Faunal Diversity in India: Acanthocephala
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FAUNAL DIVERSITY IN INDIA
J. R. B. Alfred
A. K. Das
A. K. Sanyal.
Zoological Survey of India,
( J. R. B. Alfred was
Director, Zoological Survey of India)
Phylum Acanthocephala form a compact group of endoparasitic worms parasitizing all groups of vertebrates and some of invertebrates such as molluscs, crustaceans and insects. Its taxonomic position in the animal kingdom had been uncertain till Lankester (1900) proposed to consider Acanthocephala as a phylum. Later, Van Cleave justified the proposal through convincing arguments. Presently, Lankester's proposal has been widely accepted. The shape of these worms are fusiform, oval, or cylindrical and their, size ranges from less than one millimetre to more than one metre. They are found in intestine, body cavity, mesentery, etc. These worms lack digestive tract in all stages of development and their nutrition takes place through body wall. The body is divided into the presoma including proboscis and neck, and the body proper or trunk. The body wall consists of cuticle, syncytial hypodermis and parietal muscle layer which is delimited from the body cavity by connective tissue lining. Through hypodermis pass numerous lateral lacunae and smaller lacunae forming a lacunar system which is having a considerable taxonomic value. The body has a unique proboscis armed with numerous hooks of different sizes and shapes except in Apororllyncllus lIe11lignatlzi, whose proboscis has pits in place of hooks. The arrangement of hooks on the proboscis, the number of spiral or longitudinal rows, and the number of hooks in the rows are of considerable significance in the taxonomy of Acanthocephala. Cylindrical or sub-cylindrical muscular proboscis, and paired contractile lemnisci are also of great taxonomic value.
Of the reproductive system, position of female genital aperture, position of testes, nature and number of cement glands and presence of Saeftigen's pouch help to determine the individual taxon.
Status Of The Taxon
Global and Indian Status
From the end of 17th century until 1932, about 260 species of Acanthocephala were described from the global envirorunent. At present, there are 800 species known from all over the world.
Till 1890 all the species of Acanthocephala were contained in a single genus Ecllil1orhyl1chus. Subsequently, Lankester's proposal to consider the Acanthocephala as a phylum was accepted by all (Kholodkovaskii, Skrjabin and Sultz, Van Cleave, Petrotscherko, Golvan, Schmidt and many other modem taxonomists). Today this phylum is represented by four classes, namely, Eoacanthocephala, Palaecanthocephala Archiacanthocephala and Polyacanthocephala.
The studies on Acanthocephala in India are made sporadically. As a result, majority of the animal hosts are still lying Wlexplored. Very recently (1990), a checklist on Acanthocephala of fishes of India and Pakistan, published by V. Gupta and G. Sinha added to the knowledge of Indian Acanthocephala and their distribution. Acanthocephalan diversity in India and the world is shown in Table 1. Table -1 Acanthocephalan diversity in India and world
Distribution of Acanthocephala is largely dependent upon the distribution of their hosts. The animal groups which harbour Acanthocephala at larval or 96 Faunal Diversity in India adult stage are found in both terrestrial and qauatic ecosystems. Some of the Acanthocephala are higWy pathogenic causing various diseases to swine, rodents and ducks. Attempts were made by Meyer (1933) to show country-wise distribution of Acanthocephalan species. Later, Petrotschenko (1956) indicated the zoogeographical distribution of these species by demarating several geographical regions. However, based on C\,urent report a state-wise and host-wise distribution of acanthocephalan genera and species are shown in Table 2. Table -2 State-wise and host-wise distribution of number of genera and species of Indian Hosts Fish Amphibia Reptilia Bird Mammal State G. sp. G. sp. G. sp. G. sp. G. sp. Andaman 11 55 Andhra Pradesh 4 5 1 1 Arunachal 1 1 Assam 1 1 Bihar 6 9 11 Delhi 1 1 Goa 22 Jammu & 3 14 22 Kashmir Kamataka 22 11 Kerala 16 32 3 4 Madhya Pradesh 22 Maharastra 3 3 2 51 1 Manipur 1 1 Meghalaya 3 6 1 1 3 10 Orissa 10 16 1 1 Punjab 2 6 3 4 3 16 3 4 Rajasthan 2 3 Sikkim 11 Tamil Nadu 11 15 4 5 2 2 Tripura 3 4 22 4 10 11 UttarPradesh 5 18 1 3 6 5 12 3 23 West Bengal 9 20 123 3 G = genus, sp = species. BHATIACHARYA : AcantlJOcl!l'llala
Out of 229 species of AcantIJoceplJala reported from India 203 species have been known from the indian region only. This indicates a very high percentage of endemicity of these parasites in the present state of knowledge on this group. The remaining 26 species (2 species from fishes, 1 species from Amphibia, 17 species from birds and 6 species from mammals) have been reported from other countries.
As a rule, parasites cause great harm to the hosts' body. Acanthocephala being endoparasites are not in exception. ===Pathogenicity in relation to wild life===: The pathogenic effect of Acanthocephala on the host animals is diverse. Through their body surface they absorb a considerable quantity of nutritive substance from host's body, hindering growth of the host and causing emaciation. Secondly, the thorny proboscis, the hold-fast of the parasite, pierces the intestinal wall of the host causing degeneration. Sometimes it causes peritonitis and usually results in the death .of infected animals. This is observed particularly in macracanthorhynchiasis of swine. Acanthocephala also cause much harm to poultry with the disease polymorphiasis and filicoliasis in ducks and geese. Commercially valuable animals, such as, birds (ducks, geese, etc.), fishes (salmon, carps, etc.) and many other domestic and wild animals are reported to have been incurring losses with the infection of acanthocephala.
Pathogenicity in relation to man
Acanthocephala are not obligatoy parasites of man but reports on human infection by Macracanthorhynchus hirudinaceus of swine in many cases show high pathogenicity of the parasite, causing acute anaemia which often leads the diseased to death. Monilifonnis monilifonnis of rodents is also another parasite found to parasitize man, causing considerable harm to him. Two other species of Acanthocephala, viz., AcanthoceplJalus buJonis of amphibia and Corynosoma streemosum of fish have also been reported to parasitize man. Such reports are available only from Czechslovakia, Italy, Indonesia and Alaska not from India.
Conservation And Future Study
Since acanthocephala are endoparasites, their survival and conservation solely depend upon those of their host species. This group of animal is underexplored. Further,. life cycle and pathogenicity of these parasites are also least known. In future, emphasis is to be given to study these aspects besides taxonomic study of this group.
Amin, O. M. 1989. Key to the families and subfamilies of Acanthocephala with the creation of a new class (Polyacanthocephala) and a new order (Polyacanthorhynchida). J. ParasitoI. 73(6).
Golvan, Y. J. 1969. "Systematique des Acanthocephales (Acanthocephala Rudolphi, 1801) L'ordre des Palaeacanthocephala Meyer, 1931. Memoires du Museum National D'IJistorie Naturel/a. Serie A. Zoologie, Tome LVII.
Meyer, A. 1933. Bronn's Klassen and ordungen des Tievreichs Acanthocephala, Vol. 4: 582 pp. Petrotschencho, V. I. 1958. Acanthocephala of domestic and wild animals TI. 435 pp. Petrotschencho, V. I. 1958. Acanthocephala of domestic and wild animals TIl, 458 pp. Schmidt G. D. 1980. Foundations of Parasitology (Phylum Acanthocephala: Thorny headed worms). Chapter-32. Yamaguti, S. 1963. Systema helminthum, vol. 5 Inter Science Pub!. N. Y. 423 pp.
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Acanthocephala or thorny headed wonn is an important group of animal endoparasites occurring widely in adult stages, and are of direct economic importance to man. Their life history involves one or more hosts,generally invertebrates or lower vertebrates. Very little is known about the damage caused by these worms and their impact on the economy of man.
Redi (1684) •and Leenwenhock (1695) were the earliest workers on record to have noticed Acanthocephala. However the term Acanthocephali (meaning organisms with hooks on the head) was first used by Koelrenther (1771) for a group of helminths and he named a fish parasite as Acanthocephalus. In 1776, Zoega and Muller i~dependently named the same parasite as Echinorhynchus. Later, Muller (1777-1788) described several forms under the genus Echinorhynchus.
Zeder (1800) gave these worms the name Haken wurmer (hooked worms) because of the characteristic hooks and spines invariably associated with the head. Rudolphi (1808) established the order Canthocephala to accommodate these worms, variously known as Acanthocephali and Hoken wormer. Till 1890, the order contained a single genus Echinorhynchus Zoega
The rapid accumulation of diverse forms under a single genus necessitated the reappraisal of the group. Hamann (1892) was the first to attempt a systematic classification of the order Acanthocephala, recognising a number of genera and families. He established the genera Gigantorhynchus and Neorhynchus, bringing several species included earlier in Echinorhynchus under his new genera. He further erected two new families viz. Gigantorhynchidae and Neorhynchidae to accommodate each of his two new genera.
Since then rapid progress in systematics of this group was made by a number of parasitologists, especially, Luhe (19 04, 1912) Kostylew (1915), Travassos (1917, 1926), Ward (1913), Southwell (1933), Van Cleave (1923) and Maplestone (1926). Earlier classifications of this group were based on two main premises, some workers divided the taxonomic hierarchy on the basis of nature of nuclei in the hypoderm and lamnisci; others like Southwell and Macfie (1925) took others cognisance of the nature and character of the prostate glands and proposed a tentative classification using this character. Based on the structure of probosis hooks and body spines, Tbapar (1927) divided the class Acanthocephala in three orders viz. Apororhynchidea, Ecbinorhynchidea and Acanthogyridea.
Van Cleave (1928) erected the families Pallicentidae and Hebosomatidae, utilising not only the proboscis hooks and body spines, but also laying special stress on the nature of nuclei in the subcuticula. Myer (1933) accommodated twelve families and fifty eight genera known till then into two new orders, viz. Palaeacanthocephala and Archiacanthocephala, elevating the order Acanthocephala to the status of a class. He relied in his scheme of classification on the presence or absence of trunk spines and nature of lacunar system.
Van Cleave (1936) added another order, Eoacanthocephala and divided it into two suborders Gyrncanthocephala and Neoacanthocephala. The same author (VanCleave 1948) made a comprehensive review of the whole group and elevated. Acanthocephala to the status of a separale phylum. Hyman (1951) agreed with Van Cleave and recognised, Acanthocephala as phylum but did not accept Van Cleave s classification as such She recognised only three orders in it. viz. Archiacanthocephala, Palaeacanthocephala and Eoacanlbocephala and discarded the two classes of Van Cleave. Petroschenko (1956) did not accept the status of phylum and divided class Acanthocephala into three subclasses viz. Neoechinorhynchinea, Echinorhynchinea and Gigantorhynchinea. Golvan (1958, 1961) modified the classification of Acanthocephala laying too much emphasis on the number of cement glands, ignoring the significance of trunk spines. Yamaguti (1963) proposed a modified classification of the group. He accepted Acanthocephala as a class under phylum Nematehelminthes and recognised four orders namely( 1) Apororhynchidea Thapar, 1927, (2)Neoechinorhynchoidea Southwell and Macfie 1925), (3) Eminorhynchoidea Southwell and Macfie 1925 and (4) Gigantorhynchoidea Southwell and Macfie 1925.
The position of Acanthocephala in systematic hierarchy has been controversial and as yet there is no agreed opinion among parasitologists about its status.
Cuvier (1817) included these parasites among natworms•. Leuckart (1848) considered Acanthocephala near to cestodes and included both of these as two separate orders in the class Anenteraeti. Vogt (1851) distinguished two groups among these worms, viz. flatworms and round worms calling the former as Platyhelmia and the latter as Nematelmia which contained gregarines nematodes, acanthocephalans and goraiaceans.
Gegenbaur (1869) amended Vogt's Nematelmia to Nemathelminthes. Schneider (1866) combined these and some other 'groups into Chaetognatha on the basis of musculature. Hamann (1891) went into the question of the relationship of Acanthocephala on the basis of similarity between the embryo and pseudometamerism present in Moniliformis and some other larger forms and considered them more related to cestoda than to Nematoda with which they shared in behaviour of nuclei and similarity in musculature. Choloakovsky (1897) agreed with Hamann's contention and considered cestode and Acanthocephala to have common origin. Grobben (1908) proposed a new phylum Aschelminthes to include Acanthocephala along with Nematoda, NematomorphaJGastrotricha and Rotifera, etc. More or less this arrangement was also accepted by Rauther (.1931) and Meyer (1932, 1933). Van Oeave (1924, 1941) discussed the relationship of Acanthocephala and concluded that this was related to Cestoda. He p~oposed elevation of Acanthocephala to the status of Phylum near Platyhelminthes. Recently, Amin (1982, 1985) discussed the classification of Acanthocephala and treated it as a phylum. This view was accepted by Arai (1989). The nearness of Acanthocephala to Platyhelminthes is more or less accepted now.
Work done in India
The earliest record of acanthocephalan parasites from India was made by Chandlez (1923) who described Centrorhynchus erraticus from the intestine of a cat from Calcutta. Thapar (1927) was the first Indian Zoologist to have initiated studies on Acanthocephala. He not only described a new genus and species. viz. Acanthogyrus acanthogyrus from the intestine of a carp, Calla catla. at Lucknow but also proposed a tentative classification of the group. Datta (1927) described a new species of Echinorhynchus from crow. Subramanian (1927, 1936) worked out the acanthocephalan parasites from Burma. Van Cleave (1928) erected a new family Filisomidae to accommodate a new genus and species Filisoma indicum obtained from the intestine of a fish collected from Chilka Lake, Orissa. Venna and Datta (1928, 1932) reported acanthocephalan parasites of birds. Further contributions to our knowledge of Indian Acanthocephala were made during thirties and forties by several workers.
Bhalerao (1931. 1937) described a new species of Pallissentis from the intestine at a fish from Nagour, and another from fowl. Moghe (1931) made general reference to the group. These studies were followed by a host of workers in subsequent years. Thus Datta (1936, 1937, 1940. 1947. 1953) added several forms including a number. of new species and two new genera (Mehrarhynchus and Raosentis) from fishes. Further contributions on tht. acanthocephalan fauna of India were made by Podder (1937, 1938, 1941), Datta and Podder (1938), Sen (1938). Kaw (1941, 1948, 1957), Das (1950, 1952, 1957), Moghe and Das (1953), Sarkar (1953, 1954, 1956), Datta and SOOla (1955. 1956), SOO18 and Sen (1956), Agarwal (1958), Farooqi (1958), Gupta (1958), Tripathi (1959), Pal (1963). Nath and Pande (1963), Sahai, Sinha and Rai (1967, 1971), Gupta and Lata (1968). Soota, Sriyastava and Ghosh (1970) described Acanthocephala from Andaman and Nicobar Island.
These papers deal with adult parasites, mostly described from fishes, but also include forms obtained from other vertebrates like amphibia, reptiles, birds and mammals. In recent years, very little work has been done on this group and it is confmed to fish Acanthocephala. Notable contributions have been made by Sood (1972), George and Nadakal (1973), Sen.(197S), Jain and Gupta (1979), Farooqi (1980), Soota and Bhatacharya (1981, 1982), Agrawal and Singh (1982) and few others. Verina, (1973) gave the key to the species of the genus Acanthogentis. Anantaraman and Anantaraman (1975) gave a synopsis of the Acanthocephala described from India till then. Rengaraju and Das (1976) reviewed the taxonomy of Acanthocephala and mostly adopted Van Cleave's classification without going into its merits. Rengaraju (1979) described Porrorchis indicus. Not much works has been done in India on the larval and juvenile stages of Acanthocephala. Sita (1949) studied the life history of Moniliformis moniliformis a parasite ofmt.
Gupta (1950) described developmental stages and juveniles of Centrorhynchus ptyasus parasitising rat snake. Ptyas mucogus. Das (1950,1952,1957) recorded several new as well as known species of Acanthocephala from amphibians and birds dealing with adult, juvenile and larval stages of several species belonging to genera Controrhynchus, Meiorhynchus, Pseudoporrorchis, Arhythmorhynchus. Ghosh and Chauhan (1975) reviewed the work dorie in the Z.S.I. in last fifty years.
Estimation or Taxa
Twenty five families distributed under four orders have been recorded SO far from various parts of the globe. Estimated number of families, genem and species in India are detailed below: No. of families GeIua Species 15 24 110
The work done on this group in the country is too meagre when viewed in the light of host species available in the country. The work has been sporadic and systematic survey based on host Species and their distribution is desirable.
M. Hafeezullah, C.B Srivastava, S. B. Bhattacharya, R. K. Ghosh, 'M' Block, New Alipore, Calcutta -700 053.
T. D. Soota, 234/4, A.J.C. Bose Road, Calcutta -700 020.
H. U. Farooqi, Zoology DeptL, AMU, Aligarh. N. Agrawal, Zool. Deptt, Lucknow Univ., Lucknow. V. Rangaraju, Zoology Deptt., Sholapur College Sholapur. H..S. Singh, Zoology Deptt., Meerut University, Meerut. M. L. Sood, Zoology DeptL, Punjab Agricultural Univ., Ludhiana. V. Gupta, Zool. Deptt., Lucknow Univ, Lucknow.
Richard L. Buckner, School of Life Sciences University of Nebraska Lincohn, Nebraska 68588. B. B. Nickol, School of Life Science, University of Nebraska Lincoln, Nabraska 68588. J. A. Jackson, Zoology Department, Mississippi State University, Mississippi 39762. H. P. Arai. Dept. Bio. Sc., University of Calgory, Calgary, Alberta T2NIN4. V. T. Petrochenko, Institute of helminthology, Academy of Sciences of USSR, Leningra. Y. J. Golvan, Inv. Div., Paris Museunl, Paris, Fance. L. Margolis, Deptt. Fish &Oceans BioI. Sc., Br. Pacific Bio. StD., Nanaimo, Br. Columbia. VaR5 K6.
G. D. Schmiat, Dept. BioI., Colorago State Cell•GreeleY"Colorado 80631.
Amin, O. M. 1985. Classification in Crompton &Nickol (Ed.) Biology of Acanthocephala. Cambridge Univ. Press.
Bykhovskaya, I. E. et ale 1962. Key to the parasites of freshwater fisbes of the USSR. (Eng. Translation). Academy of Sciences, USSR.
Glovan, Y J. 1961. Le phylum des Acanthocephala (3 note) La classe de Palaeacanthocephala Meyer, 1931. Ann. Parasit., 35(1-2): 138-165; 35(3): 350-386; 35(5-6): 713-723; 36(1¬2): 76-91. Hyman, L. H. 1951. The Invertebrates : Acantbocepahala Aschelmenthes, and Entoprocta 1 00. 572 pp.
Meyer, A. 1932. Acanthocephala. Bronnis klass n. Ordungen les Tierreiche 4 ~bl 2, Buch 2 Leaf I. 1-332 Akademische Verlag., Leipzig Meyer, A 1933. Acanthocephala (concludea) Bronn's klass, U Ordingen aes Tierreiche v 4 Abt. 2.,
Buch 2 Lief 2 : 233-582; Akad. Verlag" Leipzig. Peuroschenko, V. I. 1958. Acanthocephala of domestic and wild animals. P.l. 435 pp. PeuroscheJiko, V. I. 1958. Acanthocephala of domestic and wild animals. Pt 2: 458 pp. Tripathi, Y R. 1959. Studies on parasites of Indian fishes, V. Acanthocephala. Rec. Indian.,
Mus., S4 (1-2) (1956): 61-98. Yamaguti. 1963. Systema Helminthum, Vol. 5: Interscence Pub!. N. Y.423 pp.