Siphonaptera: India
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Siphonaptera
Introduction
The order Siphonaptera (siphon =wbe,_ aptera =without wings) includes secondarily wingless insects known as fleas. They are most closely related to orders Mecoptera and Diptera. The larvae of fleas resemble those of dipterous type but more generalised. They are believed to have common origin with the superfamily Staphylinioidea (Coleoptera).
The Siphonaptera are small, wingless insects, body strongly compressed laterally and generally heavily selerotized. Eyes present, sometimes vestigial or.absent. Antennae short and stout, when at rest lying within the antennal fossae. Mouth parts adapted for piercing and sucking. Thoracic segments distinct, free and not fused, legs well developed, coxae large and tarsi five-jointed. Abdomen large. Metamorphosis complete. Adults ectoparasitic on warm blooded animals. Sexes almost alike but female larger in size having a less abruptly tip-tilted tail and without coiled internal horny attachments of the genital organs which are present in the male.
The fleas are facultive rather than obligatory blood sucking ectoparasites on warm blooded animals. They are brown insects with their body size varying 1.00 to 5.6 mm. in length. The chamcteristic shape of the body, covered with backwardly directed spines and bristles, makes fleas very well adapted for gliding among hair and fur of mammals and feathers of birds, which are their common hosts. They also live in the nests of these hosts. They usually walk but can jump up to 20 cm. high with a horizontal range of about 30 cm. On the average they live from three to four months but females may live up to two years. The pupal stage varies in its duration from 5-7days to 354-450 days in some European species. Human flea can live without food f\lr months or even as long as a year. The female must feed on the blood of its normal host before. it can lay fertile eggs. They like warmth, therefore, they leave the host soon after it dies and becomes cold.
They are most abundant in the tropics but also occur in the temperate and even in the polar regions. In the most recent times, they have been carried into Sahara Desert and many Pacific Islands where they were unknown untill the advent of civilized man. Though they occur as ectoparasites on warm blooded animals, but indoor fleas hide in the cracks and crevices of buildings or in the litter found in the grain godownC) etc.
Economic Importance
The Siphonaptera are of considerable economic importance as th~y are vectors of the most important bacterial disease caused by them, the bubonic plague. This is a disease primarily of rodents and has its ancestral home in the plains of Africa and Asia. The disease became distributed ever more widely even into both Americas and now there are reservoirs from which it might break out into epidemics at any time. In Europe, from 1341-1350 about 25 million persons lost lives due to epidemic called Black Death. Also, towards the end of the 19th Century, smaller epidemics suddenly struck in several parts of the world, the worst in India, where in course of about 20 years some 10 million persons died.
Even today this disease, whenever occurs, takes thousands of victims annually in tropical countries. The Indian rat flea, Xenopsylla cheopsis (Rothschild) is the chief transmitter of Bacillus (Pasteurella) pestis which causes human bubonic plague. After a large number of rats die with this disease, it is only then that the disease spreads to man through flea bite. Pulex irritans Linnaeus is another human flea but it also attacks cats, dogs and horses. It is a cosmopolitan species but its favourite host is the pig. Ctenocephalides felis (Bouche) and C. canis (ewtis) are cat and dog fleas and their larvae consume eggs of the dog tapeworm, Dipylidium caninum, which is occasionally found in human being, particularly in children. Echidnophaga gallinacea (Westwood) is an important pest of poUltry.
Estimation of Taxa
There are more than 2,000 species and subspecies known from the World distributed over 150 genera and 17 families belonging to two superfamilies, viz., Pulicoidea and Ceratophylloidea. Pulicoidea are characterised by the outer internal ridge of mid-coxa being absent, the hind-tibia without an apical tooth on outside and sensilliun with 8 or 14 pits on each side. On the other hand, Ceratophylloidea include majority of the families and have the outer internal ridge of mid¬coxa usually present, hind-tibia usually with pointed external apical tooth and sensillium with 14, more often 16 or even more pits on each side. Fifty two species and subspecies, comprising only a little over 2% of the total Siphonaptera (known from the world), are so far recorded from India, which belong to 27 genera, eight families and two superfamilies.
Historical Resume
i) Pre-1900
Linnaeus (1758) placed fleas along with other apterous insects in the order Aptera De Geer (1778) established a new order Suctoria for fleas by excluding them from the special sectionof the order Aptera. Retzius (1783) treated Suctoria in the Saltatoria or jumping insects but Latreille (1796) adopted S uctoria as a group for the fleas. Leach (1815) included fleas in the Aptera in Suctoria, to which he designated Century Medamoptera and referred to them Suctoria of Latreille. Samoulle (1819) followed Linnaeus and Lamarck in recognizing the order.Aptera and considered Suctoria of Latreille as its synonym. Latreille (1825) erected order Siphonaptera and Kirby (1826) also established order Aphaniptera for the fleas. Kirby (1826) synonymised Aptera ofLinnaeus and Lamarck, Rhynchota of Fabricius and Suctoria of Latreille to Aphaniptera. Stephens (1829) treated fleas in the order Aphaniptera Kirby, but erected the family Pediculidae for them. Other names were given to these insects by several workers e.g. Burmeister (1829) called them 'Pulicina', Walker (1851) refeued to them as 'Suctoria' •where as Haeckal (1896) mentioned them 'Pulicida' But the name of the order 'Siphonaptera' as designated by Latreille (1825) is widely accepted and followed which has precedence over the name 'Aphaniptera' of Kirby (1826) which is also sometimes used.
ii) 1901-1947
During the frrst decade of the century, Baker (1904 and 1905) contributed two important works, 'A revision •of American Siphonaptera, or fleas, together with a complete list and bibliography of the group' and 'The classification of the American Siphonaptera' Taxonomic contributions including manuals and monographs were owing to Jordan &Rothschild (1908), Rothschild (1915), Essig (1926, 1931), Ewing ~1929), Sharif (1930), Jordan (1933), Wagner (1934), Iyenger (1935), Ioff (1936), Fox (1940), Ewing &Fox (1943) and Costa Lima &Hathaway (1946). Among these, Iyenger (Ioc. cit.) provided identification of rat fleas ofIndia.
Works on the morphology of Siphonaptera were mainly by Perfilgew (1926), Wagner (1932), Sharif (1935, 1937b), and Snodgrass (1946). Minchin (1915) studied anatomy of the fleas. Knowledge on the biology of Siphonaptera was due to Beier (1936) and Sharif (1937a).
Observations on bionomics of fleas were made by Mitzmain (191 0) whe~eas larvae were studied by Bacot and Ridewood (1914) and Sikes (1930). Schroeder (1926) discovered fossils of Siphonaptera of genera P alaeopsylla Wagner in Baltaic amber and Pulex (1) in Lower Oligocene of Aix, France.
iii) 1948-1990
During this period, significant contributions were made on the studies of Siphonaptera. Jordan (1948) dealt with fleas as Suctoria in Smart's 'Insects of Medical Importance' Brumpt (1949) studied parasitological aspects of the fleas. Weidner (1953) and Wenk (1953) worked out their biology and anatomy, respectively. Works of Munshi (1960) and Chandra (1971) refer to histology and anatomy of rat fleas, X. cheopsis and X. astia. The emphasis on the studies of plague and effects of insecticides (DDT, BRC, dieldrin, Organophosphorus etc.) on these fleas is evident in the works of Mohan (1960, 1962), Seal (1960), Sharma & Joshi (1961), Krishnamurthy &Joshi (1962), Mohan et ale (1962), Krishnamurthy et ale (1966) and Joshi et ale (1967). Seal &Bhattacharjee (1961) and Chandra (1971) studied bionomics and seaSonal prevalence ofcertain fleas from Calcutta and Kollar (Kamataka). Bai (1972), Prasad (1972,.1973, 1976) and Bai &Prasad (1976) worked on host-relationship, histochemistry, egg yolk: and influence of sex hormones of host on the fecundity of two species. Yurgenson (1982) estimated the physiological age of the fleas.
Reike (1970) recorded two fossils fleas from the Lower Cretaceous of Gyppsland (Australia), of which oue resembles Echidnophaga of family Pulicidae and other is quite unusual having long antennae.
Faunistic works on fleas of Canada, Centra America and Mexico, and Japan, were due to Holland (1949), Traub (1950) and Sakaguti & Jameson (1962). Smirnova (1950) worked on the systematics of the larval stages of some species of fleas. Hopkins (1951, 1952 and 1958) contributed towards the scientific n~es, notes on synonymy and also on Order-group and family¬group names for these fleas. Monographs and handbookS on Siphonaptera were due to Smit (1954) and loff & Scalon (1954). Rothschild & Hopkins (1953-1966) classified the order Siphonaptera into 17 families under two superfamilies, viz., Pulicoidea and Ceratophylloidea. • An illustrated Catalogue of the Rothschild Collection of fleas (Siphonaptera) in the British Museum' by Hopkins &Rothschild (1952-1971) running into five volumes, was a monuinental work. Other important contributions on fleas from different parts of the world were by Hubbard (1940-1967), Abn &Soh (1973), Haig (1978), Keifer (1979), Okereke et ale (1980), Morris (1981), White &White (1981), Edwards et al. (1982), Arnett (1983), Beaucournu &Fain (1983), Rothschild &Haddow (1983) and Darskaya &Suvorova (1984).
Taxonomic works on Siphonaptera from India and adjoining countries were mainly by Traub (1966), Traub &Evans (1967), Lewis (1968, 1970, 1972, 1973), Kulkarni &Bhatt (1972), Lewis et ale (1972), Iyenger (1973), Smit &Rosick (1973), Kulkarni et ale (1974), Smit (1974, 1977) and Michel &Dick (1978). Of these, works by Iyenger and Kulkarni et al., (loc. cit.) reported more than 50 species and subspecies including new records and new species of fleas from India. Singh (1980) referred to the collection and preservation of the fleas.
Studies from Different Environs
The Siphonaptera of India has been poorly studied. Species are known from States of Arunachal Pradesh, Jammu &Kashmir, Kamataka, Kerala, Maharashtra, Manipur, Punjab, Sikkim, Tamil Nadu and West Bengal. There are also certain reports on their occurrence in temperate parts and western Himalaya in India.
Considering the richness of hosts, i.e. species of mammals and birds, and the diversity' of the climatic and geographic conditions in India, there is a possibility of existence of a very large number of species of Siphonaptera. It is therefore, expected that future collecing from the unexplored parts of India particularly those harbouring mammalian and avian fauna, will reveal the Siphonaptera fauna more than known at present.
Classified Treatment
Rothschild &Hopkins (1953-1966) classified the order Siphonaptera into 17 families under two superfamilies, namely Pulicoidea and Ceratophylloidea. Majority of the families belong to superfamily Ceratophylloidea. Iyenger (1973) and Kulkarni et ale (1974) reported 52 species and species and subspecies belonging to 27 genera and 8 families (9 subfamilies) so far known from India. Classification upto subfamilies is given below : Superfamily Pulicoidea 1. Family Tungidae 2. Family Pulicidae (i) Subfamily Pulicinae (ii) Subfamily Archaeopsyllinae
(iii) Subfamily Xenopsyllinae Superfamily Ceratophylloidea 1.Family Vennipsyllidae 2.Family Pygiopsyllidae 3Family Hystricopsyllidae (i)Subfamily Rhadinopsyllinae (ii)Subfamily Neopsyllinae (iii)Subfamily Doratopsyllinae (iv)Subfamily Ctenophthalminae 4.Family Ischnopsyllidae (i)Subfamily Thaumapsyllinae (ii)Subfamily Ischnopsyllinae 5.Family Ceratophyllidae 6.Family Leptopsyllidae
Expertise India
K. G. Narayana Pillai, University of Kerala, Trivandrum (Kerala).
S. M. Kulkarni, National Institute of Virology, Pone (Maharashtra).
Abroad
R. Iyengar, World Health Organisation, P.O. Box 2048 Baghda~ Imq. Robert Traub, Siphonaptera Section, Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. -20560 (U.S.A.).
Selected References
Hopkins, G. H. E. 1951. The scientific name for the fleas. Entomologist, 84 : 208-214.
Hopkins, G. H. E. 1952. Notes on synonymy in Siphonaptera. J. Washington Acad. Sci., 42 : 363-365.
Hopkins, G. H. E. 1958. Order-group and family-group names for the fleas. Ann. Mag. Nat. Hist., 13 (1) : 479-487.
Hopkin,S, G. H. E. &Rothschild, M. 1952-71. An illustrated Catalogue of the Rothschild Collection offleas (Siphonaptera) in the British Museum (Natural History). 1952, 1 : 361 pp.; 1956. 2 : 445 pp.: 1962, 3 : 560 pp.; 1966, 4 : 549 pp.; 1971, S : 530 pp.
Iyenger, R. 1973: The Siphonaptera of the Indian Subregion. Oriental Insects Suppl., No.3: 102 pp., 219 figs. Jordan, K. (1948). Suctoria.In : Smart, J., Insects ofMedical Importance, 2nd. ed. Jarrold &Sons Norwich, England, 295 pp.
Kulkarni, S. M., Bhat, H. R. & Dhanda, V. 1974. A Survey of Haematophagous arthropods in Western Himalayas, Sikkim &hill districts ofW. Bengal-Fleas (Siphonaptera). Indian J. med. Res. 62 (7) : 1061-1088.
Sharif, M. 1930. A revision of the Indian Siphonaptera, Part I. Family Pulicidae. Rec. Indian Mus. 29-62.
Smit, F. G. A. M. 1954. Identification of fleas. In : Poll i tzer , R., Plague, World Health Organisation Monograph No. 22, 698 pp.
Smit, F. G. A. M. 1957. Handbook for the
