Oligochaeta: India
Contents |
Oligochaeta: India
Introduction
The evolution of the Oligochaeta is obscured due to paucity of fossil records. Nevertheless, their origin has been inferred from studies on the distribution, ecology and comparative anatomy of extant species. On the basis of fossil tracks, Valentine (1980) suggested that vormiform coelomates originated about 700 million years ago. The Annelida were possibly well established at at the junction of the Precambrian and Cambrian periods (570 million years). The differentiation of freshwater oligochaetes from sea annelids probably took place in the early Palaeozoic. Sims (1980) assumed that ancestors of present day terrestrial fonns (earthworms) were established in the undivided palaeocontinent of Pangaea which• was formed at the end of the Palaeozoic and the beginning of the Mesozoic.
Oligochaetes are cylindrical, bilaterally symmetrical coelomate worms with internal and external metameric segmentation. They lack any appendages and suckers but posses a few hook¬like chaetae embedded in the skin with which they gain hold on the substratum. Hence the name Oligochaeta (oligo. few; chaetae. bristles). They are hermaphrodites and•ova are fertilized in cocoons seCreted by a clitell um. Development is direct, without a free larval stage in the life cycle. On the basis of size and habitat, oligochaetes are often distinguished into two convenient groups: MicTodrili (small mainly aquatic worms including the terrestrial family Enchytraeidae) and Megadrili (larger, mostly terrestrial worms and their aquatic representatives).
Oligochaetes have close affinities with leeches and to indicate this relationship Michaelsen (1928) combined the orders Oligochaeta and Hirudinea into Class Clitellata. Both the Oligochaeta and Hirudinea have been lately classified as distinct classes of the super-class Clitellata of the phylum Annelida, which also comprises the class Polychaeta. On the basis of phyletic relationships, Sawyer (1986) has proposed to include the Clitellate as a subphylum of the phylum Uniramia; other subphyla being Onchophora (Class Onychophora), Myriapoda (Class Chilopoda and Class Diploda) and Hcxapoda (insects; Class Apterygota and Class Pterygota). The subphylum Clitellata included two classes: Oligochaeta and Hirudinea; the polychaetes with certain traits formed a distinct group from Uniramia. However, this report follows the traditional classification of the segmented worms: phylum Annelida, comprising a super-class Clitellata (Class Oligochaeta and Class Hirudinea) and the class Polychaeta. The latest classification of Oligochaeta as proposed by Brinkhurst and Jamieson (1971) and modified by Sims (1980, 1982) is given in the following table.
More than 4200 species of oligochaetes are known in the world. Of these, 280 are Microdrili and the remaining about 3920 belong to Megadrili (earthworms). The number of genera have been estimated between 75 and 80 for Microdrili, and between 235 and 240 for Megadrili.
Oligochaetes are found in all types of aquatic and terrestrial habitats. Aquatic Oligochaeta comprise marine, brackish and fresh-water species. The terrestrial forms (earthworms) cannot tolerate dry conclitions and their burrows may be as deep as 3 metres (e.g. Drawida grandis) to reach moisture. They are mostly nocturnal and emerge from their burrows at nightfall. The process of soil formation leads to horizontal layers and earthworms inhabiting different layers are usually categorized into three groups: (i) surface of litter dwellers (epiges), (ii) topsoil inhabitants (endoges) which arc less pigmented and also less active, and (iii) subsoil dwellers (aneciques) which are unpigmented or light pigmented and slow-moving. Some worms are arboreal and inhabit accumulated detritus in the leafaxils of banana, palm, bamboo trees, etc. Organic materials like compost, manure, forest litter and humus, municipal dumps, soils wetted with effluents and J. M. Julka, Zoological Survey of India, Solan kitchen drainage are highly attractive to some species.Some live under snow and can tolerate extreme cold condition on high mountains. Latest classification of Ollgochaeta Class Order/suborder Superfamily Oligochaeta Lwnbriculida Moniligastraida Haplotaxida Haplotaxina Tubificina : Alluroiina Lumbricina : Eilchytraeoidea Tubificoidea Biwadriloidea Criodriloidea Lumbricoidea Glossoscolecoidea Megascolecoidea
Earthworms are known to be friends of farmers. They enhance soil fertility by changing its pbysico-cbemical properties. They construct extensive bUI1'OWS and deposit the ingested material as swface or subswface casts. In this way, large quantities of soil from deeper layers are brought to the swface and decaying organic matter is taken down into soil. Soils with worms remain loose and have a greater capacity to retain air and water. Earthwonns enhance microbial activity by breaking up organic matter during humidification. A large number of worms die during unfavourable period when chemical demand in soil is maximum because of growing vegetation. Microbial decomposition of dead worms releases considerable amount of nitrogen and other nutrients (25% nitrogen in the form of nitrate, 48% ammonia, 3% soluble organic compounds, 27% unaccounted). Addition of oligochaetes in sewage sludge and sludge amended soils hastens in sludge decomposition and stimulates metabolic activity of bacterial population therein. They are also important in redistributing certain toxic heavy metals in polluted waters and soils by their burrowing, feeding and excretory activities. Because of their feeding habits, earthworms have been utilized for the production of vermicompost through venniculture techniques. Earthworm tissue is rich in proteins • and with suitable processing it could augment or even supplant traditional feeds for livestock and aquaculture. Maoris in New Zealan<J and the natives of New Guinea considers wonns ~delicacy.
Earthworms have been used in folk-medicine in treating certain diseases. Some species of aquatic oligochaetes are good bioindicators of polluted waters.
Earthworms have also been reported to cause damage to crops. They seize leaves of growing plants and pull them into burrows. often killing the plants. Intensive burrowing activity of wonns sometimes retard germination, growth and root development of vegetable crops. They also act as essential intermediate hosts to protozoan, cestode and nematode parasites of birds and mammals. There are repo~ of worms causing soil erosion on hill slopes by bringing fme soil to the surface. Castings of some species on drying become cement-like hard clods,which affect normal percolation in soil and productivity of vegetable crops, especially root crops like carrot, turnip, radish, etc.
Historical Resume
Carlos Linnaeus (1757) listed 2 annelid species in the 10th edition of Systema Naturae: an oligochaete (Lumbricus te"esris) and a polychaete (Lumbricus marinus). Otto Friderich Muller and Jean-Baptiste Pierre Antoine Leclercque de Monel Lamarck recognised separate species of aquatic oligochaetes (microdriles) between 1774 and 1816. Diversity in earthworms (megadriles) was brought to light by Savingny in 1826, when he described 20 lumbricid species from the Paris region in France. In the Indian subcontinent, these organisms attracted the attention of individ~al European naturalists during the fifth decade of nineteenth century. Collections were made from different areas and preserved for future studies.
In the meanwhile, Charles Darwin's (1881) work on the formation of vegetable mould through the action of worms had generated global interest on oligochaete studies. Thus individual efforts began much before the establishment of organised institutions like the Zoological Survey of India.
i.) Pre-1900
The significance of earthworms has been recognized in this country since ancient times as preparations made from them were used in the Unani System of Medicine for treating certain diseases.Modem scientific studies on oligochaetes in our subcontinent were initiated sometime during the later half of the last century. Robert Templeton, a British Zoologist, was perhaps the fust one to undertake systematic collection and study our oligochaete fauna, when he discovered Megascolex caeruleus in 1844 from Sri Lanka. Schmarda (1861) and Vaillant (1867) followed Templeton in describing new species of both aquatic and terrestrial worms from the island.
Efforts of Edmond Perrier (1872) resulted in the description of Moniligastor deshayes gen. et sp. nov. from Kerala on the Indian mainland. But it was only during the last two decades of the nineteenth century that unexplored fauna of our subcontinent attracted serious attention of famous European Oligochaetologists: Beddard (1882-1905), Bourne (1886-1894), Benham (1893), Michaelsen (1897¬1936), Ude (1893) and Fedarb (1898).
Frank Evers Beddard,an expert oligochaete taxonomist from England, characterised several species from the 'British India' between 1883 and 1893. He consolidated the then existing knowledge on world oligochaetes (including India) in his 'Monograph on Oligochaeta' which was published in 1895 and fonned basic work on the-group until the end of last century. Beddard (1902) revised the genus Amyntas (Amynthas) and for the first time surveyed earthworms of the Maldive and Laccadive Islands in 1903. He (1905) discovered a tiny enchytraeid oligochaete, Henlea lefroyi, which destroyed the eggs of locust Acridium sp. Besides taxonomic studies, Beddard (1882) published an account on the anatomy and histology of an Indian earthworm.
Base on a few characters, Alfred Gibbs Bourne (1886) named certain species from the Nilgiri and Sheveroy Hills in southern India. He (1894) brought out slightly detailed descriptions of oligochaetes but these were considered inadequate. Bourne's (1889) other taxonomic contribution was a short paper on earthworms of the western Himalayas and Dehra Dun. IBourne was also interested in anatomical studies and examined the circulatory system in earthwonns (1891).
Rosa, an Italian Zoologist who had accomplished notaple work on European oligochaetes, concentrated his investigations (1888 and 1890) on Leonardo Fea' s collection of Burmese earthworms in the Muso civico di Storia Naturale Giacomo Doria,Genova, Italy. A new species Megascolex templetonianus was revealed by him from Sri Lanka in 1892. Contributions of Benham (1893), Ude (1893) and Fedarb (1898a, 1898b) were limited to the discovery of a few new species from various parts of the country.
The knowledge on oligochaetes of India was of a somewhat sporadic nature throughout this period. Oligochaete species were mainly distinguished by external characteristics with little stress on their anatomy. Wilhelm Mir.haelsen, a German Oligochaetologist and whose work circled the globe, started working on Indian oligochaetes towards the end of last century. His rust major contribution (1897) was the publication of a comprehensive account on earthwonns of Sri Lanka. Species were better characterized with emphasis on both external and internal diagnostic features.
ii) 1901 to 1947
This period is considered as glorious one in the historY of Indian oligochaete taxonomy. It is marked by investigations of world-renowned workers like Wilhelm Michaelsen, LL Col John Stepbenson and Gordon Enoch Gates. These luminaries later on fonnulated basic classifications of Oligochaeta of the world. Another important development was establishment of the Zoological Survey of India in 1916, which led to organized faunistic survey in the country.
Michaelsen's (1900) monograph on Oligochaeta was published in Das Tierreich, which dealt with descriptions of all• known species of the world. This work, though out-dated, is still considered as a valuable reference guide for oligochaete taxonomy. Initially,.Michaelsen (1903, 1904) directed his studies on oligochaetes of Sri Lanka. Gradually, he extended his research activities to other parts in the subcontinent. Michaelsen (1907-1910) reported several new oligochaete taxa primarily based on the material collected by the staff of the Indian Museum and lent by Dr. Nelson Annandale, the Superintendent of the museum at that time. Most of the 'types' of Michaelsen's species were deposited in Zoologisches Institute and Zoologisches Museum, Universitat Hamburg, Germany but some specimens of the 'type series' had also been registered in the collection of the Indian Museum. Michaelsen (1909) published a consolidated account on the Oligochaeta of India and adjacent countries in the Memoirs of the Indian Museum.
This work formed the basis for future taxonomic swdies on Indian oligchaetes, as it included keys for identification and descriptions of all known species in our subcontinent till that date. From 1910 till his death in 1937, Michaelsen contributed only 2 notable publications on our fauna, since he concentrated his researches on the Ethiopian and Neotropical fauna dwing this period. He described earthworms, including new species, of Travancore and Coorg regions in 1913 and 1920 respectively.
Thomas Nelson Annandale (1905-1906), the then Superintendent of the Indian Museum, described two species of aquatic microdrile worms of the Naididae as commensal on freshwater sponges, snails and bryozoans.
Lt. Col. John Stephenson of Indian Medical service (IMS) and Professor of Zoology at Govemment College,Lahore dominated oligochaetological studies for about two decades from 1910 to 1930. He began his initial investigations on aquatic oligochaetes of undivided Punjab (1902¬1910) and Travancore (1910). Stephenson (1911-1922) recognized several new taxa ofboth aquatic and terrestrial oligochaetes from various parts of the country on the material collected by ~estaff of the Indian Museum, Bombay Natural History Society and himself.Dr. Baini Prashad, a student of Stephenson, also carried out extensive collecting for the latter's studies. Stephenson (1923) published a monograph on the Oligochaeta of this region under 'The Fauna of British India (including adjacent countries), series.
Besides taxonomic descriptions, this monograph discussed phylogenetic relationships of Indian oligochaetes for the rust time. Stephenson (1924) studied cavemicolous oligochaetes collected by Dr. S.W. Kemp from the Siju cave in the Garo Hills, Meghalaya. Between 1924 and 1931, Stephenson contributed a fe.w publications on these organisms in our subcontinent. Majority of Stephenson's papers were published in the Memoirs and Records of the Indian Museum. The 'Types' of Stephenson's species were deposited in the BritishMuseum (Natural History) and the Indian Museum; those in the latter now form a part of the National Zoological Collections in the Zoological Survey of India. The most outstanding work of Stephenson was the publication of another monograph 'The Oligocha"eta' in 1930, which is stin an indispensable source for morphological, ecological, biological and taxonomic details of oligochaetes of the world. Luigi Mardis Cognetti (1911) recognized new species of Megascolex. Plutellus and Drawida from Kerdla.
Much interest was also generated to work out the morphology of Indian earthworms in 1920's. Dr. Baini Prashad dealt with the anatomy of Pherelima posthuma at Government College, Lahore under ~e supervision of Stephenson. Studies were conducted on the lymph, alimentary and. calciferous glands by Thapar (1918, 1932), and Stephenson and Prashad (1919). Balll concentrated on nephridial studies, while Stephenson and Haru Ram (1919) investigated the morphology and development of prostate glands in the Megascolecidae. Prof. Karm Nanrln Bahl (1924) of Lucknow University in his presidential address to the Zoology Section of the Indian Science Congress at Bangalore stressed the need for preparing m~oirson Indian animal types.
The onus of writing the first memoir on Pheretima fell on Bahl himself. This memoir, published in 1926, is still considered an excellent work on detailed morphology of Metaphire (=Pheretima) posthuma. Twenty four years after its first appearance, Bahl's monograph has passed into revised second (1930), third (1943) and fourth (1950) editions, which indicates its popularity as a guide to teachers and students of Indian zoology. Bahl (1922-1947) worked out detailed morpholgy, development and physiology of excretory system in a number of Indian worms. His contributions on oligochaete nephridia formed in part basis of Jamieson's (1971) latest classification of earthworms of the world. Mode of reproduction in aquatic oligochaetes was explained by Mehra (1920, 1927).
Rao (1920-1927) gave an account of anatomy of Drawida species and history of Lampito mauritii. and discovered new taxa of Glyphidrilus from Karnataka in south India. Aiyer (1925¬1929) studied taxonomy of aquatic and terrestrial Oligochaeta of Tmvancore.
Gordon Enoch Gates, an American Zoologist, joined Judson College at Rangoon in Burma in 1924 at a time when Stephenson's Fauna ofBritish India (Oligochaea) had just appeared. Gates was fascinated by the existence of great diversity among the Burmese earthwprms,and took up studies on oligochaete taxonomy. He dominated in this field for about 55 years (1925-1982) not only in our subcontinent but also allover the world. Between 1925 and 1943, he published a series of papers on earthworms of Burma and recognised new anatomical characteristics of taxonomic importance. The. Second World War not only resulted in the destruction of G~tes'collections, library, most of his records and manuscripts but also termination of earthworm collecting in Burma. He continued to publish papers on the Burmese earthwonns from 1952 to 1962 based on his notes and material that was rescued during the war. Gates' work on earthworms of Burma culminated in the publication of a comprehensive monograph in 1972, which also included descriptions of species from northeast India, and Andaman and Nicobar Islands.
Gates (1934-1940) undertook revisionary studies on• Indian genera mainly based on material borrowed from the Zoological Survey of India and British Museum. These studies revealed several new genera: Scolioscolides. Nellogaster. Barogaster, Lennogaster. Rillogaster. Pellogaster, Priodochaeta. Priodoscolex and Travoscolides. Because of second world war, Gates shifted his activity Bunna to India at Aijahabad between 1943 and 1-951. He thoroughly surveyed earthwonns of Allahabad sector in the Gangetic plains,parts of central India, Debra D\1n and some northern hill stations.He discovered new genera, Bhalia. Calebiella and Deccania during these explorations. Gates (1941-1945) also worked out earthworms of Sri Lanka. Extens'ive taxonomic studies on oligochaertes in the Indian subcontinent led Gates to formulate a.new classification of earthworms' based on stable somatic characters.
Cemosvitov (1937) identified a small collection of Indian Lumbricidae. Anatomy of Glyphidrilus annandalei was described by Nair (1938-1939). Saksena (1943) detennined new types of muscle fibres in Pheretima posthuma. Histochemical studies on ova of this species were undertaken by Nath and Bhatja (1944). Enteronephric nephridia and physiology of excretion attracted the attention of Vati (1945, 1947) and Hora (1946) respectively.
iii) 1948-1990
Taxonomic studies Indian earthworms suffered a set back between 1947 and 1970. Gates had left this subcontinent to extend his research activities on north American worms at the University of Maine in U.S.A. But based on his earlier studies, he could revise Indian Lumbricidae in 1958 and Octochaetidae in 1962. Investigations in the Zoological Survey of India were neglected in the beginning of this period in the absence of an oligochaetologist. Scientists and staff of the survey were occupied in rearranging collections which had suffered dwing the Varuna floods at Varanasi in 1943. Meanwhile, centres of advanced research in different zoological disciplines had been established in the country at various institutions and universities, and oligochaetes being convenient laboratory organisms became targets of investigations.
Jatinder Mohan Julka renewed systematic studies on earthwonns in the Zoological Survey of India in 1965. His interest on these organisms was sustained due to Gates' guidance through correspondence. To begin with, Julka and his colleagues published notes on the regional distribution of oligochaetes and their occurrence in various parts of the country (1967-1977, with Halder; 1970-1972, with Soota). Julka participated in two multidisciplinary scientific surveys expeditions to unexplored areas of Arunachal Pradesh: Daphabum region of Lohit district (1969-70) and Subansiri district (1974-75). Several new taxa were recognized from the earlier material (1975) and also from material collected during these expeditions (1976 and 1981).
Julka undertook investigations on earthworms of Orissa (1976 and 1978; 1987 the last one with Senapati). For the flJ'St time from India, he recorded two peregrine species: Lumbricus castaneus from Himachal Pradesh in 1979 and Nematogenia panamaensis from Kerala in 1990. (in collaboration with Paliwal). Julka (1982) published a comprehensive account on earthworms of Andaman and Nicobar IslandS. He bro~ght to light two new monotypic genera from Karnataka, and new genus•with 3 new species from Meg~alaya between 1982 and 1988. Earthworm resources of India and their distribution were assessed by Julka and Paliwal (1986). Stephenson's Fauna of British India on Oligochaeta became obsolete• with the recognition of new taxonomic characteristics and taxa. Julka took up revision of Indian earthworms and to begin with he produced a monograph on the family Octochaetidae (1988) under the 'Fauna ofIndia' series. This work dealt with descriptions of 128 species, including 6 new genera and 16 new species. Affmities of Peninsular worms were traced with the Malayan fauna by Jayaram (1949). Rao (1969) recorded a marine species of enchytraeids among interstili:alfauna inhabiting beech sands of Orissa. Notes on earthwo~s of Debra Dun and the Himalayas were made by Soota (1970), and Soota and Halder (1977-1981) respectively.
Besides taxonomic studies, investigations on other aspects of Oligochaetes have been carried out in the Zoological Survey of India. Mukherjee (1969) found water birds feeding upon aquatic and terrestrial worms. Infestations of protozoan parasites in earthworms has been the subject of study by Biswas and Mukherjee (1974), Mandal and Nair (1975-1976), Mukherjee (1980), and Mukherjee and Chakraborty (1975-1987). At the High Altitude Zoology Field Station of Z.S.I. at Solan, Julka and his colleagues have been contributing substantially to the knowledge on their ecology and biology since 1984: seasonal activity and population dynamics (with Mukherjee and Paliwal), mass migration (with Chandra and Mukherjee) and effect on C/N ratio of soil (with Mukherjee). Mukherjee and Julka (1981-1984) recorded soil Protozoa in the intestine of earthwonns.
Role of earthwonns in soil fertility attracted the attention of Shrilchande and Pathak (1948) at Bombay University, and Dubash and Ganti (1963) at Institute of Science, Bombay. Dubash and Tembe (1959) published methods of their culture. They in collaboration with Ganti (1961) brought out a comprehensive review on India earthwonns. Following research at Bombay, serious studies on these organisms were undertaken at other institutions/universities in Maharashtra. Functional morphology of calciferous glands was studied by Kashyap and Ranade (1952), and Joshi and Kelker (1953) at Pune University. Histochemical investigations were undertaken by Kamat (1955-1962) at Maharashtra Association for Cultivation of Science Research Institute at Pune and by Varute (1970-1972) at Shiva ji University, Kolhapur. Hanumante and Nagabushanam (1977-1979) of Marathawada University at Aurangabad contributed significantly on neurosecretion, osmotic behaviour and regeneration in Perionyx excavatus. Srinivasulu (1986) investigated association of mycoflora with the digestive tract of an earthwonn.
The work of Nijhavan and Kanwar (1952) on physicochemicai properties of wonn castings at Punjab Agriculture University, Ludhiana is still useful in understanding their effect on soil productivity. Malhotra (1957) and Nath et at. (1958) of Punjab University carried out histochemical studies in Pheretima posthuma. Menon made significant contributions on various, systems of Eutyphoeus (1968, with Sareen and Kaur; 1969, with Singal; 1969, with Singal and, Sharma; 1969, with Kaur and Singal; 1973, with Sareen •and• Mittal). For the first time, cytological studies were initiated on Indian oligochaetes by Handa(1969-1971;1976, with Shanna and Sohi). Sareen and his students (1970-1976) continued cytological observations on their reproductive organs.
Vasisht (1977-1981) concentrated his studies on digestive system in north Indian aquatic oligochaetes. Neurohisoology in oligochaetes was studied by Satija and Carg in 1973 and 1976. Some information on worms of neighboring areas was available from the works of Das tt ale (1964) and Sharma and Kaul (1974) in Jammu and Kashmir, and Kapur and Kapil (1986) in Haryana. Sharma and Madan (1983) of Indian Institute of Technology, New Delhi published an excellent review of worms in relation to soil health and pollution control.
Khambata and Bhat (1953-1957) initiated researches on intestinal microflora of earthworms at Kasturba Medical College at Manipal in Karnataka. Bhat and his students (1974-1975) further developed suitable techniques to ascertain their dietary and migratory habits.Saroja (1959-1964) and Rao (1962-1963) of Kamataka University devoted their energies to examine physiology of oxygen consumption and low temperature acclimation in earthworms. Under the supervision of Rao, Kale (1972-1976) continued studies on effect of thennal acclimation on oligochaete metabolism.
Ecological studies on earthworms were undertaken at'the University of Agricultural Sciences, Bangalore by Krishnamoorthy, Kale and Bano betWeen 1977 and 1979. Kale and Krishnamoorthy (1978, 1981) studied their distributional pattern in different habitats and their influence on soil fertility. Krishnamoorthy (1976-1989) carried out investigations on behaviour and comparative ecology of earthworms in grassland and woodland sites. Kale and Bano (1986) concentrated their researches on vermicomposting, which resulted in developing organic fertilizers from worm casts (vermicompost named as Vee Compo E•. 83 U.A.S.) and worm tissue as source of protein in the food of poultry and aquaculture.
K. Vanamala Naidu (1961-1967) of Government Arts and Science College, Chittoor in Andhm Pradesh undertook taxonomic studies on aquatic oligochaetes which had remained neglected since the publication of Stephenson's Fauna of British India in 1923. He revised the Aelosomatidae, Naididae and Tubificidae. His painstakingly prepared checklist of freshwater Oligochaeta of the Indian subcontinent and Tibet is still an important reference work.
Physiological studies on earthworms attracted the attention of Tandan (1951) and Gupta (1976¬1979) at Lucknow University. Agarwal and Shanna (1977) contributed towards the knowledge on neurosecretory cells of cerebral ganglion of P heretima posthuma at Debra Dun. Taxonomy and ecology of the Enchytraeidae were studied by Lal (1981) at Benaras Hindu University. Mukherjee (i986, with Janardan Singh) carried out a comparison of different quadrat sizes.and extraction methods for estimating earthworm populations. Use of earthworms as a fish feed was advocated by Srivastawa in 1986. Histochemical studies on prostate glands and gizzard were undertaken by Sharan (1971), and Vinayak and Prashad (1975-1978) at Patna University.
Since 1976, Subbarao and his students have been pursuing ecological and physiological studies on a littoral oligochaete at Andhra University. A new protozoan ciliate from brackish water oligochaete was described by Joseph and Hanumantharao (1980). Effects of fue on nutrients of worm casts and fertilizers on population density attracted the attention of Vikram Reddy (1983¬1987) at Kakatiya University. Prabhoo (1961.. 1964) revealed the existence of 5 new enchytraeids from Kerala soils. Taxonomic stud'ies on these worms are still neglected in our country as compared to other regions of the world.
Protein constituents of cuticle in earthworms were analyzed by Rajulu and his colleague (1968¬1973) at Madras University. Arunachalam and his co-workers (1978-1988) carried out investigations on the biology, ecology and histochemistry of a peregrine species, Pontoscolex corethrurus at Madurai University. Bioluminescence in Lampito mauritii was observed by Kaleemurrahman (1981) at National College, Madras. Distribution and population structure of earthworms were surveyed by Ismail (1985, with Murthy; 1990, with Ramakrishnan and Azar). Growth as a factor in harvesting worms was studied by Ismail and Alawdeen in 1986. Other oligochaete researchers in Tamil Nadu during this period were: Radha (1982, taxonomy and bioecology of enchytraeids). Rajaram et ale (1986, ecology of aquatic oligochaetes) and Vardaraj (1986, effect of insecticides on earthworm metabolism).
Physiological and ecological studies on central Indian earthworms were undertaken by As~ (1968 &1977) and Chauhan (1978-1980) respectively. Dev and Vyas (1972) analysed alkaline phosphates in the alimentary canal and specmatbecae of Barogasler annandalei.
Prof. Madhab Chandra Dash initiated researches on oligochaete ecology at Berhampur University in Orissa. His investigations included estimation of worm' populations by chemical extraction methods in 1973 (with Patra), ecology and taxonomy of the Enchytraeidae between 1973 and 1978 (with Thambi and Nanda), comparison of primary plant production with secondary production of oligochaetes in 1974 (with Patra and Thambi) and population of oligochaetes in 1974 (with Patra and Thambi) and population dynamics and energy budget of Lampito mawiti; in 1973-1977 (with Patra). Subsequently, Dash shifted his activites to Sambalpur University where he developed an excellent centre of oligochaete research. Initially, studies were undertaken on various aspects:
Histochemical changes during developmental stage of Lampito mauritii (1977, with Senapati, Hota and Guru), morphology and production of cocoons and emergence pattern (1979-1980, with Senapati), fungal and nematode feeding (1979-1980, with Behem, Mishra, Nanda and Senapati), respiration (1979-1984, with Mishra,and process of decomposition and soil turnover (1980-1986, with Senapati,Rana, Panda, H.K. Dash and Behura). Different types of digestive enzymes in the Enchytraeidae were studied by Dash and his students in 1981. Dash and his co¬workers (Senapati, H.K. Dash, Patra and Nanda in particular) also contributed substantially to the knowledge on wormcast production, population dynamics and energetics in different habital$ between 1979 and 1983. Dash (1983) published a monograph on the biology of Enchyttaeidae.
Bikram Keseri Senapati and his students have conducted research on population biology,reproductive strategy and secondary production of earthworms. Another aspect of Senapati's investigations has been effect of insecticides and organic matter accumulation on earthworm activity between 1986 and 1987 (with Pani,Sahu and Mishra). Studies of Senapati and Dash (1982, 1986) on earthworms in relation to decompositipn and waste utilization would be useful in developing vennitechnology in the country. Mishr~ another student of Dash, has been concenU'ating studies on effects of insecticides and starvation on earthworm metabolism since 1986.
Reddy (1978, with Alfred), and Ramakrishnan (1988, with Mishra; 1989, with Bhaduria) undertook investigations on population dynamics of earthwonns in different habitats at Northeast Hill University, Shillong.
Studies from Different Environs
Aquatic oligochaetes usually do not have any habitat preference in relation to physico-chemical parameters of environment or specific plant association or any other readily recognizable ecological criteria (Brinkhilrst and Jamieson, 1971). Studies on aquatic wonns of different environs (family Tubificidae) in our region are very few most of these deal with the taxonomy of littoral fonns. Researches on benthic species have remained more or less neglected. Stephenson (1903-1920) recorded several species from small fresh water bodies like ponds, tanks, pools, ditches, etc. from northwest and central India, Western Ghats and Sri Lanka, primarily based on the material collected by Annandale of the Indian Museum. Oligochaetes of a tank within the Indian Museum compound at Calcutta were extensively explored by Annandale '(1905-1906), Michaelsen (1909) and Stephenson (1911-1912).
The other notable contributions on worms of these habitats were those of Aiyer (1929) in Kcrala and Naidu (1961-1963) in Andhra Pradesh respectively. The lacustrine fauna attracted the -attention of Oligochactologists only on a few occasions: Bhimtal and Nainital lakes (Michaelsen, 1909; Stephenson, 1916), Villayami lake in Kerala (Aiyer, 1929), Lot Tak lake in Manipur (Stephension, 1907) and Inle lake in Bwma (Stephenson, 1918).
Among freshwater streams, the Bagga stream at Cuddapah in Andhra Pradesh was thoroughly explored f~r oligochaeotological studies by NaiQu (1961-1963). Sporadic records of one or two species had been made from other habitats: sewage canal (Naidu 1961), Ravi, Gaumati and Handri rivers (Stephenson, 1914, 1920, Naidu, 1961) and brackish water Chilka lake (Stephenson, 1917, 1921). Stephenson (1923) compiled a list of several aquatic oligochaetes as commensals on sponges, snails and bryozpans of standing waters.
Another group of Microdrile oligochaetes, the Enchytraeidae (pot worms), occurs in terrestrial littoral and marine habitats. They are abundant in acidic soils with high organic matter. The work on Indian Enchytraeidae is limited to a few research papers. Dash (1978-1980, with Thambi and Nanda) carried out studies on the taxonomy and ecology of these worms in grasslands and deciduous forests of Orissa. Prabhoo (1961-1964) and Radha (1982, unpublished) studied their systematics in South Indian soils. Ecological investigations on pot worms of a tropical grassland were undertaken by Lal (1981, unpublished). A salt-tolerant species, Stephensoniella (=Enchutraeus) barkudensis had been recorded from the bracksih water Chilka lake and Ennor estuary. A couple of species were encountered among interstitial fauna on sea beaches by Aiyer (1929) and Rao (1969).
Most of Indian Oligochaete publications are on earthworms. Certain areas of the country have been surveyed extensively. Evergreen forests. with considerable rainfall and large amount of organic mauer in the soil in eastern Himalayas harbour rich earthworm fauna. Several new species were described from these forests in Meghalaya (Stephenson, 1920 Julka, 1988), Arunachal Pradesh Stephenson, 1914; Julka, 1976, 1981) and Darjeeling district of West Bengal (Michaelsen, 1909; Stephenson,1920, Julka, 1975). Mass migration of worms in hill soils of Nagaland and Himalchal Pradesh was observed by Reddy (1980) and Julka et ale (1984) respectively. Studies on earthwonns had been conducted on different types of forests: subtropical pine forest in Meghalaya (Reddy and Alfred, 1977), mixed broad-leaf and pine forest in Himachal Pradesh (Julka and Mukherjee, 1984), subtropical woodland in Orissa (Mishra and Dash, 1984) and peninsular woodlands in Karnataka (Krishnamoorthy, 1985, 1988). Aiyer (1929) and Julka (1986, with Chandra) examined earthworms of tropical rain forests in Kerala. New megadrile taxa were revealed from the deciduQus forests and pastures in the western Ghats by Stephenson (1920), Gates (1945) and Julka (1982, with Rao; 1983). Earthworms of eastern Ghat soils were studied by Senapati and Dash (1982), while whose of broad-leaf subtropical forests in Doon Valley were surveyed by Gates (1945) and Soota (1970).
Earthwonn fauna of pasture, arable and deciduous forest soils were explored by Julka (1975-1978), and Julka and Senapati (1987). Anthropochorous lumbricids from high altitudes of the Labaul valley and other parts of Himachal Pradesh were reported by Julka (1979, 1981) and Soota and Halder (1980).
Earthworms of aluvial soils in the Gangetic plains attracted the attention of Dates (1945-1947). Comprehensive accounts on northern peninsular soils were published by Gates in 1945 and 1956. Stephenson (1923), Gates (1960) and Julka (1988) studied the megadrile fauna of laterite soils in Maharashtra.
Investigations on the activities, biomass and energy budget of earthworms in pastures were undertaken by Roy (1957) and Desh et ale (1977-1983). Studies on these organisms in agroecosystems were carried out by Senapati el ale (1986-1988), and Reddy and Goudnarayan (1987). Recently, Julka and Paliwal (1990) have investigated seasonal changes in the population of worms in an orchard. Earthworms of shifting cultivation (Jhum) in northeast India were studied by Mishra and Ramakrishnan (1988) and Bhaduria and Ramakrishnan (1989). Stephenson (1924) described cavemicolous oligochaetes from the Siju cave in the Garo Hills in Meghalaya.
Oligochaetological studies on peninsular soils were carried out to some extent by Michaelsen (1909), Gates (1945), Kale and Krishnamoorthy (1978), and Julka (1988). Earthworms of acidic soils in Palni Hills attracted the attention of Michaelsen (1909), Jamieson (1977) and Julka
(1988). Several species from the insular ecosystem of Andaman and Nicobar Islands have also been reported (Julka, 1982). Bionomics and salinity tolerance of a littoral oligochaete on Andhra cOast were investigated by Subbarao and Ganapati (1972-1978). Earthworms of diverse ecosystems were thoroughly explored by Gales in a series of papers published between 1925 and 1972. He also studied worms from the plains and hills of Sri Lanka in 1941 and 1945.
Estimation of Taxa
Oligochaete fauna in our region is represented by two orders: Moniligastrida and Haplotaxida. Out of twenty seven recognized families, fourteen families comprising 91 genera and 585 species occur in the Indian subcontinent. The National Zoological Collections in the Zoological Survey of India comprise about 362 species belonging to 64 genera and 11 families. The family-wise break-up of oligochaetes in the Indian Subcontinent is as follows:
These comprise aquatic forms and tiny terresttial pot worms of the Enchytraeidae belonging to suborder Tubificina of Haplotaxida. Microdriles in our subcontinent are represented bytbree families of aquatic oligochaetes namely Naididae,Tubificidae and Phraeodrilidae, and terrestrial pot worms of the Enchytraeidae. An approximate estimate of different categories of microdriles in the Indian subcontinent and world (figures in parenthesis) is given below:
Although aquatic oligochaetes of southern hemisphere are poorly known,there appears to be a greater diversity in northern hemisphere. Most Indian species of aquatic oligochaetes are cosmopolitan. Among these are: naidid species of Chaetogasler, Nais, Slavina, Stylaria, Haemonais. Dero and Pristina, and tubificid species of TUbifex. Limnodrilus and Aulodrilus. However, a few species of the Naididae (Chaetogaster limnaei gengaiensis, Dero indica, D. plumosa. Aulophorus humanae and A. indieus) seem to be endemic and show close affinities with the fauna of southern hemisphere. Distributional range of certain species like Allonais i~qualis. A. peelinata. A. gwaliorensis and A. paraquayensis extends to Africa, south America and other parts of Asia. Cultures of Limnodrilus ho//meisleri, a cosmopolitan tubificid, are commonly used as feed for aquarium fishes in several parts of the country, especially in West Bengal and Orissa.
The Phraeodrilidae is restricted to southern hemisphere. Only one species, P hraeodrilus z~ylanicus. has been recorded from Sri Lanka in this subcontinent. Studies on tropical Enchytraeidae are fairly sparse. Until 1961, twelve species belonging to five genera, Enchytraeus. Achaeta, Fridericia, Stephensoniella and /lemienchytraeus, were known from Indian soils. With.the record of three more genera, Marionina. Propappus and Hemijridericia. a few more species were added to our knowledge of enchytraeids.As a result, a total of 21 species are known from India as compared to about 500 in the world. Almost all Indian enchytraeid genera have world-wide distribution, but with endemic species in this region. The true picture of endemicity of Indian species could be inferred only by further extensive exploration of enchytaeids in the subcontinent
Like elsewhere, earthworms form bulk of oligochaete fauna in India and adjacent countries. They belong to order Moniligastrida and suborder Tubificina of Haplotaxida. They are represented by 508 species and 67 genera in the Indian subcontinent as compared to about 3320 species and 240 genera in the world. Details of different taxa in our region are as follows (figures in parenthesis denote world fauna) :
Moniligastridae 4(5) 97(113) Criodrilidae 1(1) 1(1) Lumbricidae 8(21) 16(302) Glossoscolecidae 1(25) 1(200) Almidae 1(5) 4(40) Ocnerodrilidae 8(21) 16(108) Acanthodrilidae 3(27) 34(482) Octochactidae 26(45) 145(430) Mcgascolccidae 14(25) 195(1000) Eudrilidae 1(45) 1(500) 67(220)* 508(3176)* • Excluding 20 genera and 144 species belonging to those families not found in the Indian region.
Majority of megadriles are endemic in our subcontinent. A few peregrine species have been introduced presumably in soil around roots of exotic plants. Generic distribution of peregrine species among 8 families is : Lumbricidae (8), Ocnerodrilidae (4), Megascolecidae (2), Acanthodrilidae (2), Eudrilidae (I), Clossoscolecidae (1), Criodrilidae (1) and Octochaetidae (1).
Among the Moniligaslridae, drawida contains the maximum number of 79 species with endemicity in South and northeast India and Bwma. Some of the longest Indian worms (over one metre in length) belong to this genus e.g. Drawida nilamburensis and D. grandis. Other genera of this family found in our region are : M oniligaster (8 spp) in southern portion of peninsular India, Desmogaster (8 spp) and Hastirogaster (2 spp) in Burma.
The Ocnerodrilidae comprises two subfamilies: Ocnerodrilinae and Malabariinae. All southeast Asian genera of the Ocnerodrilinae came originally from south America or Africa (Gates, 1972). 1bese are Gordiodrilus, Ocnerodrilus. Euke"ia and Nematogenia, each being represented by one or two circummundane spccies.Monospecific Curgiona is known to occur only in south India. Gates (1972) believed its home range to be in Africa. The Malabariinae genera, Malabaria (4 spp), Animal Resources ofIndia. Deccania (1 sp) and Thatonia (5 spp.), are endemic and probably evolved in the Indian peninsula. Indian acanthodrilids are represented by a peregrine species, each of Microscolex and Pomodrilus. and several endemics of Plutellus (32 spp).
The Octochaetidae has largest number of genera (26) in the Indian region. The number of known 6ctochaetid species is 145 which are numerica~y second to megascolecids in this region. Excepting Dichogaster (represented by 5 anthropochorous species), all Indian octochaetid genera have endemic species. Most of these genera have endemic species. Most of these genera are found only in the indian subcontinent, and can be referred to two distinct groups : north, northeast and Bwmese group of Eutyphoeus (45 spp.) Bahlia (1 sp), Calebiella (1 sp), and Scolioscolides (1 sp), and a second group of peninsular India comprising 22 genera with majority of species belonging to Hoplochaetella, Octochaetona, 'Celeriella, Wahoscolex, Lennogaster and Ramiella.
The Megascolecidae contains 14 genera represented by the largest number of 193 species. The pheretimoid group comprising Amynlhas (33 spp.) and Metaphire (26 spp.) is endemic in Bunna and Andaman and Nicobar Islands; the other group being represented by one or two peregrine species of Pilhemera and PolYpherelima. Tonoscolex (16 spp.), and Nelloscolex (2 spp.) are endemic in Burma and northeast India. Recently, another endemic genus Kanchuria with 4 species has been discovered from the Garo and Khasi Hills in Meghalaya.Perionyx with 53 species has discontinuous distribution in western and eastern Himalayas, Burma and peninsular India, but majority of species are found in the Darjceling district of West Bengal in the eastern Himalayas. Other megascolecid genera, M egasco/ex (33 spp.), Notoscolex (11 spp.), Lampito (8 spp.), Lannoscolex (1 sp.) and Troyia (1 sp.) belong to the Indian peninsula.
The ethiopian Eudrilidae is represented by one or two records of a circummundane species, Eudrilus eugenae.
Classified Treatment
Order Moniligastrida Family Moiligastridae
Stephenson (1923) and Gates (1934, 1945, 1965, 1972) described several sPecies of Drawida from various parts of the country. Recently, Julka (1976, 1978) revealed three new species of this genus from Arunachal Pradesh. Senapati et ale (1979) studied seasonal dynamics and emergence pattern of D. c~lebi. Effects of insecticides on population and reproductive biology were estimated by Pani and Senapati in 1986. Revised descriptions of all known species of M oniligaster were published by Gates (1940). An up-to-date key for the identification of all moniligastrid genera, and descriptions of moniligastrid spccies from Burma and northeast India can be found in the monographic work of Gates (1972). Order Haplotasida Suborder Tubificina Superfamily Enchytraeoidea Family Enchytraeidae
About 21 genera comprising 500 species are known in the world, but only 1 genera and 21 species are reported from Indian soils. Stephenson (1923) gave descriptions of a few species in the Fauna of British India.Prabhoo (1961, 1964) described 5 new species of Achaeta from Kerala. A few new species of different genera were also discovered by Thambi et ale (1978) and Lal et ale (1981). Ecological studies on enchytraeids of grasslands were carried out by Thambi et ale (1978) and Dash el ale (1978). Dash's studies on enchytraeids culminated in the publication of a comprehensive monograph "Biology of Enchytracidaeu dealing with their morpholgy, taxonomy, biology, physiology and ecology. Superfamily Tubificoidea Family Naididae
Stephenson (1923) consolidated information on naidids known from India and adjacent countries. Aiyer (1929) discovered a new genus Stephensoniella from aquatic habitats in Kerala. He also gave detailed descriptions of certain species of Aulophorus, Dero, Nais, Pristina and Slavina. Detailed descriptions of south Indian species are available in the works of Naidu (1961-1963). Sharma et ale (1976) I)rought to light the chromosome number in Dero indica and Aulophorus furcatus. Revised descriptions of the world species and keys for their identification have been provided by Brinkhurst and Jamieson (1971). Family Tubificidae
Taxonomy of Indian tubificids has been dealt with by Stephenson (1923) and Naidu (1965). An up-to-date account on their morphology, ecology, biology and taxonomy has been presented by Brinkhurst and Jameison (1971) in a monograph, "Aquatic Oligochaeta of the world" Chromosomal patterns in two species, Limnodrilus hoffmeisteri and Branchiura sowerbyi, were studied by Sharma et ale (1976). Ecology of Tubifex tubi/ex in the Veli lake on southwest Indian coast has been worked out by Shobana and Nair (1983). Family Phrncodrilidae
This family is restricted to southern hemisphere with only one species, P hraeodrilius zeylanicus, known from our region (Sri Lanka). Superfamily Criodriloidea Family Criodrilidae
The family is endemic in the palaearctic region. Immature specimens of circummundane Criodrilus lacuum collected from different Indian localities possibly belong to Glyphidrilus. Superfamily LUlnbricoidca Family Lwnbricidac
Endemics of this family are restricted to temperate ones of North America and Eurasia. However, several species have been transported to almost all continents presumably in soil around roots of plants carried by man. Successful colonization of these has taken place in southeast Asia only at elevations with temperate-like climate. They are now widely distributed in the western Himalayas and Darjccling Hills. Cemosvitov (1973) and Gates (1939) redescribed lumbricid spccies in the collection of the Zoological Survey of India. Gates (1958) further published revised descriptions of species known from the Indian mainland. A comprehensive account of all known species in the Indian subcontinent is available in Gates' (1972) monograph on Burmese earthworms. Julka (1981) reported anthropochorous lumbricids from Lahaul &Spiti in Himachal Pradesh with an average elevation of 3000 In. Superfamily Glossosclecoidca Fanlily Glossoscolccidae
This family is represented by a single neolIopicalperegrine species, Pontoscolex corethrurus, from various localities in Bunna, Andaman and Nicobar Islands and Indian lowlands. At places, particularly in rubber plantations of lower Burma and kerala, it has become numerically dominant species. Its biology has been worked out by Arunachalam (1978) in the Palni Hills of south India. Family Almidae
A single genus, Glyphidrilus, with four endemic species occurs in India and adjacent countries. All these species are limicolous and have discontinuous distribution. Revised descriptions of Indian species have been given in the works of Brinkhurst and Jamieson (1971) and Gates (1972). Naie (1938) published a detailed account on the anatomy of Glyphidrilus anandalei. Superfamily Megascolccoidea Family Ocnerodrilidae
About 21 genera comprising 108 spccies are known in the world. Of these, seven genera with 15 species occur in India. Taxonomic descriptions and distributions of these are available in the works of Gates (1949, 1956, 1972) and Julka (1976; 1990, with Paliwal). Indian Ocnerodrilids are represented by neotrpical Eukerria, ethiopian Gordiodrilus and Nematogenia,and neotropical and ethipian Oenerodrilus circummundane pergrine species. Monospecific Curgiona known only from southern tip of western Ghats may also be of ethiopian origin. Other ocnerodrilid genera belonging to the subfamily' Malabariinae are definitely endemic in this subcontinent and presumably evolved in the peninsular India. These genera are M alabaria in the Gangetic plains, western Ghats, peninsular plateaus and Burma; Deeeania in northern and southern portions of western Ghats; Thatonia in north and east plateaus of the peninsula. Family Acanthodrilidae
This family comprises about 27 genera and 480 species in the world. Indian acanthodrilids are represented by three genera and 18 species. Gates (1972) published the latest key for identification of oriental gcnera and species but deals with detailed descriptions of Burmese species only. Descriptions of Plulellus species from the Indian mainland and Sri Lanka are available in the works of Stephenson (1923) and Julka (1975, 1976, 1981). Two genera, Pontodrilus and Microseolex arc represented by circummundane species' P. bermudensis and M. phosphorous. Subbarao and his colleagues (1972-1978) have provided information on salinity tolerance and bionomics of P. bermudensis in the coastal region of Andhra Pradesh. Family Octochaetidae
Taxonomy of Indian octochaetids has been worked out in detail. Stephenson (1923) distinguished genera of this family (as defined now) among three subfamilies of the Megascolecidae. Gates was not convinced with this classification and undertook revisionary studies on octochaetid genera. Between 1937 and 1940 he recognized several new octochaetid genera Seolioseolides, Barogaster, Lennogaster, Rillogaster, Pellogasler, Priodoehaeta, Priodoscolex. Travoscolides and Celeriella. Gates (1962) transferred most species of Oetoehaetoides to his new geneus Oetoehaelona. Julka (1988) published a comprehensive monograph on the Octochaetidae under Fauna of India series. In this work, he designed Indian species of Ilowaseolex to a distinct genus Wahoseolex, and also rccognised another five new genera from various parts of the country. Descriptions of Burmese species have been provided by Gates (1972) in his monograph. Jamieson (1977) described two new species of Celeriella from Palni hills. Family Megascolecidae.
This family is the largest one in having 1000 known species in the world. In our subcontinent it is represented by 25 genera and 127 species. Descriptions of all known species from Burma, northeast India and Andaman and Nicobar Island have been dealt with by Gates in 1972. He revised the Indian spccies of Pherelima in 1937 and Lampilo in 1938. He discovered two new genera : Nellogaster in 1938 and Nelloseo/ex in 1939. Sims and Easton (1972) revised the world species of Pheretima and recognised new genera of Pheretimoid complex: Pithemera, Metapheretima and Metaphire. Easton (1976) studied atonolny and distribution of Metapheretima elongata species complex of Indo-Australian region. Jamieson (1977) discovered a new genus, Troyia, from Palni Hills. ~new genus Kanchuria and its three new species were brought to light by Julka (1988) from the Garo and Khasi hills in Meghalaya. Since Bahl's work, P heretima posthuma has attracted the attention of several histochemists and physiologists in the country .. Various aspects of another common species Lampito mauritii have also been worked out. Family Eudrilidae
The family is restricted to the ethiopian region. But one species Eudrilus eugenae has acquiroo worldwide distribution in the tropical regions presumably through the agency of man. In India this species has been collected on one or two occasions. Recently, culture of this species have been utilized for the production of vermicompost at University of Agricultural Sciences at Bangalore.
Current Studies
Taxonomy and ecological studies on earthworms of fragile western Himalayan ecosystem are being vigorously pursued at the High Altitude Zoology Field Station of the Zoological Survey of India at Solan in Himachal Pradesh. In connection with the writing up of the second volume on megadrile oligochaetes under Fauna of India series, topotypes of several species from various parts of tile country are under study.
Ecological studies at HAZFS ZSI, Solan include investigations on population dynamics, biomass and habitat preference of earthworms in Himachal Pradesh. Protozoan parasites of these organisms are also being studies.
Studies on ecology and energy budget of earthworms are being carried out at Sambalpur University in Orissa. Agriculture ScienJists at the University of Agricultural Sciences at Bangalore are conducting investigations on their basic ecology and vermicomposting techniques. Some ecological researches on earthwonn ecology are in progress at Kakatiya University in °Andhara Prades~ and G.B. Pant Institute of Himalayan Ecology at Almora.
Expertise India
In ZSI
1.M. Julka, High Altitude Zoology Field Station, Zoological Survey of India, Solan, Himachal Pradesh
T.D. Soota, Zoological Survey of India, 234/4, A.1.C. Bose Road, Calcutta 700 020.
K.R. Halder, R.N. Mukherjee &R. Paliwal, Zoological Survey of India, M-Block, New Alipore, Calcutta 700 053
Elsewhere
M.e. Dash, B.K. Senapati &P.C. Mishra, School of Life Sciences, Sambalpur University, Jyoti Vihar, Burla (Orissa) M.V. Reddy, School of Environmental Studies, Kakatiya University, Warangal (A.P.) R.V.S.S.R. Subba Rao, Department of Zoology, Andhra University, Waltair (A.P.) R.V. Krishnamoorthy; R.D. Kale &Kubra Bano, University of Agricultural Sciences, O.K.V.K., Bangalore (Karnataka) S.A.'Ismail, National College Madras, Madras (Tamil Nadu) P. Dey, Ihargram Raj College, Jhargram, DistL Midnapore (West Bengal). D.P. Halder, Kalyani Agriculture University, Kalyani (West Bengal) Janardhan Singh, Department of Entomology, College of Agriculture, Benaras Hindu University, Varanasi. P.S. Ramakrishnan, Department of Environment Biology, Jawaharlal Nehru University, New Delhi. Tunecra Kumar Bhaduria, G.B. Pant Institute of Ecology &Environment, Kosi, Dislt. Almora. (U.P.) S.M. Handa &M.L. Sareen, Punjab University, Chandigarh.
Abroad
M.B. Bouche, C/o Station-dc-Recherche, Sur la Faune du Sol, 7, rue Sully, F-21034 Dijon Cedex France.
Brinkhurst, Institute of Ocean Sciences, Sidney, British Columbia, Canada. Hrabe, Hrabe Oligochaeta Collection Obranci Mira 70, C S 60200, CeskO$lovenska
B.G.M. Jamieson, Zoology Dcpartment, University of Queensland, Australia -4067 Kr-zysztof Kaspr-zak, Wydzial Ochrony Srodowiska, Urzedu Wojewodzkiego, AI. Stalingradzka 16/1860-967 Poznan K.E. Lee, Division of Soils, C.,S.I.R.O., Adelaide, Australia Edu Bois Reymond Marcus, Rua Sorocaba47 02146 Sao Paulo, Brasil Gilberto Righi, Departamento de Zoologia Instituto de "Biociencius Universidade de Sao Paulo C.P. 11.230-209 01000 Sao Paulo, Brasil MJ. Song, Kyung Pook National University, Deparunent of Biology, Taegu 630, Republic of Korea. R.W. Sims &E.G. Easton, British Museum (Natural History) London, U.K. J.A. Talavcera, Departamento de Zoologia Facultad de Biologia Universidade-de-La Laguna, Tcnerife, Isles Canariacs. Tan Tian-Juc Section of Anatomy, Human Agricultural College, Changsha, China Thai Tran Bai, Pedagogical Institute of Hanoi Vietnam. Zhong Yuam-Hui, DeparLmcnt of Biology, Sichuan University, Chengdu, China. I. Zazonc, Kctendra Hudinasloa a Zoologic CS -949 01 Nitra, Ceskoslovensko. A. Zicsi, Ticrsystematischen inslitut, der horian -Eotouois Universital Puskin Ultac 3 H-8000 Budapest, Magtyursos7..ag.
Selected References
Bahl, K.N. 1950. The Indian Zoological Memoirs. I. Pheretima. 4th. edition, 84 pp. Lucknow Publishing Housc, Lucknow. Brinkhurst, R.O. &Jamieson B.G.M. 1971. Aquatic Oligochaetes of the world. xii + 860 pp. University of TronLo press, ToronLo and Buffalo. Dash, M.C. 1983. The Biology of Enchylraeidae (Oligochaeta), vii + 171 pp. International book distributors, Dchradun.
Gates, G .E., 1972. Burmese Earthworms. An introduction to the systematics and biology of megadrile oligochaetes with special reference to southest Asia. Trans. Am. phil. Soc., 62 (7) : 1-326. Julka, J.M. 1988. Megadrile Oligqchaeta (Earthworms) : Haplotaxida : Lumbricina : Megascolecoidea : Octochaelidae. The Fauna ofIndia and the adjacent 'countries, xiv + 400 pp. Zoological Survey of India. Michaelsen, W. 1910. Oligochaeta. Das Tierreich., : 1-575.
Sims, R.W. 1980. A ClassifICation and the distribution of earthworms of suborder Lumbricina (Haplotaxida : Oligochaeta). Bull. Br. Mus. Nat. Hist. (zool), 39 (2) : 103 -124. Sims, R.W. &Easton, E.G., 1972. A numerical revision of the earthworm genus Pheretima (Megascolccidae : Oligochaeta) with the recognisation of new genera and an appendix on the earthworms collected by the Royal Society North Borneo Expedition. Bioi. J. Linn. Soc., 4(3) : 169-268.
Stephenson, J. 1923, Oligochaeta. The Fauna ofBritish India, including Ceylon and Burma, xxiv
518 pp., Taylor and Francis Ltd., London. Stephenson. J. 1930. The Oligochaeta, xvi + 978 pp., Clarendon Press, LondoD.
