Florida Nematology

History of Nematology

History of Entomophilic


The genus Steinernema

Mole crickets control

Mole cricket nematode

Symbiotic bacteria

Termite nematode

ditic stage of

Nematode identification




History of Insect Parasitic Nematodes

This document was constructed and is maintained by KHUONG B. NGUYEN
Entomology and Nematology Department
University of Florida


This document was prepared for students in the course "Insect Parasitic Nematodes"
Entomology & Nematology Department, University of Florida

It was reported that entomophilic (entos = insect, philos =  like) nematodes were found in about 27 families of 8 orders, namely Rhabditida, Tylenchida, Aphelenchida, Strongylida, Oxyurida, Ascaridida, Spirurida,, and Mermithida.

termiteSteinernemaS. scapteriscipulchrocephalaepiptygma
  Neosteinernema                 Adults and juveniles of           Mole crickets parasitized        Pulchrocephala sp.        Face view, tail and epiptigm
 parasite of termite                   Steinernema                 by Steinernema scapterisci         found in mole crickets        of Steinernema females


- Aldrovandus  found dead grasshoppers with worms emerging from their bodies (in De Animalibus Insectis 1623). He was the one who first used the word Vermes.
- Lister (1672) described similar worms from a plant in his garden and compared his findings to that of Aldrovandus.
- Reaumur (1742) described a worm that was undoubtedly Sphaerularia  bombi Dufour.
- Gould (1747) graphically described the emergence of worms, probably mermithids, from ants.
- Carl von Linnaeus (1707-1778), in his Systema Naturae, listed eight genera in the Vermes Intestini (1758).  Two of these were truly parasitic worms, and the name of the third, Gordius, is associated with Linnaeus.
- A Lutheran pastor, J.A.E. Goeze (1731-1793), was the first to study nematodes seriously under the microscope and described the vinegar eelworm (1782).  He began to distinguish between the various kinds of worms.  He also described the emergence of mermithids from soil following a heavy rain (1782).
Rudolphi(1771-1832). Rudolphi (1819) included 350 species belonging to 11 genera in his works Entozoorum Synopsis.  He also gave us the scientific name, Nematoidea.
 - Siebold(1804-1885) studies introduced the concept of a life cycle involving different kinds of hosts, with different ways of penetration. With studies of Charvet (1834), Berthold (1843), and Dujardin (1842)], Siebold established the Gordiacea (in 1843), he included mermithids in the group.  Under the title "Ueber die Fadenwürmer der Insekten" in six works (1842-1858), von Siebold described and noted 233 nematode species from insects.
- Leuckart (1822-1898) clarified Linnaeus' groups and established Vermes on a firm basis akin to that which we use today (Fig. 3).  Rudolphi (1809), with true perspicacity, recognized Nematoidea as separate from Acanthocephalea, Trematodea, and Cestoidea, but Leuckart (1887) established them as separate groups.
Hope (1839) wrote "The genera and species of insects infected by filariae" and complicated an already complex puzzle concerning the identity of filariids, mermithids, and gordiids.
- Bremser (1824), in his "New Atlas of Intestinal Worms" recorded Leblond's discussion of the finding of Audouin of mermithid worms in cockchafers in France.  Another French worker,  Dujardin (1801-1860), was also a pioneer in the study of nematodes in insects in France.  He described Mermis nigrescens in 1842 and Mermis aquatilis in 1845.
- In Scotland in 1861, Sir John Bulloch described a worm that was obviously Sphaerularia bombi.  In 1853, Meissner (1829-1905) described Mermis albicans in detail.
- In 1851, Karl Diesing (1800-1867) published Systema Helminthum, with 175 insect nematode records and involving five entomophilic genera.  He listed 118 species of Gordius, 17 of Mermis, including M. nigrescens and M. albicans and Sphaerularia bombi as members of the same suborder and tribe.  Not surprisingly, he considered Sphaerularia as a genus inquirendum.  He assigned twelve species to Anguillula, nine of which had been assigned to Oxyuris spp., and found in intestines of insects.  Diesing recognized two genera, Gordius and Mermis, but kept them in the same suborder and tribe.  It remained for Vejdovsky (1886) to separate Nematoda and Nematomorpha .
- Braun in 1883 defined the Mermithidae, a definition that still stands.
- Linstow (1842-1916), in Berlin, initiated a series of papers that extended from 1860 to 1914. The admirable practice of naming new genera by the use of a new prefix with -mermis originated with von Linstow and was first used in establishing the name Paramermis in 1898.  In 1878 , he published a Compendium der Helminthologie, which lists several entomophilic nematodes.
 - Schneider (1831-1890) provided one of the first classifications of nematodes (1866).  He accepted only M. nigrescens and named a new species M. lacinulata.  He separated Mermis from Gordius, but left Sphaerularia bombi with Gordius in the same group.  In the same monograph, he reveals his curiosity about S. bombi by including a chapter on its development.
- Late in the nineteenth century, three Russian researchers, Fedchenko (1874, 1886), Keppen (1870, 1881, 1882), and Radkevitch (1869), gave us information on the entomophilic nematode fauna of that vast country.  They created a tradition that was carried into the next century by several researchers.  Dr. E.S. Kirjanova(1900-1976) was one of these.  She worked on plant, animal, and insect nematodes, but had a special interest in the Nematomorpha.  She supervised 25 dissertations, and described 100 new species of nematodes and Nematomorpha.
- Writing about insect nematologists in Russia, Filipjev must be considered as a prominent nematologist for nematological science. When writing about Filipjev in the book "Plant and Insect Parasitic Nematodes", Nickle and Welch stated "If Rudolphi is recognized as the Father of Helminthology, then Ivan Nikolaevich Filipjev (1889-1940) may certainly be named the father of Insect Nematology. Not only did he contribute indirectly through his work on nematode classification, but directly by bringing together the scattered information on insect nematodes and incorporated his own findings into the synthesis.  This formed the second section (about 80 pages) of a monograph published in Russian in 1934 entitled "Nematodes that are harmful and useful in Agriculture" (Filipjev, 1934).  Sections of this book were sent to Dr. Prof. J.H. Schuurmans-Stekhoven in Belgium who, as junior author, had the original Russian translated into French and then into the "American Language".  This was published in Leiden under the title A Manual of Agriculture Helminthology in 1941.
 Also in 1934, Filipjev's "Classification of the free-living nematodes and their relation to parasitic ones" was published in English by the Smithsonian Miscellaneous Collections (1934).  Micoletzky (1925) and Cobb (1919) dropped their classification in favor of Filipjev's.  Western workers lost contact with Filipjev, and Kirjanova (1959) reported his death on October 22, 1940.
Filipjev's book on harmful and helpful helminths in agriculture combines both plant and insect nematodes..
Filipjev (1934) included those nematodes that have insects as "intermediate" hosts, the Spirurida and Filariida as well as Nematomorpha.  The limits of a scientific field have no sharp boundaries.  Poinar (1975), for example, includes the Spirurids, Filarids, and Nematomorphs, whereas this book combines the insect-parasitic nematodes and the plant-parasitic nematodes."
 -Hall (1929) presented "Arthropods as an intermediate host for helminths" but regrettably had no references.  Stiles' and Hassall' s 1920 compendium of generic and specific names was praised when published as much as it is esteemed today.  It is a valuable volume filled with names and dates, and remarkably free of errors.  It is known as the Index Catalogue of Medical and Veterinary Zoology, Roundworms.  Checklists of more recent dates include .
- Gilbert Fuchs published a series of papers on bark beetle nematodes.  The series extended from 1914 to 1938
- Yatsenkowsky (1924) in the USSR provided evidence that a small number of nematodes could cause castration of the bark beetle hosts, and heavy infections killed the beetles. Polozhentsev and Negrobov (1967) listed 400 insect species that are intermediate and definitive hosts for trematodes, cestodes, acanthocephalans, and nematodes.  Another Russian worker, Ipatyeva (1970), provided a list of nematodes associated with the Scarabaeoidea..
- In the United States, Cobb(1927), from his experience with three entomophilic nematodes, stressed the potential of nematodes in controlling insects.  In Britain, Oldham (1933), on the basis of his own experience and that of T. Goodey (1930) on Tylenchinemaoscinellae, made the same recommendation.  Both T. Goodey (1951) and J.B. Goodey (1963) produced systematic books containing nematodes associated with insects.
- Detailed morphological and taxonomic studies were strongly pursued in the early decade of this century.  Dadai (1911), Daday (1913), and Hagmeier (1912) established many of the type species for future genera. Cobb (1859-1932), Steiner (1886-1961), and Christie (1889-1978), three Americans, established insect nematology in North America in single, dual, and triple authorships in a long series of papers.  Their " Agamermis decaudata Cobb, Steiner, and Christie 1923; a nema parasite in grasshoppers and other insects" is recognized as a classic, as pointed out by Gerald Thorne (1961).
- Two other Americans, Rudolf William Glaserand Norman R. Stoll , had outstanding contributions.  Glaser (1888-1947) was the first to mention the potential of Neoaplectana nematodes for biological control, and gave much effort to mass-culture techniques and was the first researcher to culture N. glaseri on an artificial culture in vitro.  Norman Stoll (1892-1976) continued Glaser's work on vermiculture (Stoll, 1959).
- Steinerdescribed S. kraussei (1923) and S. glaseri (1929)
P. Bovien (1933, 1937, 1944), made three excellent contributions, in one of them he described Neoaplectana affinis and N. bibionis . Also in this contribution, he suggested the interrelationship among bacteria, nematodes, and insects (1937).
S.R. Dutkyand W.S. Hough (1955), and the Czechoslovakian, J. Weiser (1955), simultaneously found a nematode in the Codling moth that the former designated by its accession number, DD-136, and the latter named the worm Neoaplectana carpocapsae.  Dutky, and later Weiser, confirmed the presence of an associated bacterium.
- J.R. Christie(1974) reviewed the parasites of invertebrates, noting many newly discovered species, chiefly in insects.
- Poinarhas worked with different kinds of insect parasitic nematodes, especially mermithids and rhabditids. He is a "key milestone" in the development of entomopathogenic nematodes. He has authored or coauthored five books and more than 400 publications on nematode systematics, structure, bionomics, paleontology, natural enemies, epizootiology, and host parasite relationships. At the present time (2002), he is still working in evolutionary of nematodes at Oregon State University.
- Nickle  provided lengthy papers (1967a, 1972, 1973, 1974) showing worms in situ and reviews on taxonomy and biology of insect parasitic nematodes. He has retired from USDA since 1995.
 - Shephard (1974) published the compendium of abstracts, this is of considerable value.
 - Massey(1974) produced a monograph on the taxonomy of nematodes of bark beetles in the United States.
- In the United States, insect pathology was beginning to develop from late 1940's. Steinhaus (1949) devoted a chapter to nematodes. Welch (1956, 1963, 1965) revived interest and reviewed progress in the field, which he named entomophilic nematology.
- In Europe (late 1940's and early 1950's) at the University of Erlangen, Professor H.J. Stammer established a school of insect nematology and from which outstanding students, such as Drs. H. Korner , E. Liebersperger, G. Osche, W. Riihm , and F. Wachek, had great contribution to insect nematology.
 -In Brazil, L. Travassos has a long series of publications (1925-1965) contributing to our knowledge of the oxyurids and thelastomatids in Diplopoda, Chilopoda, and Insecta.  Her 1953 paper is a typical example of her work.  P.T. Artigas published his work in "Systematica dos nematoideos dos arthopodes", 1929. This is a very good document for workers in the field. He also had some good contributions in 1926, 1928.

Additional workers in the field of entomophilic nematodes (retired + active) :

    In the United States: J.J. Petersen, A.A. Johnson, H.K. Kaya, J.G. Stoffolano, C.J. Geden, E.G. Platzer, D.P. Molloy, R.Gaugler, M. Klein, G.C. Smart,
    K. B.  Nguyen, D. Shapiro-Ilan, S. P. Stock, P. S. Grewal, E. Lewis, C. Manion, R. Gibblin-Davis, A. M. Koppenhofer, B. J. Adams, L. A. Lacey.
    In Canada: J.M. Webster, C.H.S. Thong, J.R. Finney, and R. Gordon, M. L. Adamson, Guy Belair.
    In United Kingdom: W.M. Hominick and P.N. Richardson, D. J. Hunt, A. Reid.
    In Austria: H. Kaiser.
    In France: C. Laumond, D. Van Waerebeke.
    In Russia:  P.A. Polozhentsev, A.K. Artyukovsky, I.A. Rubtsov, S.L. Lazarevskaya,  G.V. Veremchuk, S. Spiridonov.
    In Ireland: A. Burnell, C. Griffin, B. Dowds, M. Downes.
    In Germany: R.U Ehlers, D. Sturhan.
    In New Zealand: W. Wouts.
    In Australia:  R. Bedding and A. Akhurst, J Curran.
    In Brazil: G.R. Kloss, N. Cordeiro.
    In India: M. A.Basir, S.N. Singh, P. N. Rao, M. N. Farooqui, S. Ganguly.
    In Japan: N. Ishibashi, E. Kondo, Y. Mamiya, M. Yoshida, S. Yamanka.
    In Israel: I. Glazer, D. Segal.
    In Czechoslovakia: Z. Mracek
    In China: J. Liu, J. Heng...
    In Hungary: A. Fodor.


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        Artigas, P. 1926. Nematoides de invertebrados. Boletim Biologico , São Paulo 1:1-13.
        Artigas, T.T. 1929. Systematica dos nematoideos dos arthropodes. Faculdade de Medicina de SaoPaulo. 114 pp.
        Berthold, A. A. (1843). Uber den Bau des Wasserkalbes (Gordius aquaticus). Abhandl. K. Gesellsch Wissensch. Gottingen Math-Phys. CL. 1: 1-18.
         Bovien, P. (1933). On a new nematode, Scatonema wülkeri gen. et. sp. n. parasitic in the body cavity of Scatopse fuscipes Meig. (Diptera, Nema-tocera). Vidensk Medd. Dansk Naturh. Foren. Kdben'havn. 94: 13-32.
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Updated 1 January 2010