Hymenoptera Ichneumonidae: India
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The family Ichneumonidae o~ the order Hymenoptera is one of the ~argest of all animal groups as it includes numerically more species than those of entire vertebrates, and among insects ranks only next to the family Curculionidae of the Order Coleoptera. Of the total species of insects known from the world, 5-8% belong to the family Ichneumonidae. They have mimicking colour pattern, usually in the form of black spots and stripes and are associated with foliage. They are usually parasitic in cocoons of the .order Lepidoptera, but also .parasitize cocoons of S ymphyta, Braconidae, other Ichneumonidae, Neuroptera, Diptera and spiders. They kill the host by feeding on body fluids of the host, then spin their own cocoons, pupate and emerge as adult ichrieumonids. By their parasitic habit they destroy a large number of agricultural and forest insect pests, thereby preventing the undue increase of pest species. They serve as an important agent for the biological control of pests. Thus, knowledge of their speciation, distribution and biology has potential practical value.
The oldest fossils of Ichneumonidae are known from the early Cretaceous. It is estimated that 60,000 species of ichneumonids are known from the world, which are classified in 26 subfamilies. There ar~ 4140 species of ichneumonides in the Indo-Australian region, of which about 1200 species occur in the Indian-subregion.
The family Ichneumonidae is readily recognised by the costal and sub-costal veins touching or fused with each other, and also in the presence of second recurrent vein in the fore wing. In Braconidae and Stephanidae also costal and subcoastal ve.ins agree with Ichneumonidae, but in these families second recurrent vein is always lacking. Some ichneumonids lack second recurrent vein, but then they are recognised by the characters of tibial spurs. The ichneumon ids .have long antennae, with more than 16 segments. The only other hymenopterans with 16 antennal segments are Braconidae, Stephanidae, Pamphilidae, Trigonalidae and Sclerogibbidae. The other distinguishing feature of Ichneumonidae is the mandible (and also in most braconids) having only two teeth, but ther are exceptions in some ichneumonids where the upper tooth is subdivided and the lower tooth entirely absent
The family Ichneumonidae is divided into following 26 subfamilies (those with an asterisk (*) are not known to occur in India) : 1. Pimplinae, 2. Tryphoninae, 3. Eucerotinae, 4. Labeninae, 5. Xoridinae, 6. Agriotypinae, 7. Banchinae, 8. Ctenopelmatinae, 9. Porizontinae, 10. Cremastinae, 11. Tersilochinae, 12. Ophioninae, 13. Mesochorinae, 14. Metopinae, 15. Anomaloninae, 16. Gravenhorstiinae, 17. Acaenitinae, 18. Hclictinae*, 19. Orthocentrinae, 20. Diplozontinae, 21. Phygadeuontinae, 22. Hemigasterinae, 23. Mesosteninae, 24. Ichneumoninae, 25. Lycorininae*, 26. Phrudinac*.
LinnaetJs (1758) placed all Ichncumonidae, Braconidae and other Terebrantia in a single genus Ichneumon. Since then new genera have been added to the family up to 1850, followed by their rapid increase by the end of the century. Table showing number of subfamilies, genera &species known from India Subfamilies Genera Species
Gravenhorst (1829) published a classification of the family Ichneumonidae in 'Ichneumonologia Europaea' His genera were super-generic gro.ups containing subunits which were treated nomenclaturally as true genera, but designated by him as "families or subgenera" To translate his terminology to modem usage, Gravenhorst's "genera" will be called subfamilies, and his "families or subgenera" as genera. Therefore, his subfamilies were: 1. Ichneumon; 2. Tryphon; 3. Trogus; 4. Aloneya; 5. Cryptus; 6. Pimpla; 7. Metopius; 8. Bassus; 9. Banchus; 10. Ophion; 11. Hellwigia; 12. Acocnites; and 13. Xorides.
Wesmael (1844) in his classification divided the family into 36 groups and these were accorded family rank by later workers. Ashmead (1890) reduced the "families" of Foerster to tribes. Foerster (1850-76) proposed 489 generic and species names.
Addition of more new taxa and their classification in more natural groups were significant contributions made during this period by several workers namely: Fabricius, (1798-1804); Latreille (1809); Nees (1815-1818); Gravenhorst, (1823-1829); Haliday, (1832-1838); Kirby, (1837); Boi (1841-1856); Wesmael (1844-1859); Holmgren (1855-1889); Cresson (1856-1873); Kriechbaumer, (1872-1902); Thomson (1873-1893); Brulle (1688); Dalla Torre (1888-1902); Saussure (1892) and Tosquinet (1896-1903). Ichneumonidae 453
The early three decades of this century were dominated by Cameron, (1895-1925); Schmiedeknect (1900-1933); Moley (1903-1917); and Viereck, (1911-1925). Their work included many new taxa, most of them valid even today. Cushman and Rohwer (1920) broughtout a new classification of the tribes of Ichneumoninae ("Pimplinae',). Cushman also made important studies on the ichneumonids upto 1947. Roman (1903-1943) improved upon the classification of Thomson and Schmiedeknect for European species. Seyrig (1932-1934) published an account of Madagascan Ichneumonids.
The fourth and fifth decades of this century were taken over by Heinrich (1934, 1938) on Madagascan ichneumonids. Townes (1944) published a catalogue and revised classification of Nearctic ichneumonids.
The knowledge of Ichneumonidae was also due to the contributions of the following workers namely: Szepligeti (1900-1916); Kukujev (1901-1904); Perkins (1902-1915); Schulz (1906¬1911); Brues (1910-1913); Viereck (1911-1925); Enderlin (1912-1921); Cushman (1915-1947); Clement (1924-1938); Meyer (1927-1935); Heinrich (1930-1965); Betrem (1932-1941); Blanchard (1936-1947) and Cheesman (1938-1953).
During the period addition to new taxa and information on various other aspects were by the efforts of Hedwig (1949-1961); Burk (1952); Gupta et al., (1957-1989); Perkins (1958-1963); Porter (1963-1967); Tobias (1963); Dash (1964); Jonathan (1967-1982) and Fitton (1976-1985). On account of major and significant contributions by Townes (1958-1989) during these four decades, these were often referred to as the "Townes era of ichneumonid research" Up to 1952, his work was confmed to Nearctic fauna, followed by a taxonomic arrangement and catalogue on the Indo-Australian ichneumonids in 1961, on the Palaearctic ichneumonids in 1965 and also on Neotropic ichneumonids in 1966. Townes (1967) undertook a more extensive work on the generic and higher classification of Ichneumonidae on the world basis. His four volumes on "the gener of Ichneumonidae" during 1969-1971 were widely appreciated. Townes and Townes (1973) also ~rought out a catalogue and reclassification ofEthiopian Ichneumonidae.
Studies from Different Environs
Till 1955, out knowledge of Ichneumonidae of Indian subregion was scanty. Morley's "Fauna of British India, Hymenoptera Vol III (part I) Ichneumonidae", published in 1913, did not include important groups such as the Mesostenini (Cryptinae) and Ichneumoninae. Cameron (1901-1902), Cushman and Rohwer (1920), Gupta (1955-1975), Gupta et al., (1966-1975), Perkins (1943), Townes et ale (1944-1984) published elaborate works on the Indo-Australian Ichneumonidae.
Gupta and collaborators (M. K. Kamath, J. K. Jonathan, D. T. Tikar, Sharda Maheshwary, M. L. Gupta, Girish Chandra and Santosh Gupta) have published. several monographs and other revisions on the Oriental Ichneumonidae, jointly as well as independently. Gauld and Mitchell contribued much to our knowledge of tl\e Anomaloninae and Ophioninae of the Orient. ,Kaur (1979), Rao (1972), Nikam (1972-1984) also contributed to the knowledge of Ichneumonidae of the Indian subregion.
Several surveys have been conducted to collect ichneumonid fauna from all parts of India, particularly Assam, Jammu and Kashmir, Himachal Pradesh, Meghalaya, Tripura, Uttar Pradesh, West Bengal (mainly Darjiling Hills), Punjab, Haryana, Delhi, Rajasthan, Kerala and Karnataka. Of these, collections made from Meghalaya, Tripura and Darjiling Hills are represented by a good number in National collections at the Zoological Survey of India.
There are revisionary works on the taxonomy of Indian Ichneumonidae by Jonathan (1973, 1980) and Kaur and Jonathan (1976, 1979). These studies were based on the material available at the Zoological Survey of India as well as material borrowed from various museums in India and abroad. These contributions included taxonomic descriptions of 25 genera and 325 species belonging to subfamilies Mesosteninae and Tryphoninae. Of these. 204 species and subspecies are new to science. The new taxa constitute an increase of 17% of the known species of Ichneumonidae from India. The results of the studies of ichneumonid fauna of States of Meghalaya, Tripura, and West Bengal are under publication.
J. K. Jonathan, Zoological Survey of India, M-Block, New Alipore, Calcutta 700 053.
P. K. Nikam, Department of Zoology, Marathwada University, Aurangabad.
Girish Chandra, Department of Zoology, K. M. College, University of Delhi, Delhi.
M. L. Gupta, Department of Zoology, Deshbandu College, Kalkaji, New Delhi.
S. Maheshwary. Department of Zoology, Miranda College, Delhi.
V. K. Gupta, Director, Entomology and Nematology Department, University of Florida, 8005 1.56th Avenue, Gainesville, Florida 32608, U.S:A. 2.G. Fitton, Department of Entomology, British Museum (Natural history), Cromwell Road. London SW7, 5BD, England. 3.D. Gauld, Hope department of Entomology, University Museum, Oxford, OX 1, 3 PW, England.
Setsuya Momoi, Entomological Laboratory, Faculty of Agriculture, Kobe University, Kobe, Japan.
Ashmead, W. H. 1990. Classification of the ichneumon flies, or the superfamily Ichneumonidae. Proc. U.S. Natl. Mus.• 23 : 1-192.
Gupta, S. &Gupta" V. K. 1983. Ichneumonologia Orientalis, Oriental Ins. Monogr., 10 : 1-313.
Gupta, V. K. &Maheshwary, S. 1977. Ichneumonologia Orientalis, Part IV. The tribe Porozontini (= Campoplegini) (Hymenoptera: Ichneumonidae). Oriental Ins. Monogr., 5 : 1-267.
Gupta, V. K. &Tikar, D. T. 1976. Ichneumonologia Orientalis or a Monographic Study of Ichneumonidae of the Oriental Region, Part 1. The tribe Pimplini (Hymenoptera: Ichneumonidae : Pimplinae). Orienta/Ins. Monogr.• 1 : 1-312.
Jonathan J. K. 1980. The Isotima-Complex. Rec. Zool. Surv. India. Occ. Paper No. 17 : 1-146.
Jonathan, ]. K. &Gupta, V. K. 1973. Ichneumonologia Orientals, Part III. The Gorypbus-complex (Hymenoptera: Ichneumonidae). Oriental Ins. Monogr., 3 : 1-203.
Kamatb, M. K. &Gupta, V. K. 1972. Ichneumonologia Orientalis, Part ll. The Tribe Ryssini (Hymenoptera: Ichneumonidae). Oriental Ins. Monogr., 2 : 1-300. Kaur, R. & Jonathan, J. K. 1979. Ichneumonologia Orientalis, Part Vllt The Tribe Phytodietini from India (Hymenoptera: Ichneumonidae). Oriental Ins. Monogr., 9 : 1-276.
Morley, C. 1913. Fauna of British India, including Ceylon and Burma: Hymenoptera. Vol. Ill. Ichneumonidae. : 531 pp. Taylor &Francis, London. Townes, H. 1969-71. the genera of Ichneumonidae, Parts 1-4. Mem. Amer. Enl. Inst., Nos. 11, 12, 13, 17.