidae | Philippines | Throughout Southeast Asia, Australia (before 1942), Hawaii (2000), Sri Lanka | Gay and Watson, 1982; Scheffrahn et al., 2000; Hemachandra et al., 2012 | | Cryptotermes domesticus | Kalotermitidae | Southeast Asia | China, Australia (before 1942), Panama in Central America | Gay and Watson, 1982; Evans, 2010 | | Cryptotermes dudleyi | Kalotermitidae | Southeast Asia | India (Orissa in Lower Bengal) and Bangladesh (before 1950), East Africa, northern Australia (before 1942), Caribbean islands (Jamaica and Trinidad), South America (Panama, Costa Rica, Colombia and Brazil) | Williams, 1976; Gay and Watson, 1982; Constantino, 1998; Scheffrahn and Krecek, 1999; Constantino, 2002; Schabel 2006 | | Cryptotermes havilandi | Kalotermitidae | Tropical West Africa | Caribbean Islands, Brazil, East Africa, Tanzania, Indian Ocean (Madagascar), India and Bangladesh | Maiti, 1983; Bose, 1984; Constantino, 1998; Scheffrahn and Krecek, 1999; Schabel 2006; Scheffrahn et al., 2009 | | Heterotermes convexinotatus | Rhinotermitidae | South America, from Mexico to Argentina | Greater (Hispaniola and Puerto Rico) and Lesser Antilles (Antigua, Barbados etc) and in Gal'apagos | Peck, 2001; Szalanski, 2004 | | Heterotermes perfidus* | Rhinotermitidae | Unknown origin | south Atlantic Ocean | – | | Heterotermes philippinensis | Rhinotermitidae | Philippines | Madagascar and Mauritius | Cachan, 1949 | | Heterotermes tenuis | Rhinotermitidae | Central America | Lesser Antilles; Trinidad and Tobago | Szalanski, 2004 | | Heterotermes sp. | Rhinotermitidae | Probably Caribbean Island | Miami, Florida | Szalanski, 2004 | | Reticulitermes flavipes | Rhinotermitidae | Eastern United States | Canada, Bahamas (1998), Europe (Vienna- 1837), Germany (before 1937), and South western France (before 1840), South America (Uruguay -1960), Chile (1986) | Grace et al., 1991; Scheffrahn et al., 1999; Dronnet et al., 2005; Austin et al., 2005; Su et al., 2006 | | Reticulitermes grassei | Rhinotermitidae | Southwestern Europe (France and Spain) | Britain (Saunton, 1994), Faial Island of the Azores (2000) | Jenkins et al., 2001; DeHeer et al., 2005 | | Coptotermes acinaciformis | Rhinotermitidae | Australia | Auckland and New Plymouth, New Zealand (1930), Suva, Fiji (1939) | Phillip et al., 2008; | | Coptotermes curvignathus | Rhinotermitidae | Southeast Asia | Southern China | Xie et al., 2001 | | Coptotermes formosanus** | Rhinotermitidae | Southern China and Taiwan | Continental United States (≃ 1950s), Japan (1700), Hawaii (1907), South Africa (1974) | King and Spink, 1969; Vargo et al., 2003; Sun et al., 2007 | | Coptotermes frenchi | Rhinotermitidae | Southern Australia | New Zealand | Miller, 1941 | | Coptotermes gestroi | Rhinotermitidae | Southeast Asia | Mauritius (1936), R'eunion Island (1957), Taiwan (2001), Hawaii (1963), Polynesia 1999) and Micronesia and Fiji (before 2009). Mexico (2000), Florida (1996), Greater Antilles and lesser Antilles (1937), India | Roonwal and Chhotani, 1965; Scheffrahn et al., 1994; Kirton and Brown, 2003; Tsai and Chen., 2003 ; Vargo et al., 2003; Scheffrahn and Su., 2005; Li et al., 2009 | | Coptotermes sjostedti | Rhinotermitidae | Tropical West Africa | Lesser Antilles (Guadeloupe Island, 1999) | Scheffrahn et al., 2005 | | Coptotermes truncates* | Rhinotermitidae | Unknown origin | Madagascar, Republic of Seychelles (1897) | – | | Nasutitermes corniger | Termitidae | Central to South America and the Caribbean Islands | Abaco Island of the Bahamas, Florida (before 2001), New Guinea | Scheffrahn et al., 2005; Evans, 2010 |
|
| Not a valid species; | The establishment and spread of this species in the United States is the best-documented termite invasion | | | Table 2.: Potential invasive Isoptera into India*
| | Country | Native or invasive species distributed at present | | Brazil | Nasutitermes corniger, Incisitermes immigrans, Cryptotermes havilandi, Cryptotermes brevis | | Cameroon | Coptotermes sjostedti, Cryptotermes havilandi, Cryptotermes brevis | | Central African Republic | Coptotermes sjostedti | | Colombia | Heterotermes convexinotatus, Heterotermes tenuis | | Congo | Cryptotermes havilandi | | Costa Rica | Incisitermes immigrans, Nasutitermes corniger | | Ecuador | Incisitermes immigrans, Nasutitermes corniger | | Equitorial Guinea | Coptotermes sjostedti, Cryptotermes havilandi | | Ghana | Coptotermes sjostedti, Cryptotermes havilandi | | Malaysia | Coptotermes curvignathus, Coptotermes gestroi | | France | Reticulitermes flavipes | | Guyana | Nasutitermes corniger, Termes hispaniolae, Cryptotermes brevis | | Mexico | Incisitermes minor, Heterotermes convexinotatus | | Papua New Guinea | Mastotermes darwiniensis, Cryptotermes cynocephalus | | Suriname | Nasutitermes corniger, Coptotermes testaceus | | Trinidad and Tobago | Cryptotermes brevis, Cryptotermes domesticus, Cryptotermes dudleyi, Cryptotermes havilandi, Coptotermes testaceus, Heterotermes tenuis, Heterotermes convexinotatus, Nasutitermes corniger, Termes hispaniolae | | USA | Cryptotermes brevis, Coptotermes formosanus and C. gestroi | | Benin | Coptotermes sjostedti, Cryptotermes havilandi | | *Ecuador | Heterotermes sp. | | Gabon | Coptotermes sjostedti, Cryptotermes havilandi, Cryptotermes brevis | | Honduras | Incisitermes immigrans, Nasutitermes corniger | | Indonesia | Coptotermes curvignathus, Coptotermes gestroi | | Kenya | Cryptotermes dudleyi | | New Zealand | Cryptotermes brevis, Kalotermes banksiae Coptotermes acinaciformis, Porotermes adamsoni, Glyptotermes brevicornis | | Nigeria | Coptotermes sjostedti, Cryptotermes havilandi | | Panama | Incisitermes immigrans, Nasutitermes corniger | | Philippines | Heterotermes philippinensis, Cryptotermes cynocephalus, Cryptotermes havilandi | | Sri Lanka*** | Cryptotermes cynocephalus, Cryptotermes domesticus, Cryptotermes perforans, Coptotermes formosanus | | Sudan** | Coptotermes sjostedti | | Panama | Cryptotermes dudleyi, Cryptotermes domesticus | | South Africa | Coptotermes formosanus, Cryptotermes brevis | | Suriname | Nasutitermes corniger, | | Togo | Coptotermes sjostedti, Cryptotermes havilandi, Cryptotermes brevis | | Uruguay | Reticulitermes flavipes |
|
| Based on Evans et al. (2013); | Dahlsjö et al., 2020; | Pears et al., 1995; | Hemachandra et al., 2015 | | | Table 3.: NCBI GenBank database available for Indian Isoptera
| | Species | Gene | NCBI GenBank accession number | Authors | Remarks | | Anacanthotermes viarum | 28S ribosomal RNA | JQ957910 | Deivendran et al., 2012 | | | Glyptotermes ceylonicus | 12S ribosomal RNA | MZ191059 | Joseph and Mathew, 2021 | | | mtCOII | MZ191060 | Joseph and Mathew, 2021 | | | Neotermes koshunensis | 12S ribosomal RNA | KM657485 | Ramya et al., 2014 | | | Neotermes nilumburensis | 16S ribosomal RNA | MZ234128 | Kalleshwaraswamy et al., 2021 | | | Neotermes viraktamathi | 16S ribosomal RNA | MW699667 | Kalleshwaraswamy et al., 2021 | | | Coptotermes gestroi | 16S ribosomal RNA | MZ540797 | Ranjith et al., 2021 | | | mtCOI | MW575256 | Ashika and Venkatesan, 2021 | | | Coptotermes heimi | | KT828711, KT828711 | Mahapatro and Singh., 2016 | | | 12S ribosomal RNA | KT828710, KT828709 | | | KU665478 | | | | | KR078331 | Mahapatro and Singh, 2015 | | | | KT820660 | Mahapatro and Singh, 2015 | | | | EU553818 | Sobti et al., 2008 | | | | EU553816 | Sobti et al., 2008 | | | | AY558908 | Scheffrahn et al., 2004 | | | 16S ribosomal RNA | MW680951 | Kalleshwaraswamy et al., 2021 | | | GQ422885 | Salunke et al., 2009 | | | GQ422882 | Salunke et al., 2009 | | | | EU553817 | Sobti et al., 2008 | | | 28S ribosomal RNA | JQ957911 | Deivendran et al., 2012 | | | Coptotermes sjoestedti | 16S ribosomal RNA | MN540914 | Mahadeva swamy et al., 2019 | | | Coptotermes testaceus | | MK559593 | Mahadeva Swamy et al., 2019 | | | 16S ribosomal | MK559592 | Mahadeva Swamy et al., 2019 | | | RNA | MK559591 | Mahadeva Swamy et al., 2019 | | | MK559590 | Mahadeva Swamy et al., 2019 | | | Coptotermes sp. | mtCOII | MN913607 | Tiwari et al., 2020 | | | Heterotermes balwanthi | 16S ribosomal RNA | KU574658 | Vidyashree et al., 2016 | | | Heterotermes indicola | 12S ribosomal RNA | KT820657 | Mahapatro and Singh., 2016 | | | KF170428 | Mahapatro and Kumar, 2013 | | | KF769546 | Poonia and Sharma, 2013 | | | KM077441 | Poonia and Sharma, 2014 | | | 16S ribosomal | HF968496 | Mahapatro, G.K. and Kumar, 2013 | | | MZ183973 | Kalleshwaraswamy et al., 2021 | | | RNA | KF170427 | Mahapatro and Kumar, 2013 | | | KF769554 | Poonia and Sharma, 2013 | | | Heterotermes malabaricus | 16S ribosomal RNA | KU574645 | Vidyashree et al., 2016 | | | MZ540855 | Ranjith et al., 2021 | | | MZ558168 | Santhrupthi et al., 2021 | | | OK285073 | Joseph and Mathew, 2021 | | | mtCOII | OK284904 | Joseph and Mathew, 2021 | | | Hypotermes makhamensis | Mt CoI | KT898536 | Patel and Jadhav., 2015 | This species is not reported from India (native to Thailand, Vietnam and Cambodia). Hence needs collection details and morphological authentication | | KT898532 | | KT898526 | | KT724954 | | Mt CoI | KY614388 | Murthy and Lubna., 2017 | | Hypotermes xenotermitis | 12S ribosomal RNA | KY825251 | Murthy and Lubna, 2017 | | KU687341 | Murthy et al., 2016 | As per the authors, specimens were collected from south India. This species is not reported from south India (reported only from North India). Needs morphological authenticity and morphometry data. | | KY293420 | Murthy and Lubna, 2016 | | KT224387 | Murthy et al., 2016 | | KX646190 | Murthy et al., 2016 | | KU687340 | Murthy et al., 2016 | | KT898553 to KT898566 | | | KT898535 to KT898551 | | | mtCOI | KT898527 to KT898531 | | | KT898507 to KT898525 | Patel and Jadhav, 2015 | | KT887698 to KT887717 | | | 16.KT879833 to KT879849 | | | KT879830, KT724955, KT898533 | | | Macrotermes convulsionarius | 16S ribosomal RNA gene | MZ540857 | Ranjith et al., 2021 | | 28S ribosomal RNA | JQ957912 | Deivendran et al., 2012 | | Microtermes incertoides | 16S ribosomal RNA | MZ571483 | Ranjith et al., 2021 | | Microtermes mycophagus | KU665477 | Mahapatro and Singh, 2016 | | KP765715 | Mahapatro and Singh, 2015 | | KP748241 | Mahapatro and Singh, 2015 | | KF769547 | Poonia and Sharma, 2013 | | KT820658 | Mahapatro and Singh, 2015 | | KM657479 | Ramya et al., 2014 | | JX263668 | Singla et al., 2012 | | JX045651 | Singla et al., 2012 | | EU553821 | Sobti et al., 2008 | | EU553819 | Sobti et al., 2008 | | 16S ribosomal RNA | KF769555 | Poonia and Sharma, 2013 | | EU553822 | Sobti et al., 2008 | | EU553820 | Sobti et al., 2008 | | Microtermes obesi | 12S ribosomal RNA | EU551158 | Sobti et al., 2008 | | KM657488 | Ramya et al., 2015 | | KT820661 | Mahapatro and Singh, 2015 | | 16S ribosomal RNA | MZ558085 | Santhrupthi et al., 2021 | | EU306616 | Sobti et al., 2007 | | KU574654 | Vidyashree et al., 2016 | | mtCOI | EU242522 | Sobti et al., 2007 | | EU306614 | Sobti et al., 2007 | | mtND1 | EU306613 | | EU306615 | | Microtermes unicolor Microtermes sp. | 12S ribosomal RNA | JX263667 | Singla et al., 2012 | | | ITS-2 | KX495579 | Murthy et al., 2016 | | 16S ribosomal RNA | KM275840 | Poonia and Sharma, 2014 | | 12S ribosomal RNA | KF703855 | Poonia and Sharma, 2013 | | Odontotermes anamallensis | 16S ribosomal RNA | MZ562513 | Santhrupthi et al., 2021 | | Odontotermes assmuthi | 16S ribosomal RNA | KF769556 | Poonia and Sharma, 2013 | | KU574651 | Vidyashree et al., 2016 | | MZ558070 | Santhrupthi et al., 2021 | | 28S ribosomal | JF792835 | Deivendran and Suresh, 2011 | | RNA | FJ966379 | Suresh et al., 2009 | | Odontotermes bellahuniensis | 16S ribosomal RNA | KU574650 | Vidyashree et al., 2016 | | Odontotermes bhagwatii | | KF769552 | Poonia and Sharma, 2013 | | 12S ribosomal | KM523663 | Ramya et al., 2014 | | RNA | KM523662 | Ramya et al., 2014 | | EU551161 | Sobti et al., 2008 | | 16S ribosomal RNA | KF769559 | Poonia and Sharma, 2013 | | EU258632 | Kumari et al., 2007 | | EU258631 | Kumari et al., 2007 | | mtCOII | EU242525 | Sobti et al., 2008 | | mtND1 | EU262586 | Sobti et al., 2007 | | EU262585 | Sobti et al., 2007 | | EU262587 | Sobti et al., 2008 | | Odontotermes boveni | 16S ribosomal RNA | MZ344981 | Kalleshwaraswamy et al., 2021 | | Odontotermes brunneus | 12S ribosomal RNA | KT820659 | Mahapatro and Singh., 2016 | | 16S ribosomal RNA | KF792982 | Poonia and Sharma., 2017 | | MZ540811 | Ranjith et al., 2021 | | 28S ribosomal RNA | JF792836 | Deivendran and Suresh, 2011 | | 12S ribosomal | KF769549 | Poonia and Sharma, 2013 | | RNA | JX263664 | Singla et al., 2001-2 | | Odontotermes ceylonicus | 12S ribosomal RNA | KY908410 | Murthy et al., 2017 | | Odontotermes escherichi | 12S ribosomal RNA | KY495155 | Murthy and Lubna., 2017 | Not reported from India (recorded only from Sri Lanka). Needs morphological authenticity | | mtCOI | KT224389 | Murthy et al., 2015 | | KY495154 | Murthy and Lubna., 2017 | | Odontotermes feae | | KU947966 | Mahapatro and Singh, 2016 | | 12S ribosomal RNA | KY908402 | Murthy et al., 2017 | | KY676779 | Murthy and Lubna, 2017 | | KY908403 | Murthy et al., 2018 | | KR296660 | Mahapatro et al., 2015 | | 16S ribosomal RNA | KU574649 | Vidyashree et al., 2016 | | Odontotermes | 12S ribosomal | KM523667 | Ramya et al., 2015 | | gurdaspurensis | RNA | KM523664 | | | Odontotermes horni | 12S ribosomal RNA | EU551159 | Sobti, et al., 2008 | | | GQ422892 | Salunke et al., 2009 | | GQ422890 | Salunke et al., 2009 | | GQ422889 | Salunke et al., 2009 | | GQ422888 | Salunke et al., 2009 | | 16S ribosomal RNA | GQ422887 | Salunke et al., 2009 | | GQ422886 | Salunke et al., 2009 | | GQ422879 | Salunke et al., 2009 | | EU258629 | Kumari et al., 2007 | | KU574646 | Vidyashree et al., 2016 | | EU258630 | Sobti et al., 2007 | | 28S ribosomal RNA | JF792837 | Deivendran and Suresh, 2011 | | mtCOII | EU242523 | Sobti, et al., 2007 | | Odontotermes obesus | 12S ribosomal | KY908407 | Murthy et al., 2017 | | RNA | KP410731 | Mahapatro et al., 2015 | | EU551160 | Sobti et al., 2008 | | 16S ribosomal | KU574648 | Vidyashree et al., 2016 | | RNA | MZ423304 | Kalleshwaraswamy et al., 2021 | | MN511317 | Amina etal., 2019 | | mtCOI | KY474376 | Murthy and Lubna, 2007 | | MZ823814 | Ranjith et al., 2021 | | mtCOII | EU242524 | Sobti, et al., 2007 | | mtND1 | EU262594 | Sobti, et al., 2007 | | Odontotermes longignathus | Mt CoI | KY930907; | Murthy and Lubna., 2019 | This species is not reported from India (native to south east Asia). Needs morphological authenticity | | KY930908; | | | KY775488 | | | 12S ribosomal | KY495156; | | | RNA gene | KY563712 | Murthy and Lubna., 2017 | | Mt CoI | MN205551 | Alina et al., 2019 | | Odontotermes parvidens | 12S ribosomal RNA | KF769551 | Poonia and Sharma, 2013 | | 16S ribosomal RNA | KF769558 | Poonia and Sharma, 2013 | | Odontotermes redemanni | 12S ribosomal RNA | KF769553 | Poonia and Sharma, 2013 | | 16S ribosomal | KU574647 | Vidyashree et al., 2016 | | RNA | KF792983 | Poonia and Sharma., 2017 | | Odontotermes wallonensis Odontotermes yadevi | 28S ribosomal | JF792838 | Deivendran et al., 2012 | | RNA | | 16S ribosomal | KU574656, | Vidyashree et al., 2016 | | RNA | KU574655 | Kalleshwaraswamy et al., 2021 | | MZ189521 | | Euhamitermes hamatus | Not reported from India (recorded only from Thailand, Malaysia, Singpur, Bangladesh). Needs morphological authenticity | | 12S ribosomal RNA | KM657484 | Ramya et al., 2015 | | Eurytermes buddha | 16S ribosomal RNA | MW678776 | Kalleshwaraswamy et al., 2021 | | mtCOI | MW664866 | Kalleshwaraswamy et al., 2021 | | Ampoulitermes wynaadensis | 12S ribosomal RNA | MZ044682 | Kalleshwaraswamy et al., 2021 | | MZ044681 | Kalleshwaraswamy et al., 2021 | | Grallatotermes niger | 16S ribosomal RNA | MZ262750 | Kalleshwaraswamy et al., 2021 | | Nasutitermes anamalaiensis | 16S ribosomal RNA | KU574659 | Vidyashree et al., 2016 | | MW692352 | Kalleshwaraswamy et al., 2021 | | MW694353 | Kalleshwaraswamy et al., 2021 | | Nasutitermes brunneus | 16S ribosomal | MZ262702 | Kalleshwaraswamy et al., 2021 | | RNA | MZ540856 | Ranjith et al., 2021 | | Nasutitermes indicola | 16S ribosomal RNA | KU574660 | Vidyashree et al., 2016 | | Nasutitermes kali | 16S ribosomal RNA | MZ262715 | Kalleshwaraswamy et al., 2021 | | Nasutitermes octopilis | 12S ribosomal RNA gene | KM657478 | Ramya et al., 2015 | Not reported from India (recorded only from Africa-Guyana). Needs morphological authenticity | | Trinervitermes_ biformis Trinervitermes togoensis | 16S ribosomal | KU574657 | Vidyashree et al., 2016 | | RNA | MZ558074 | Santhrupthi et al., 2021 | This species is not reported from India (native to Africa). Needs morphological authenticity. | | 12S ribosomal | KY569523 | | | RNA gene | KX711183 | Murthy and Lubna., 2017 | | And MtCo1 | | Murthy etal., 2016 | | Amitermes belli | 12S ribosomal RNA | KR078330 | Mahapatro and Singh., 2015 | | 16S ribosomal RNA | MZ269706 | Priya and Gupta, 2021 | | Angulitermes sp. | 12S ribosomal RNA | KP780274 | Mahapatro and Singh, 2015 | | Dicuspiditermes achankovili | mtCOI | MT272760 | Amina et al., 2020 | | MT272755 | Amina et al., 2020 | | MT272750 | Amina et al., 2020 | | Dicuspiditermes fontanellus Dicuspiditermes gravelyi | 16S ribosomal RNA | MZ270643 | Kalleshwaraswamy et al., 2021 | | 12S ribosomal | MZ825163 | Ranjith et al., 2021 | | RNA | MZ825164 | Ranjith et al., 2021 | | 16S ribosomal RNA | MZ270644 | Kalleshwaraswamy et al., 2021 Ranjith et al., 2021 | | | MZ823812 | | Dicuspiditermes obtusus Homallotermes pilosus | 16S ribosomal RNA | MZ270642 | Kalleshwaraswamy et al., 2021 | | mtCO1 | MT272758 | Amina et al., 2020 | | MT272753 | Amina et al., 2020 | | MT272747 | Amina et al., 2020 | | Indocapritermes aruni Krishnacapritermes thakuri | mtCOI | MT272742 | Amina et al., 2020 | | mtCOI | MT272759 | Amina et al., 2020 Amina et al., 2019 | | MN507713 to MN507729 | | Krishnacapritermes dineshan | mtCOI | MN507708 to | Amina et al., 2019 | | MN507712 | | Labiocapritermes distortus | 16S ribosomal RNA | MZ558073 | Santhrupthi et al., 2021 | | | MT272756 | Amina et al., 2020 | | mtCOI | MT272752 | Amina et al., 2020 | | MT272744 | Amina et al., 2020 | | MT272743 | Amina et al., 2020 | | Microcerotermes beesoni | 12S ribosomal RNA | JX263665 | Singla et al., 2012 | | Microcerotermes fletcheri | 16S ribosomal RNA | KU574652 | Vidyashree et al., 2016 | | Microcerotermes pakistanicus | 16S ribosomal RNA | KU574653 | Vidyashree et al., 2016 | | MZ414220 | Kalleshwaraswamy et al., 2021 | | MZ427480 | Kalleshwaraswamy et al., 2021 | | Pericapritermes topslipensis | mtCOI | MT272762 | Amina et al., 2020 | | MT272761 | Amina et al., 2020 | | MT272749 | Amina et al., 2020 | | MT272745 | Amina et al., 2020 | | MT272741 | Amina et al., 2020 | | Procapritermes keralai | mtCOI | MT272748 | Amina et al., 2020 | | MT272757 | Amina et al., 2020 | | Pseudocapritermes fletcheri | 16S ribosomal RNA | MW686909 | Kalleshwaraswamy et al., 2021 | | MW687081 | Kalleshwaraswamy et al., 2021 | | MT272754 | Amina et al., 2020 | | mtCOI | MT272751 | Amina et al., 2020 | | MT272746 | Amina et al., 2020 | | MW672524 | Kalleshwaraswamy et al., 2021 | | Synhamitermes quadriceps | 16S ribosomal RNA | MW680954 | Kalleshwaraswamy et al., 2021 | | mtCOI | MW680303 | Kalleshwaraswamy et al., 2021 | | Neotermes koshunensis | 12S ribosomal RNA gene | KM657485 | Ramya et al., 2015 | Not reported from India (recorded only from Japan). Needs morphological authenticity | | Rinacapritermes abundans | mtCOI | MT274296 | Amina et al. 2022 | | | Rinacapritermes silvius | mtCOI | MT274294 | Amina et al. 2022 | |
| |
|