Isoptera: India
This is an extract from |
Note: This article is likely to have several spelling mistakes that occurred during scanning. If these errors are reported as messages to the Facebook page, Indpaedia.com your help will be gratefully acknowledged.
Contents |
Isoptera
Introduction
Termites or white ants belong to a small onJer of insects, Isoptera. They are one of the most fascinating group of insects due to their primitiveness in orgin in the Cretaceous period, ubiquitous in distribution in the tropics, polymorphic ~aste system, cryptobiotic mode of life helping them pass unnoticed, superb architectural. ability in making nests, unique ecological adjustment, and for their immense economic importance. In search of food and shelter, termites cause considerable damage to wood in forest and agricultmal crops, to timbers in storage and in buildings, to books, clothes and other articles of cellulosic orgin. It is estimated that the amount of loss thus caused by termites in India alone runs to several millions of rupees per year and the loss allover the world must be something colossal.
India being a tropical country with varied physiography. climate and vegetaion, has more than 200 species of tennites, distributed from the plains to the high altitude of the Himalaya. Tennite distribution is generally limited by certain combination of factors, such as, tropical climate, restricted diet of cellulose, consequence of soft body forcing them to live in closed nests, vulnerability to predators and weak flight. The phylogeny of the termite genera is better known than many other groups of animals of comparable size. This is due to constellation of multiple characters available from imago, soldier and worker castes, availability of more than 60 fossil species, receprocal phylogeny of termitophiles, symbiotic flagellates, etc. Moreover, phylogeny in many instances is correlated with ecological, geographical and palaeontological parameters.
The study of termites has been used to illustrate various evolutionary principles, such as, homology, covergence, parallelism, divergence, progressive and regressive, and adaptive evolutionary concepts.
Large number of invertebrate (beetles, flieSl and others) including some fungi, have well adopted to living either in te~ite gut (flagellates) or in their nest from the very bigining of their evolution, leading to interesting relationships. Such intimate relationship with termitophiles and symbiotes has thrown light to understand the reciprocal evolution of both the host termites and their associates.
The cryPtobiotic mode of life in termites has made them unique subject of study. The main atttaction is due to their role in disintegration of wood and cellulose, their place in the food chain of many tropical animals, their close association with fungi, termitophiles and parasites, their unique symbiotic relationship with protozoans and .bacteria; their complex behaviour and nest constuction, their mechanism of caste differentiation, intricate social life, etc.
The 'recognition of the species it) termites is very difficult due to minor variations in morphological characters in all the castes, namely, imagos, soldiers and workers. However, the soldiers, primarily adapted for defence of the colony, exhibit comparatively significant characters for taxonomic identity. As such the museum collections represented by soldiers, are essential for taxonomic purpose. Imagoes and workers associated with the soldiers are identified indirecdy in most of of the cases. However, the generic identification is easy as compared to that of species.
Historical Resume
i) Pre-1900
India has the oldest citation of reference on termites in literature by the name as ghuna (wood destroyer) particularly in the ancient sanskrit text, the Rig Veda (ca. 1350 B.C.); and as kasthaharilca in the Ramayana and other sanskrit literature. The oldest scientific name of a termite species, Termes fatale "from Indiae" appeared in "Systema Naturae" 10th edition, which is considered as the starting point of animal taxonomy published by Linnaeus (1758). Linnaeus recognized three species of termites, and put them under .two different orders, namely, Neuroptera and Aptera. However, the species, Termesfatale is actually from Surinam, South America, and not from India.
Work from• India was initiated in 1779 by Konig on a mound building termite ~south India. This was later on identified as Odontotermes redemanni (Wasmann) by Roonwal (1970) althougb others referred to it as Termes fatalis K6nig (H~gen, 1858; Green, 1913). The other three species dealt with by K6nig (1779) are now in Anacanthotermes Jacobson, Macrotermes Holmgren and Hospitalitermes Holmgren. After KOnig's publication, only 16 species were brought to light upto the end of 19th Century by Rambur (1842); Walker (1853); Hagen (1858, 1859); Brauer (198~); Was mann (1893, 1896) and others.
Classification : Since the time of Linnaeus, the termites were assigned to different Orders, Suborders and Tribes, until Brulle (1832) put them in the order 'Isopteres' The placement of termites in this order was not accepted by the then workers (Latereille, 1802; Walker, 1852 and Hagen 1858). However, after a lapse of many years, Comstock and Comstock (1895) revived the status of the order Isoptera. Credit goes to Hagen (1858) who recognised for the fust time four genera under the Order, namely, Kalotermes Hagen, Termopsis Desneux, Hodotermes Hagen and Termes Linnaeus, and to Froggatta (1896) for recognising four sub-families namely Calotermitinae, Rhinotermitinae, Glyptotermitinae and Termitinae under a single family Termitidae. Thus, the foundation of termite classification was laid during this period.
Catalogues, lists, etc. : The list of Neuropterous insects (including termites) were published by Walker (1853) and Hagen (1858). The latter author also published a synopsis of Neuroptera of Sri Lanka in successive years (1858 and 1859).
ii) 1901-1947
The resident zoologists in India, mainly attached to Imperial Agricultural Research Institute, Pusa; Forest Research Institute, Debra Dun; and Zoological Survey of India, Calcutta, realised the importance of studying these noxius insects and got them identified from foreign experts. Termites from some parts of India and Sri Lanka were worked out by Wasmann (1902) and Desneux (1904, 1906, 1908); those from Sri Lanka by Holmgren (1911), from south India by Holmgren (1912, 1913), from different areas of India by Holmgren and Holmgren (1917),' from Abor Hills by Silvestri (1914), from Barkuda Island by Silvestri (1923), from Sri Lanka by Bugnion•(1912, 1913, 1914), Bugnion and Ferrier (1911), Bugnion and Popoff (1910), John (1925), and Kemner (1926, 1932); from India by Snyder (1933, 1934), from India and Burma by Gardner (1944) and others. As a result, an addition of about 130 species was made during this period.
References to termites were made by different •workers in their books etc. Lefroy (1909) in 'Indian Insect Life' listed 20 species grouped in three genera. Fletcher (1914) listed only three species and Ayyar (1940) in his book on South Indian insects made some casual mention of termites. However, valuable biological informatipn on 50 species is available in the book 'The Ecology and Control ofForest Insects ofIndia and the neighbouring countries' by Beeson (1941).
Classification : At the begining of 20th century, Desneux (1904) compiled the available knowledge in the 'Genera Insectorum' under Isoptera, with inclusion of a single family Tennitidae containing three subfamilies, Calotermitinae, Mastotermitinae and Termitinae. However, Mastotermitinae gained the status of a family later on (Silvestri, 1909). Synder (1949) published a more sound classificatory catalogue, following the contributions of various workers including those from the Indian region (Holmgren, 1910, 1911; Green, 1913; Snyder, 1920, 1933; Light, 1921; Margabandhu, 1934, etc.).
Phylogeny : The phylogenetic aspects of termites were first discussed by Holmgren (1911, 1912) including some genera from the Indian region, who put emphasis on the characteris~c of imago-wodcer mandibles. This has been proved an ideal conservative character on which the present day phylogenetic study is based, supported by some other characters. On the other hand, Hare (1937) based his phylogenetic study on the basis of the development of soldier's mandible, which today seems to be less sound.
Catalogues, lists, etc.: The references of some 30 species of termites from the Indian subregion, could be found in a catalogue compiled by Desneux (1904),44 species from Sri Lanka by Green (1913) and of 119 species from India and Sri Lanka by Margabandhu (1934, 1935).
iii) 1948 -1990
This is the most productive period as far as the termite research is concerned in the entire world including the Indian region. The basic impetus came from two outstanding publications, namely World Catalogue by Snyder (1949) and Phylogeny of World Termite Genera by Ahmad (1950).
The taxonomic work in India gained its maximum momentum during this period under the initiative of Roonwal frrst at the FRI, Debra Dun and later at ZSI, Calcutta and ZSI, Jodhpur. He along with his collaborators namely Sen-Sarma, Kumar Krishna, Chhotani, Chatterjee, Thakur, Thapa, Bose, Maiti, Venna, Rathore, etc, worked on different aspects. The termite research at ZSI, Calcutta flourished from 1956 under Roonwal and his students, Chhotani, Bose and Maiti, with reference to taxonomy, morphology, phylogeny, zoogeography, biology and ecology. Numerous publications were mad~, including a 'Fauna of India' volume on the lower termites by Roonwal and Chhotani (1989). Another volume in this series is awaiting publication.
Revisionary Works : At the begining, emphasis was laid to revise common genera containing disputed species, for example Neotermes Holmg. and Glyptotermes Frogg. were revised by Roonwal and Sen-Sarma (1960), Coptotermes Wasmann by Roonwal and Chhotani (1962), Cryptotermes Banks by Chhotani (1970), Glyptotermes by Chhotani (1975), Stylotermes Hol~g. and Holmg. by Mathur and Chhotani (1959), Mierocerotermes Silvestri by Prashad, Sen-Sarma and Thapa (1967), Microtermes Wasmann by Chatterjee and Thakur (1964), Hypotermes Holmg. by Chatterjee .and Thakur (1963); Eurytermes Wasmann by Roonwal and Chhotani (1966), Nasutit~rmes Dudley by Prashad and Sen-Sarma (1966), Odontotermes Holmg. by Thakur (1981) and Heterotermes Frogg. by Thakur and Sen-Sanna (1979).
State Fauna: Considering the vastness of the Indian subcontinent, State-wise study of termite fauna gained importance. The fauna of Assam region (N.E. India) was published by Roonwal and Chhotani (1962), Rajasthan by Roonwal and Bose (1964), Tripura by Sen-Sarma and Thakur (1979), West Bengal by Maiti (1983), Orissa by Chhotani and Das (1983), and South India by Bose (1984). The fauna of some neighbouring countries were also studied, for instance, the fauna of Paksitan by Ahmad (1955), Indonesia by Roonwal and Maiti (1966), Bangladesh by Akhtar (1975), Bhutan by Roonwal and Chhotani (1977), and Sabah, West Malaysia by Thapa (1982).
Many important discoveries were made during this period. The most significant is the discovery of a family Indotermitidae (Roonwal and Sen-Sarma, 1960), with type genus Indotermes. The validity of this family however, has been questioned (Krishna, 1979). A number of genera, namely, Posteleetrotermes Krishna, Epicalotermes Silvestri, Calearitermes Snyder, Psammotermes Desn., Speeulilermes Wasmann, Anoplotermes Muller, etc., were recorded for the first time from Indian region indicating zoogeographical significance. A total of 157 species were added to the known fauna, to make a total of about 300 species altogether known from the region.
Classification : The foundation of present day Isoptera classification was laid by Synder (1949) and Grasse •(1949) in two separate contributions, which have been generally accepted with some minor modifications. Both the classifications recognized six families, but Mastotermitidae was divided into two families in one (Synder, 1949) and Hodotermitidae into two families in another (Grasse, 1949). This system was further substantiated by phylogenetic study based mainly on the character of imago-worker mandibles.
Out of nine families recognised so far, except two families, Mastotermitidae (confined to Australia) and Serritermitidae (Brazil), all other families, namely Kalotermitidae, Termopsidae, Hodotermitidae, Rhinotermitidae, Stylotermitidae, lndotermitidae and Termitidae occur in the Indian region. The largest family, Termitidae, often referred to as higher termites, contains three¬fourth of the known species. The subfamilies, Termopsinae and Stylotermitinae were raised to family level by Grasse (1949) and by Chatterjee and Thakur (1964) respectively.
Phylogenetic Studies: The first phylogenetic account was proposed by Ahmad (1950) mainly based on the characteristic feature of the imago-worker mandibles. This pattern remained almost unchanged except some additional works done subsequently, e.g. by the study of entiric valve of digestive tract and wing-micro-sculpturing (Roonwal and co-workers 1974-1988).The phylogeny of Kalotermitidae was revised by Krishna (1961), Nasutitermitidae by Sands (1957), Capritermes complex by Krishna (1968), Stylotermitidae and Indotermitidae by Roonwal (1975), Nasutitermitinae by Sen-Sarm~ (1968), Anoplotermes -Speculitermes -complex by Roonwal and Chhotani (1966), and Grallatotermes -complex by ~en-~arma (1966).
Catalogues. lists. bibliographies. reviews etc. : The frrst ever world Catalogue was published by Snyder (1949), which stimulated research around the world. Catalogue of the Indian Isoptera was published by Rattan Lal and Menon (1953), and that of Isoptera collection present in F.R.I., by Roonwal and Pant (1953), which was revised by Mathur and Thapa (1962). Roonwal (1962) gave an excellent review of the systematic work done on world termites (1949-'60), appended with a catalogue and a bibliography. More recently, Chhotani (1972,1977) reviewed the taxonomic work done on termites of the Indian region appended with a catalogue and a bibliography.
Zoogeography: The first analysis of origin, dispersal and zoogeography of the world termites was made by Emerson (1955). Zoogeography of the regional fauna has been discussed by many authors, e.g. of Assam region by Roonwal and Chhotani (1965), Bhutan by Roonwal and Chhotani (1977), Rajasthan by Roonwal and Bose (1964), Andaman and Nicobar Islands by Roonwal and Bose (1965,1970), West Bengal by Maiti (1983), Bangladesh and Pakistan by Akhtar (1974,1975). Zoogeography of some genera, namely, Cryptotermes (Chhotani, 1970), Glyptotermes (Chhotani, 1975) and Odontotermes (Thakur, 1976), were also analysed. The origin, dispersal and evolution of termites in the islands of Andaman and Nicobar have been discussed by Maiti (1977,1979).
Biology: The progress on biological research of Indian species, however, does not commensurate with that of taxonomic research. Some biological notes were incorporated in the faunistic reports. Specific studies on different termites are limited to the nature of infestation in wood and wood products, nest structure, gallery pattern, swarming period, colony development,fungus comb, nest population, etc. The data on these aspects are available in the contributions of Beeson (1941), Mukherjee and Mitra (1949), Roonwal (1970,1979), Sen-Sarma et ale (1975) and Maiti (1983).
The nest structure of Coptotermes (Roonwal, 1954,1959; Roonwal and Chhotani, 1962,1967), Microcerotermes (Roonwal, 1979), Odontotermes (Roonwal, 1973) and arborial carton nests of Nasutitermes (Chhotani and Bose 1979) have been described in detail. The general biology of local termite fauna of Dehra Dun (Mathur and Sen-Sarma, 1959, 1960,1962; Chatterjee and Sen-Sarma 1962; Sen-Sarma, 1962), West Bengal (Maiti, 1983), Rajasthan (Roonwal and Verma, 1973; Roonwal and Rathore, 1974, 1975, 1976, 1978; Roonwal 1975) and penisular India (Roonwal, 1978) are studied.
Ecology: The ecological investigations on Indian termites gained less attention than it deserves. Only a few species have be.cn studied with reference to temparature and humidity responses (Sen¬Sarma, 1969, 1974, Sen-Sarma and Gupta.1968; Sen-Sarma and Misra, 1969, 1971, 1976), termite and fungus interactions (Bakshi, 1962, Mukherjee and Mitra, 1949), termite and symbiotic flagellate association (De Mello 1919,1928,1937,1941; Chakraborty and Banerjee, 1956; Das, 1972, 1974, 1983; Uttangi, 1959, 1962; Mukherjee and Maiti, 1988,1989) and swarming period (Mathur, 1962; Maiti, 1983).
Studies from Different Environs
During the post Independence period, some preliminary work on the composition and faunal strength of termite fauna of different environs has been available. The arid zone fauna of Rajasthan was studied by Roonwal and Bose (1964); that of humid zone of North -east India by Roonwal and Chhotani (1962); of temperate and montane areas of the north-west Himalaya by Chatterjee and Thakur (1967); the deltaic and the sub-Himalayan West Bengal by Maiti (1983); of Bhutan Himalaya by Roonwal and Chhotani (1977); of the peninsular tract by Bose (1984); and the insular areas of the Andaman and Nicobar by Roonwal and Bose (1965, 1970) and Maiti (1977, 1979). Studies on termites of Meghalaya and Tripura are in progress at ZSI, Calcutta.
Estimation of Taxa
Out of nine families recognised so far, seven families are represented so far in the Indian region. The scrutiny of literature shows that about 300 species under 45 genera occur as shown below :
Classified Treatment
Family Tennopsidae
Members of this family inhabit primarily wood. They are divided into three genera allover the world. In India, the family is represented by species, Archotermopsis wroughtoni (Desneux), described in 1904 from the Himalaya. This species has undergone several nomenclatural changes and its biology is worked out better than any other species.
Family Hodotennitidae
Out of five living genera known so far from the world with a total of 12 species only, one genus Anacanthotermes occurs in this region, represented by four species, of which A. baluchistanicus Akhtar is described recently from Pakistan (Akhtar, 1972). The taxonomy and distribution of all the species have been dealt recently by Roonwal and Chhotani (1989).
Family Kalotermitidae
The members of this family strictly inhabit wood in its many forms, including dead portion of live trees. Out of 24 genera known so far, nine genera represented by 56 species occur in this region, which have been extensively studied by Indian workers (Roonwal and Chhotani, 1989). Of these genera, Postelectrotermes Krishna, Calcaritermes Snyder and Epicalotermes Silvestri are recent addition to the Indian fauna (Roonwal and Maiti, 1966; Roonwal and Chhotani, 1983; Chaudhury and Ahmad, 1972). Altogether 33 species are described recently.
Family Rhinotermitidae
The members of this family feed on dead and decaying wood and live in cracks and crevices of the trees, with or without soil connection~ Some species are capable of transporation through floating logs like that of coconuts, and thus got introduced in the coastal areas of different continents. Eight genera represented by 28 species occur in this region. Out of a total of 13 genera known from the world, 12 species have been described as new to science by recent workers, vide Roonwal and Chhotani (1987).
Family S tylotermitidae
This small family contains a single living genus Stylotermes distributed in the Oriental region. It has three segmented tarsi as also in Indotermitidae and also. share some characters of Kalotermitidae and Rhinotermitidae. All the members inhabit the dead and decaying portions of standing trees. Since its establishment by Holmgren and Holmgren, 1913, the family was represented by only species Stylotermes Jletcheri Holmgren and Holmgren, until seven more species have been added by the recent workers.
Family Indotermitidae
This small family is represented by single genus Indotermes Roonwal and Sen-Sarma, (1958). Sinotermes erected by He and Xia (1981) from China has recently been merged under it (Roonwal and Chhotani, 1989). As Such, genus Indotermes contains seven species from South and South¬East Asia. All its members are subterranean in habit.
Current Studies
Active research on termites is being conducted in ZSI, and its Desert Regional Station, Jodhpur, and in FRI, Debra Dun. Recently biological investigations are also being taJcen up in the Department of Zoologyt Vishwa Bharati, Santiriiketan. Some research activities are also in progress in certain Universities in South India.
Expertise India
In ZSI
P. K. Maiti, A. R. Lahiri, N. Saha & P. H. Roy, all of ZSI, M-Block, New Alipur, Calcutta 700053. O. B. Chhotani &O. Chhotani (nee Bose), ZSI, Nizam Palace, 13th floor, 234/4, AJ.C. Bose Road, Calcutta 700 020. N. S. Rathore, ZSI, Desert Regional Station, Paota B Road, Jodhpur (Rajasthan).
Elsewhere
M. L. Thakur &R. S. Thapa, Forest Research Institute, Dehradun (U.P.). P. K. Sen Sarma, Birsa Agricultural University, Kunke, Ranchi (Bihar).
Abroad
W. A. Sand &R. M. C. Williams, British Museum (Nat. Hist.), London. IItJl*m 307 Ie. Krishna, Howard University. Barbar L. Thome American Museum of Natural History, New York. W. G. H. Coaton, J. E. Ruelle and J. L. Sheasby, Plant Protection Research Institute, Pretoria (S. Africa). M. Ahmad and M. S. Akhtar, Punjab University, Lahore (pakistan).
Selected References
Akhtar, M. S. 1975. Taxonomy and Zoogeography of the termites (Isoptera) of Bangladesh. Bull. Dept. Zoo. Univ. Panjab, Art. 7 : 1-199.
Bose, G. 1984. Termite fauna of Southern India. Occ. Pap. Rec. zool. Surv. India, No. 44 : ix + 270 pp.
Chhotani, O. B. 1980. Termite pest of agriculture in the Indian Region and their control. Tech. Monogr. 2001. Surv. India, No.4: 94 pp.
Maiti, P. K. 1983. Termite fauna (Isoptera) of West Bengal, India. Their recognition, biology and ecology. Dcc. pap. Rec. 2001. Surv. India, No. 42 : 152 pp.
Roonwal, M. L. 1970. Termites of the Oriental Region. In : Biology o/Termites (Ed. K. Krishna and F. M. Weesner), Vol. 2: 315-391.
Roonwal, M. L. & Chhotani, O. B. 1962. Termite fauna of Assam Region,.Eastem India. Proc. Natn.lnst. Sci. India, (B) 28 (4) : 281-406.
Roonwal, M. L. &Chhotani, o. B. 1989. The Fauna ofIndia and the adjacent countries, Isoptera (Termites), Vol. 1 : viii + 672 pp., 315 figs. Zoological Survey of India, Calcutta.
Roonwal, M. L. &Sen Sarma, P. K. 1960. Contributions to the systematics of Oriental Termites ICAR Ent. Monogr. No.1: xiv + 407 pp. Snyder, T. E. 1949. Catalogue of the Termites (Isoptera) of the World. Smiths. Misc. Coli., 112 : 490 pp.
Sen-Sarma, P. K., 'a a1. 1975. Wood destroying termites 0/ India (Final Tech. Rep. PL 480 Project A 7 -FS-S8) ; 198 pp.
