Keys to Species of Plant-Parasitic Nematodes

Note: Please notify Paula Agudelo pagudel@clemson.edu of any corrections or additions to this list of keys.

This list of keys and references for identifying species of selected genera of plant-parasitic nematodes was compiled to assist students and professionals in the discipline of nematology. It is not comprehensive but was compiled as a current source of information and initial perspective for those interested in species identification of selected plant-parasitic nematode genera. Nomenclature and systematics follow that of Ebsary, 1991, and genera selected are primarily those in Mai, W. F., and P. G. Mullin, with H. H. Lyon and K. Loeffler. 1996. Plant-parasitic nematodes: A pictorial key to genera. Ithaca, NY: Cornell University Press. We added older references where most appropriate and included some references that are not keys, where keys are unavailable or if particularly useful as supplemental references. For information on older references and on genera not included herein, see E. C. Bernard and S. M. Baird. 1983. Bibliography of keys for the identification of plant-parasitic and free-living terrestrial nematodes. Nematology Newsletter 29(4):9-18. Detailed study should encompass original species descriptions. We recommend the following references for studies of the principles of taxonomy behind many of the keys and for general review:

Golden, A. M. 1971. Classification of the genera and higher categories of the order Tylenchida (Nematoda). Pp. 191-232 in B. M. Zuckerman, W. F. Mai, and R. A. Rohde, eds. Plant Parasitic Nematodes. Vol. 1. Morphology, Anatomy, Taxonomy and Ecology. New York: Academic Press.

The landmark collection of manuscripts entitled: A Reappraisal of Tylenchina (Nemata) published in Revue de Nématologie, 1987-88, by contributors R. Fortuner, E. Geraert, M. Luc, A. R. Maggenti, and D. J. Raski.

Siddiqi, M. R. 1985. Tylenchida parasites of plants and insects. Slough, UK: Commonwealth Agricultural Bureau on behalf of Commonwealth Institute of Parasitology. 645p.

Ebsary, B. A., 1991. Catalog of the Order Tylenchida (Nematoda). Publication 1869/B. Ottawa, Canada: Canada Communication Group-Publishing. 196 p.

Nickle, W. R., ed. 1991. Manual of agricultural nematology. New York, NY: Marcel Dekker. 1035 p. (A valuable reference that has many species keys)

Compiled by:
S. A. Lewis, Department of Plant Pathology and Physiology, Clemson University, Clemson, SC 29634-0377

R. T. Robbins and Weimin Ye, Nematology Lab, Department of Plant Pathology, University of Arkansas, Fayetteville, AR 72701

Z. A. Handoo and A. M. Golden, USDA, ARS, Nematology Lab, Bldg. 011A, Rm. 159, BARC-West, 10300 Baltimore Avenue, Beltsville, MD 20705-2350

Thanks to Dr. E. C. Bernard for his review of this submission.

SOME IMPORTANT KEYS TO SPECIES OF SELECTED GENERA OF PLANT-PARASITIC NEMATODES

Tylenchida Thorne, 1949
Tylenchoidea Örley, 1880
Tylenchidae Örley, 1880 (See key to genera in Manual of Agricultural Nematology)
Tylenchinae Örley, 1880
Tylenchus Bastian, 1865 and related forms
Andrássy, I. 1954. Revision der gattung Tylenchus Bastian, 1865 (Tylenchidae, Nematoda). Acta Zoologica Academiae Scientiarum Hungaricae 1:5-42. (Foundation manuscript for studying Tylenchidae).

Bello, A. 1973. Étude taxonomique du genre Tylenchus Bastian 1865 (Nematoda: Tylenchinae). Nematologia Mediterranea 1:1-7.

Andrássy, I. 1979. The genera and species of the family Tylenchidae Oerley, 1880 (Nematoda). The genus Tylenchus Bastian, 1865. Acta Zoologica Academiae Scientiarum Hungaricae 25:1-33.

Siddiqi, M. R. 1979. Seven new species in a new nematode subfamily, Duosulciinae n. subfam., is proposed to accommodate species with only two incisures (or a single ridge) in the lateral field. Nematologica 25:215-236. (Key to the six genera)

Brzeski, M. W., and M. R. Sauer. 1982. Scanning electron microscopy of some Tylenchidae and Boleodoridae and reappraisal of the Boleodoridae. Nematologica 28:437-446

Sumenkova, N. I. 1984. Survey of generic and suprageneric categories of nematodes in the family Tylenchidae Orley, 1880. Taksonomiya I biologiya fitogel'mintov. 1984. 132-144. Moscow: Nauka. (A key to genera, in Russian)

Ottolenchus Husain & Khan, 1967 (Wu, 1970) Golden, 1971 synonymized this subgenus with Aglenchus, Andrássy, 1954

Brzeski, M. W. 1982. Taxonomy of Ottolenchus Husain & Khan, and a description of Coslenchus polonicus sp. n. (Nematoda: Tylenchidae). Revue de Nématologie 5:71-77.

Maqbool, M. A., and F. Shahina. 1985. Two new species of the genus Ottolenchus Husain & Khan, 1967 (Nematoda: Tylenchidae) with a key to the species and observation on O. facultativus (Szczygiel, 1969) Brzeski, 1982 from Pakistan. Revue de Nématologie 8:329-333.

Sumenkova, N. I. 1987. Taxonomy of phytonematodes of the genus Ottolenchus (Nematoda: Tylenchidae) and a description of two new species of the genus. Zoologicheskii Zhurnal 66:1805-1811.

Filenchus Andrássy, 1954
Raski, D. J., and E. Geraert. 1986. Review of the genus Filenchus Andrássy, 1954 and descriptions of six new species (Nemata: Tylenchidae). Nematologica 32:265-311. (Synonymy of Ottolenchus, Dactylotylenchus,
Lambertia, Duosulcius, Zanenchus and Discotylenchus with Filenchus)

Malenchus Andrássy, 1968
Andrássy, I. 1981. The genera and species of the family Tylenchidae Oerley, 1880 (Nematoda). The genus Malenchus Andrássy, 1968. ). Acta Zoologica Academiae Scientiarum Hungaricae 27:1-47.

Geraert, E., and D. J. Raski. 1986. Unusual Malenchus species (Nemata:Tylenchidae). Nematologica 32:27-55.

Boleodorinae Khan, 1964
Basiria Siddiqi, 1959
Karegar, A., and E. Geraert. 1997. The genus Basiria Siddiqi, 1959 (Nematoda:Tylenchida). II. Species with four lateral lines and anterior median bulb. Nematologica 43:383-406.

Karegar, A., and E. Geraert. 1998. The genus Basiria Siddiqi, 1959 (Nemata:Tylenchida) IV. General discussion, genus diagnosis and key to the species. Nematologica 44:1-13.

Boleodorus Throne, 1941
Khan, E. 1963. Boleodorus mirus n. sp. (Tylenchida: Boleodorinae n. subfam.) from Kufri, Simla (H. P.) India, with a key to the species of the genus Boleodorus Thorne, 1941. Zool. Anz. 173:336-341.

Geraert, E. 1971. Observations on the genera Boleodorus and Boleodoroides (Nematoda: Tylenchidae). Nematologica 17:263-276.

Brzeski, M. W., and M. R. Sauer. 1982. Scanning electron micrography of some Tylenchidae and Boleodoridae and reappraisal of the Boleodoridae. Nematologica 28:437-446.

(The classification for this group is presently very confused.)

Psilenchus de Man, 1921.
Kheiri, A. 1970. Two new species in the family Tylenchidae (Nematoda) from Iran, with a key to Psilenchus De Man, 1921. Nematologica 16:359-368.

Brzeski, M. W. 1989. Notes on the genus Psilenchus de Man, 1921, with description of P. klingleri sp. n. (Nematoda: Tylenchidae). Annales Zoologici 43:51-69.

Neopsilenchus Throne and Malek, 1968
Khan, F. A., and A. M. Khan. 1975. Three new species of Neopsilenchus Thorne & Malek, 1968 (Nematoda: Psilenchinae). Indian Journal of Nematology 5:15-21.

Sultan, M. S., I. Singh, P K. Sakhuja. 1987. Plant parasitic nematodes of the Punjab, II. Neopsilenchus longicaudatus n. sp. (Nematoda: Tylenchidae). Indian Journal of Nematology 17:330-332.

Shahina, F., and Maqbool, M. A. 1990. Studies on the genus Neopsilenchus (Nematoda: Tylenchidae) with description of Acusilenchus n. subgen and three new species. Pakistan Journal of Nematology. 8:49-63.

Atylenchinae Skarbilovich, 1959
Aglenchus Andrássy, 1954
Andrássy, I. 1980. The genera and species of the family Tylenchidae Oerley, 1880 (Nematoda). Acta Zoologica Academiae Scientiarum Hungaricae 26:1-20.

Coslenchus Siddiqi, 1978
Brzeski, M. W. 1978. Taxonomic notes on Coslenchus Siddiqi, 1978 (Nematoda: Tylenchidae). Annales Zoologici 28:417-436.

Siddiqi, M. R. 1980. Six new species of Coslenchus Siddiqi, 1978 (Nematoda: Tylenchidae). Nematologica 26:432-447.

Andrássy, I. 1982. The genera and species of the family Tylenchidae Orley, 1880 (Nematoda). The genus Coslenchus Siddiqi, 1978. Acta Zoologica Academiae Scientiarum Hungaricae 28:193-232.

Geraert, E., and D. J. Raski. 1988. Study of some Aglenchus and Coslenchus species (Nematoda:Tylenchida). Nematologica 34:6-46.

Tylodorinae Paramonov, 1967
Cephalenchus (J. B. Goodey, 1962) Golden, 1971
Sultan, M. S., and M. S. Jairajpuri. 1981. Two new species of the genus Cephalenchus (Goodey, 1962) Golden, 1971 with a key to species. Indian Journal of Nematology 11:165-171.

Mizukubo, T., and N. Minagawa. 1985. Taxonomic study of the genus Cephalenchus (Nematoda:Tylenchida) from Japan. Descriptions of three new species and records of C. planus Siddiqui & Khan with a key to species. Japanese Journal of Nematology 15:26-40.

Raski, D. J., and E. Geraert. 1986. Descriptions of two new species and other observations on the genus Cephalenchus Goodey, 1962 (Nemata: Tylenchida). Nematologica 32:56-78.

Eutylenchus Cobb, 1913
Brzeski, M. W. 1996. On the genus Eutylenchus Cobb, 1913 (Nematoda: Tylenchidae). Nematologica 42:1-8.

Anguinidae Nicoll, 1935 (1926)
Anguina Scopoli, 1777 (after Chizhov and Subbotin, 1990) and related genera
Note: Classification and identification of species of this genus are difficult and controversial.
Chizhov, V. N., and S. A. Subbotin. 1990. Phytoparasitic nematodes of subfamily Anguininae (Nematoda,Tylenchida). Morphology, trophic specialization, taxa. Zoologicheskii Zhurnal 69:15-26. (In Russian).

Ditylenchus Filip'ev, 1936
Brzeski, M. W. 1991. Review of the genus Ditylenchus Filipjev, 1936 (Nematoda: Anguinidae). Revue de Nématologie 14:9-59.

Dolichodoridae Chitwood, 1950
Dolichodorus Cobb, 1914
Loof, P. A. A. A., and R. D. Sharma. 1975. Dolichodorus minor n. sp. (Nematoda: Dolichodoridae) with a key to the genus Dolichodorus. Revista Theobroma (Brazil) 5:35-41.

Lewis, S. A., and A. M. Golden. 1981. Description and SEM observation of Dolichodorus marylandicus n. sp. with a key to species of Dolichodorus. Journal of Nematology 13:128-134.

Smart, G. C., Jr., N. B. Khuong. 1985. Dolichodorus miradvulvus n. sp. (Nematoda: Tylenchida) with a key to species. Journal of Nematology 17:29-37.

Belonolaimidae Whitehead, 1960
Belonolaiminae Whitehead, 1960
Belonolaimus Steiner, 1949
Rau, G. J. 1963. Three new species of Belonolaimus (Nematoda: Tylenchida) with additional data on B. Longicaudatus and B. gracilis. Proceedings of the Helminthological Society of Washington 30:119-128.

Geocenamus Thorne & Malek, 1968
Brzeski, M. W. 1991. Taxonomy of Geocenamus Thorne & Malek, 1968 (Nematoda: Belonolaimidae). Nematologica 37:125-173.

Brzeski, M. W. 1992. Species omitted in Geocenamus review. Nematologica 38:255 (Short communication).

Telotylenchinae Siddiqi, 1960
Amplimerlinius Siddiqi, 1976
Brzeski, M. W. 1985. Contribution to the knowledge of Polish plant parasitic nematodes (Nematoda). 7. Ten species new for Polish fauna. Fragmenta Faunistica 29:275-288.

Shaw, S. P., and E. Khan. 1992. Tylenchorhynchidae Golden, 1971 (Nematoda: Tylenchida) in India: 1. Amplimerlinius sikkimensis sp. n. from Sikkim with key to Amplimerlinius species and Tylenchorhynchus shivanandi sp. n. from Nagaland. Bulletin of Entomology, New Delhi 33:7-13.

Paratrophurus Arias, 1970 (Histotylenchus Siddiqi, 1971 and Telotylenchoides Siddiqi, 1971 are synonymous with Paratrophurus, according to Ebsary, 1991)

Trophurus Loof, 1956
Kleynhans, K. P., and P. Cadet. 1994. Trophurus deboeri n. sp. from sugarcane soil in Barbados and key to the species of the genus Trophurus Loof, 1956 (Nematoda: Belonolaimidae). Fundamental and Applied Nematology 17:225-230. Telotylenchus Siddiqi, 1960

Tylenchorhynchus Cobb, 1913 and related forms
Tarjan, A. C. 1973. A synopsis of the genera and species in the Tylenchorhynchinae (Tylenchoidea, Nematoda). Proceedings of the Helminthological Society of Washington 40:123-144.

Jairajpuri, M. S., and D. J. Hunt. 1984. The taxonomy of Tylenchorhynchinae (Nematoda: Tylenchida) with longitudinal lines and ridges. Systematic Parasitology 6:261-268. (Keys to Dolichorhynchus, Neodolichorhynchus, Tessellus, and to genera of subfamily)

Fortuner, R., and M. Luc. 1987. A reappraisal of Tylenchina (Nemata). 6. The family Belonolaimidae Whitehead, 1960. Revue de Nématologie 10:183-202. (Five genera synonymized with Tylenchorhynchus)

Brzeski, M. W., and C. M. Dolinski. 1998. Compendium of the genus Tylenchorhynchus Cobb, 1913 sensu lato (Nematoda: Belonolaimidae). Russian Journal of Nematology 6:189-199. (Synonymy of six genera with
Tylenchorhynchus)

Handoo, Z. A. 2000. A Key and Diagnostic Compendium to the Species of the Genus Tylenchorhynchus Cobb, 1913 (Nematoda: Belonolaimidae). Journal of Nematology 32(1):20-34.

Bitylenchus Filip'ev, 1934
Jairajpuri, M. S. 1982. Some studies on Tylenchorhynchinae: The subgenus Bitylenchus Filipjev, 1934, with description of Tylenchorhynchus (Bitylenchus) depressus n. sp. and a key to species of Bitylenchus. Mededlingen van de Faculteit Landbouwwetenschappen, Rijksuniversiteit Gent 47:765-770.

Golden, A. M., M. A. Maqbool, and Z. A. Handoo. 1987. Descriptions of two new species of Tylenchorhynchus Cobb, 1913 (Nematoda:Tylenchida), with details on morphology and variation in T. claytoni. Journal of Nematology 19:58-68. (In this manuscript Bitylenchus is synonymized with Tylenchorhynchus.)

Gomez-Barcina, A., M. R. Siddiqi, and P. Castillo. 1992. The genus Bitylenchus Filip'ev, 1934 (Nematoda: Tylenchidae) with descriptions of two new species from Spain. Journal of the Helminthological Society of Washington 59:96-110. (Bitylenchus is revivified.)

Merlinius Siddiqi, 1970
Sultan, M. S., I. Singh, and P. K. Sakhuja. 1988. Plant parasitic nematodes of Punjab, India V. Merlinius communica n. sp. with key to species of the genus and Telotylenchus elongatus n. sp. Indian Journal of Nematology 18:44-48.

Sultan, M. S., I. Singh, and P. K. Sakhuja. 1988. Addendum to the paper "Plant parasitic nematodes of Punjab, India. V. Merlinius communica n. sp. with a key to the species of the genus and Telotylenchus elongatus n. sp. Indian Journal of Nematology 18:44-48.

Nagelus Malek & Thorne, 1968
Powers, T. O., J. G. Baldwin, and A. H. Bell. 1983. Taxonomic limits of the genus Nagelus (Throne and Malek, 1968) Siddiqi, 1979 with a description of Nagelus borealis n. sp. from Alaska. Journal of Nematology 15:582-593.
(Tabular information)

Quinisulcius Siddiqi, 1971
Maqbool, M. A. 1992. Description of Quinisulcius solani n. sp. (Nematoda: Tylenchorhynchidae) with a key to the species and data on Scutylenchus koreanus from Pakistan. Journal of Nematology 14:221-225.

Hoplolaimidae Filip'ev, 1934
Hoplolaiminae Filip'ev, 1934
Aorolaimus Sher, 1963
Baujard, P., P. Castillo, M. E. Doucet, B. Martiny, and D. Mounport. 1994. Taxonomic studies on the genus Aorolaimus Sher, 1963 (Nemata: Hoplolaimidae) I. Bibliographic analysis and tentative key to species. Fundamental and Applied Nematology 17:103-115.

Helicotylenchus Steiner, 1945
Sher, S. A. 1966. Revision of the Hoplolaiminae (Nematoda). VI. Helicotylenchus (Steiner, 1945). Nematologica 12: 1-56.

Anderson, R. V. 1979. A supplemental key to species of Helicotylenchus Steiner, 1945 (Nematoda: Hoplolaimidae) described since 1972 and a description of H. oscephalus n. sp. Canadian Journal of Zoology 57:337-342.

Anderson, R. V., and E. S. Eveleigh. 1982. Description of Helicotylenchus amplius n. sp. and a key to the Canadian species of the genus (Nematoda: Hoplolaimidae). Canadian Journal of Zoology 60:318-321.

Boag, B., and M. S. Jairajpuri. 1985. Helicotylenchus scoticus n. sp. and a conspectus of the genus Helicotylenchus Steiner, 1945 (Tylenchida: Nematoda). Systematic Parasitology 7:47-58. (Compendium of 154 species)

Fotedar, D. N. and V. Kaul. 1985. A revised key to the species of the genus Helicotylenchus Steiner, 1945 (Nematoda: Rotylenchoidinae). Indian Journal of Nematology 15:138-147. (Approximately 125 species in this key)

Firoza, K., and M. A. Maqbool, M. A. 1994. A diagnostic compendium of the genus Helicotylenchus Steiner, 1945 (Nematoda: Hoplolaimidae). Pakistan Journal of Nematology 12:11-50.

Rotylenchus Filip'ev, 1936
Sher, S. A. 1965. Revision of the Hoplolaiminae (Nematoda) V. Rotylenchus Filipjev, 1936. Nematologica 11:173-198.

Boag, B., and D. J. Hooper. 1981. Rotylenchus ouensensis n. sp. (Nematoda: Hoplolaimidae) from the British Isles. Systematic Parasitology 3:119-125.

Geraert, E., and S. Barooti. 1996. Four Rotylenchus from Iran, with a key to the species. Nematologica 42:503-520.

Castillo, P., N. Vovlas, A. Gomez-Barcina and F. Lamberti. 1993. The plant parasitic nematode Rotylenchus, (a monograph). Supplement to Nematologia Mediterranea 21. 200 p.

Hoplolaimus Daday, 1905
Sher, S. A. 1963. Revision of the Hoplolaiminae (Nematoda). II. Hoplolaimus Daday, 1905 and Aorolaimus n. gen. Nematologica 9:267-295.

Jairajpuri, M. S., and Q. H. Baqri. 1973. Nematodes of high altitudes in India. I. four new species of Tylenchida. Nematologica 19:19-30.

Anderson, R. V. 1983. Morphological characteristics of Hoplolaimus indicus Sher, 1963 in Canada, a parasite of wild rice. Journal of Nematology 15:366-369.

Handoo, Z. A., and A. M. Golden. 1992. A key and diagnostic compendium to the species of the genus Hoplolaimus Daday, 1905 (Nematoda: Hoplolaimidae). Journal of Nematology 15:45-53.

Scutellonema Andrássy, 1958
Sher, S. A. 1963. Revision of the Hoplolaiminae (Nematoda) III. Scutellonema Andrássy, 1958. Nematologica 9:421-443.

Van Den Berg, E., and J. Heyns. 1973. South African Hoplolaiminae 2. The genus Scutellonema Andrassy, 1958. Phytophylactica 5:23-40.

Germani, G., J. G. Baldwin, A. H. Bell, X. Y. Wu. 1986. Revision of the genus Scutellonema Andrássy, 1958 (Nematoda: Tylenchida). Revue de Nématologie 8:289-320.

Peltamigratus Sher, 1964
Mulk, M. M., and M. R. Siddiqi. 1982. Three new species of Hoplolaimid nematodes from South America. Indian Journal of Nematology 12:124-131.

Bitencourt , C., and C. S. Huang. 1986. Brazilian Peltamigratus Sher, 1964 (Nematoda: Hoplolaimidae), with description of six new species. Revue de Nématologie 9:3-24.

Rashid, F, E. Geraert , and R. D. Sharma. 1987. The genus Peltamigratus Sher, 1964 with description of two new species (Nematoda: Hoplolaimidae). Revue de Nématologie 10:3-21.

Fortuner, R. 1987. A reappraisal of Tylenchina (Nemata). 8. The family Hoplolaimidae Filip'ev, 1934. Revue de Nématologie 10:219-232. (Peltamigratus was placed in synonymy with Aorolaimus.)

Pararotylenchus Baldwin and Bell, 1981
Baldwin and Bell, A. H. 1981. Pararotylenchus n. gen. (Pararotylenchinae n. subfam.: Hoplolaimidae) with six new species and two new combinations. Journal of Nematology 13:111-128.

Rotylenchulinae Husain & Khan, 1967
Rotylenchulus Linford & Oliverra, 1940
Robinson, A. F., R. N. Inserra, E. P. Caswell-Chen, N. Vovlas, and A. Troccoli. 1997. Rotylenchulus species: identification, distribution, host ranges, and resistance. Nematropica 27:128-180. (Useful descriptions - not a key. See Nickle, Manual of Agricultural Nematology).

Pratylenchidae Thorne, 1949
Pratylenchinae Thorne, 1949
Pratylenchus Filip'ev, 1936
Thorne, G. 1961. Principles of Nematology. New York: McGraw-Hill. (Still a good reference for becoming acquainted with the primary identifying features)

Loof, P. A. A. A., 1978. The genus Pratylenchus Filipjev, 1936 (Nematoda: Pratylenchidae): a review of its anatomy, morphology, distribution, systematics and identification. Vaxtskyddsrapporter 5:1-50. (Outstanding key)

Ryss, A. Yu. 1988. Kornevye paraziticheskie nematody semeistva Pratylenchidae (Tylenchida) mirovoi fauny. Lenningrad, Nauka. (Russian text)). (See software)

Frederick, J. J., and A. C. Tarjan. 1989. A compendium of the genus Pratylenchus Filipjev, 1936 (Nemata: Pratylenchidae). Revue de Nématologie 12:243-256.

Handoo, Z. A., and A. M. Golden. 1989. A key and diagnostic compendium to the genus Pratylenchus Filipjev, 1936 (lesion nematodes). Journal of Nematology 21:202-218.

Café Filho, A. C., and C. S. Huang. 1989. Description of Pratylenchus pseudofallax n. sp. with a key to the species of the genus Pratylenchus Filipjev, 1936 (Nematoda: Pratylenchidae). Revue de Nématologie 12:7-15.

Radopholus Thorne, 1949
Sher, S. A. 1968. Revision of the genus Radopholus Thorne, 1949 (Nematoda: Tylenchidae). Proceedings of the Helminthological Society of Washington 35:219 237.

Hoplotylus s'Jacob, 1960
Bernard, E. C., and T. L. Niblack. 1982. Review of Hoplotylus s'Jacob (Nematoda: Pratylenchidae). Nematologica 28:101-109.

Hirschmaniella Luc and Goodey, 1964
Ebsary, B. A., and R. V. Anderson. 1982. Two new species of Hirschmaniella Luc and Goody, 1963 (Nematoda: Pratylenchidae) with a key to the nominal species. Canadian Journal of Zoology 60:530-534.

Sivakumar, C. V., and E. Khan. 1982. Description of Hirschmaniella kaverni sp. n. (Radopholidae: Nematoda) with a key for identification of Hirschmaniella spp. Indian Journal of Nematology 12:86-90.

Pratylenchoides Winslow, 1958
Ryss, A. Y. 1980. Pratylenchoides ivanovae sp. n. (Nematoda: Pratylenchidae) and key to species of the genus Pratylenchoides. Parazitologiya 14:516-520.

Baldwin, J. G., et al. 1983. Contribution to the study of the genus Pratylenchoides Winslow (Nematoda: Tylenchidae). Revue de Nématologie 6:111-125.

Nacobbinae Chitwood, 1950
Nacobbus Thorne and Allen, 1944
Sher, S. A. 1970. Revision of the genus Nacobbus Thorne and Allen, 1944 (Nematoda: Tylenchoidea). Journal of Nematology 2:228-235.

Heteroderidae Filip'ev & Schuurmans Stekhoven, 1941 Mulvey, R. H., and A. M. Golden. 1983. An illustrated key to the cyst-forming genera and species of Heteroderidae in the western hemisphere with species morphometrics and distribution. Journal of Nematology 15(1): 1-59.

Wouts, W. M. 1985. Phylogenetic classification of the family Heteroderidae (Nematoda: Tylenchida). Systematic Parasitology 7:295-328.

Luc, M., A. Maggenti, and R. Fortuner. 1988. A reappraisal of Tylenchina (Nemata). 9. The family Heteroderidae Filip'ev & Schuurmans Stekhoven, 1941. Revue de Nématologie 11:159-176.

Heteroderinae Filip'ev & Schuurmans Stekhoven, 1941 Baldwin, J. G., and L. P. Schouest, Jr. 1990. Comparative detailed morphology of the Heteroderinae Filip'ev & Schuurmans Stekhoven, 1941, sensu Luc et al.
(1988): Phylogenetic systematics and revised classification. Systematic Parasitology 15:81-106.

(Also, see key to genera of Heteroderinae by Baldwin and Mundo-Ocampo on Pp. 345-346 of Manual of Agricultural Nematology)

Cryphodera Colbran, 1966
Karssen, G., and A. Van Aelst. 1999. Description of Cryphodera brinkmani n. sp. (Nematoda: Heteroderidae), a parasite of Pinus thunbergii Parlatore from Japan, including a key to the species of Cryphodera Colbran, 1966. Nematology 1:121-130.

Globodera Skarbilovich, 1959
Wouts, W. M. 1984. Globodera zelandica n. sp. (Nematoda: Heteroderidae) from New Zealand, with a key to the species of Globodera. New Zealand Journal of Zoology 11:129-135.

Heterodera Schmidt, 1871
Mulvey, R. H. 1972. Identification of Heterodera cysts by terminal and cone top structures. Canadian Journal of Zoology 50:1277-1292. (Includes good photographic plates)

Golden, A. M. 1986. Morphology and identification of cyst nematodes. Pp. 23-45 in F. Lamberti and C. E. Taylor, eds. Cyst nematodes. NATO ASI Series. Series A: Life Sciences Vol. 121. New York: Plenum Press. [Includes key to 6 genera and 59 species of cyst-forming nematodes (Heteroderidae)].

Wouts, W. M., A. Schoemaker, D. Sturhan, and P. R. Burrows. 1995. Heterodera spinicauda sp. n. (Nematoda: Heteroderidae) from mud flats in the Netherlands, with a key to the species of the H. avenae group. Nematologica 41:575-583.

Meloidogyninae Skarbilovich, 1959
Meloidogyne Goeldi, 1889
Taylor, A. L., and J. N. Sasser. 1978. Biology, identification and control of root- knot nematodes (Meloidogyne species). Raleigh: North Carolina State University Graphics.

Eisenback, J. D., H. Hirschmann, J. N. Sasser, and A. C. Triantaphyllou. 1981. A guide to the four most common species of root-knot nematodes of root-knot nematodes (Meloidogyne spp.), with a pictorial key. Raleigh: North Carolina State University.

Ebsary, B. A., and E. S. Eveleigh. 1983. Meloidogyne aquatilis n. sp. (Nematoda: Meloidogynidae) from Spartina pectinata with a key to the Canadian species of Meloidogyne. Journal of Nematology 15:349-353.

Hewlett, T. E., and A. C. Tarjan. 1983. Synopsis of the genus Meloidogyne Goeldi, 1887. Nematropica 13:79-102.

Jepson, S. B. 1987. Identification of root-knot nematodes (Meloidogyne species). Wallingford, UK: CAB International. 265 p.

Eisenback, J. D., and H. H. Triantaphyllou. 1991. Root-knot nematodes: Meloidogyne species and races. Pp. 191-274 in W. R. Nickel, ed. Manual of agricultural nematology. New York: Marcel Dekker.

Criconematoidea Taylor, 1936
Criconematidae (Taylor, 1936) Thorne, 1949
Criconematinae Taylor, 1936
Bakernema Wu, 1964
Ebsary, B. A. 1982. Bakernema yukonense n. sp. (Nematoda: Criconematidae) with keys to the species of Criconemella and Discocriconemella. Canadian Journal of Zoology 60:3033-3047.

Criconema Hofmanner & Menzel, 1914
Golden, A., and W. Friedman. 1964. Some taxonomic studies on the genus Criconema (Nematoda: Criconematidae). Proceedings of the Helminthological Society of Washington 31:47-59.

Yeates, G. W., P. A. A. A. Loof, and W. M. Wouts. 1997. Criconematidae (Nematoda: Tylenchida) from the New Zealand region: analysis of and key to Criconema (Nothocriconemella) species. New Zealand Journal of Zoology 24:153-162.

Criconemella (Mesocriconema) de Grisse & Loof, 1965 and related genera
Raski, D. J. 1952. On the morphology of Criconemoides Taylor, 1936, with descriptions of six new species (Nematoda: Criconematidae). Proceedings Helminthological Society of Washington 19:85-99.

De Grisse, A., and P. A. A. A. Loof. 1965. Revision of the genus Criconemoides (Nematoda). Mededelingen Landbouwhogeschool en Opzoekingsstations van de Staat te Gent 30:577-603. (Landmark work, keys to species of the various Criconemoides segregates)

Tarjan, A. C. 1966. A compendium of the genus Criconemoides (Criconematidae: Nemata). Proceedings of the Helminthological Society of Washington 33:109-125.

Raski, D. J., and A. M. Golden. 1966. Studies on the genus Criconemoides Taylor, 1936 with descriptions of eleven new species and Bakernema variabile n. sp. (Criconematidae: Nematoda). Nematologica 11:501-565.

Mehta, U. K., and D. J. Raski. 1971. Revision of the genus Criconema Hofmanner and Menzel, 1914 and other related genera (Criconematidae: Nematoda). Indian Journal of Nematology 1:145-198. (Important manuscript includes Ogma, Crossonema and others).

Andrássy, I. 1979. Revision of the subfamily Criconematinae Taylor, 1936 (Nematoda). Opuscula Zoologica (Budapest) 16:11-57. (Many keys to genera and species)

Ebsary, B. A. 1979. Description of Criconemoides pleriannulatus n. sp. (Nematoda: Criconematidae) and a key to species of Criconemoides. Canadian Journal of Zoology 57:1-5.

Luc, M., and D. J. Raski. 1981. Status of the genera Macroposthonia, Criconemoides, Criconemella, and Xenocriconemella (Criconematidae: Nematoda). Revue de Nématologie 4:3-21.

Hashim, Z. 1984. Re-diagnosis and a key to the species of Neolobocriconema Mehta & Raski, 1971 (Nematoda: Tylenchida), with a description of N. orlarum n. sp. from Jordan. Systematic Parasitology 6:69-73.

Loof, P. A. A. A., and De Grisse. 1989. Taxonomic and nomenclatorial observation of the genus Criconemella De Grisse & Loof, 1965 sensu Luc & Raski (Criconematidae). Meded. Fac. Landbouww. Rij. Gent 54:53-74.

Ogma Southern, 1914
Minagwa, N. 1993. Taxonomic studies of Criconematidae (Nematoda: Tylenchida) of Japan IV Ogma: part 2. Bulletin National Agro. Environmental Science 9:53-152.

Van Den Berg, E., and P. Queneherve. 1995. Ogma toparti sp.n. and two known Criconematoidea from the French Caribbean (Nemata: Tylenchina). Fundamental and Applied Nematology 18:361-369.

Hemicriconemoides Chitwood & Birchfield, 1957
Dasgupta, D. R., D. J. Raski, and S. D. Van Gundy. 1969. Revision of the genus
Hemicriconemoides Chitwood & Birchfield, 1957 (Nematoda: Criconematidae). Journal of Nematology 1: 126-145.

Germani, G., and M. Luc. 1970. Contribution a l'etude du genre Hemicriconemoides Chitwood & Birchfield, 1957 (Nematoda: Criconematidae). Cahiers ORSTOM Serie Physiologie des Plantes Tropicales Cultivees 11:133-150.

Esser, P. R. and Volvas, N. 1990. A diagnostic compendium to the genus Hemicriconemoides (Tylenchida: Criconematidae). Proceedings of the Soil and Crop Science Society of Florida 49:211-219.

Hemicycliophorinae Skarbilovich, 1959
Hemicycliophora de Man, 1921
Schoemaker, R. L. P. 1967. Hemicycliophora nyanzae n. sp. found in East Africa with a key to the species of Hemicycliophora De Man, 1921. Nematologica 3:541-546.

Loof, P. A. A. 1968. Taxonomy of Hemicycliophora species from west and central Europe (Nematoda: Criconematoidea). Mededlingen Landbouwhogeschool Wageningen 68-14:1-43.

Brzeski, M. W. 1974. Taxonomy of Hemicycliophorinae (Nematoda, Tylenchida). Zeszyty Problemowe Postepow Nauk Rolniczych 154:237-330.

Loof, P. A. A. A. 1976. The genera Hemicycliophora De Man, 1921 and Caloosia Siddiqi & Goodey, 1963 (Criconematoidea). Mededelingen van de Faculteit Landbouwwetenschappen Rijksuniversiteit Gent 41:1023-1029.

Eroshenko, A. 1976. Paraziticheski kornevye nematody. Semeistvo Hemicycliophoridae, Leningrad: Nauka Leningradskoe otd-nie. 79p.

Brzeski, M. W., and T. S. Ivanova. 1978. Taxonomic notes on Hemicycliophora de Man (Nematoda: Hemicycliophoridae). Nematologia Mediterranea 6:147-162.

Siddiqi, M. R. 1980. Taxonomy of the plant nematode superfamily Hemicycliophoroidea, with a proposal for Criconematina, a new suborder. Revue de Nématologie 3:179-199.

Van Den Berg, E. 1987. Hemicycliophora species from the western Cape Province with a key to the South African species (Hemicycliophoridae: Nematoda). Phytophylactica 19:303-307.

Tylenchulidae Skarbilovich, 1947
Tylenchulinae Skarbilovich, 1947
Tylenchulus Cobb, 1913
Inserra, R. N., N. Vovlas, J. H. O'Bannon, and R. P. Esser. 1988. Tylenchulus graminis n. sp. and T. palustris n. sp. (Tylenchulidae), from native flora of Florida, with notes on T. semipenetrans and T. furcus. Journal of Nematology 20: 266-287.

Paratylenchinae Throne, 1949
Paratylenchus Micoletzky, 1922 and Gracilacus Raski, 1962
Geraert, E. 1965. The genus Paratylenchus. Nematologica 11:301-334. (Old but contains lots of good information and comparative drawings)

Solov'eva, G. I. 1972. Parasitic nematodes of woody and herbaceous plants. Akadeiya Nauk SSSR (Lenningrad, Russia: Nauka Publishers).

Raski, D. J. 1975. Revision of the genus Paratylenchus Micoletzky, 1922 and descriptions of new species. Part I of 3 parts. Journal of Nematology 7:15-34.

Raski, D. J. 1975. Revision of the genus Paratylenchus Micoletzky, 1922 and descriptions of new species. Part II of three parts. Journal of Nematology 7:274-295.

Esser, R. P. 1992. A diagnostic compendium to species included in Paratylenchinae Thorne, 1949 and Tylenchocriconematinae Raski & Siddiqui, 1975 (Nematoda: Criconematoidea). Nematologica 38: 146-163.

Gracilacus Raski, 1962
Raski, D. J. 1975. Revision of the genus Paratylenchus Micoletzky, 1922 and descriptions of new species. Part III of three parts-Gracilacus. Journal of Nematology 8: 97-115.

Triplonchida Cobb, 1920
Diphtherophorina Coomans and Loof, 1970
Trichodoroidea Thorne, 1935
Trichodoridae Thorne, 1935 (Siddiqi, 1961)
Trichodorus Cobb, 1913 and related genera
Esser, R. P. 1971. A compendium of the genus Trichodorus (Dorylaimoidea: Diphtherophoridae). Soil and Crop Science 31:244-253.

Siddiqi, M. R. 1973. Systematics of the genus Trichodorus Cobb, 1913 (Nematoda: Dorylaimida), with descriptions of three new species. Nematologica 19:259-278. (A key to the subgenera and species of Paratrichodorus is given)

Decraemer, W. 1980. Systematics of the Trichodoridae (Nematoda) with keys to their species. Revue de Nématologie 3:81-99.

Rashid, F., D. de Waele, and A. Coomans. 1985. Trichodoridae from Brazil. Nematologica 31:289-320. (Emended diagnosis and key for Allotrichodorus)

Waele, D. de, and M. W. Brzeski. 1995. Trichodorus altaicus sp. n. (Nematoda: Trichodoridae) and a key to the species of the genus Trichodorus. Fundamental and Applied Nematology 18:181-187.

Decraemer, W. 1995. The family Trichodoridae: Stubby root and virus vector nematodes. Dordrecht, The Netherlands: Kluwer Academic Publishers. 360 p.

Taylor, C. E., and D. J. F. Brown. 1997. Nematode vectors of plant viruses. Wallingford, UK: CAB International. 286 p.

Decraemer, W., and P. Baujard. 1998. A polytomous key for the identification of species of the family Trichodoridae Thorne, 1935 (Nematoda: Triplonchida). Fundamental and Applied Nematology 21:37-62.

Decraemer, W. & Baujard, P. (1998). Additions and corrections to: Decraemer: the family Trichodridae. Fundamental and Applied Nematology, 21 (2): 207-212.

Dorylaimida Pearse, 1942
Longidoroidea Thorne, 1935
Longidoridae Thorne, 1935
Longidorinae Thorne, 1935
Longidorus Micoletzky, 1922 (Filip'ev, 1934; Thorne, 1934) and Paralongidorus Siddiqi, Hooper & Khan, 1963
Romanenko, N. D. 1978. A polytomic key for the identification of species of Longidorus Micoletzky, 1922. Fitogel'mintologicheskie issledovaniya. Moscow, "Nauka." 111-114.

Chen, Q. W., D. J. Hooper, P. A. A. Loof, and J. H. Xu. 1997. A revised polytomous key for the identification of species of the genus Longidorus Micoletzky, 1922 (Nematoda: Dorylaimoidea). Fundamental and Applied
Nematology 20:15-28.

Escuer, M., and M. Arias. 1997. Paralongidorus iberis sp. n. from Spain with a polytomous key to the species of the genus Paralongidorus Siddiqi, Hooper & Khan, 1963 (Nematoda: Longidoridae). Fundamental and Applied Nematology 20:135-148.

Taylor, C. E., and D. J. F. Brown. 1997. Nematode vectors of plant viruses. Wallingford, UK: CAB International. 286 p. (A polytomous key [= compendium = tabular key] is included for species of Longidorus and Paralongidorus. See the
authors' discussion of dichotomous and polytomous keys on pp. 58-60.

Loof, P. A. A., and Q. W. Chen. 1999. A revised polytomous key for the identification of species of the genus Longidorus Micoletzky, 1922 (Nematoda: Dorylaimoidea). Supplement I. Nematology 1:55-59.

Xiphinematidae Dalmasso, 1969
Xiphinematinae Dalmasso, 1969
Xiphinema Cobb, 1913
Lamberti, F., and T. Beleve-Zacheo. 1979. Studies on Xiphinema americanum sensu lato with descriptions of fifteen new species (Nematoda, Longidoridae). Nematologia Mediterranea 7:51-106.

Loof, P. A. A., and M. Luc. 1990. A revised polytomous key for the identification of species of the genus Xiphinema Cobb, 1913 (Nematoda: Longidoridae) with exclusion of the X. americanum-group). Systematic Parasitology 16:35-66.

Lamberti, F., and M. Carone. 1991. A dichotomous key for the identification of species of Xiphinema (Nematoda: Dorylaimida) with the X. americanum-group. Nematologia Mediterranea 19:341-348.

Loof, P. A. A., and M. Luc. 1993. A revised polytomous key for the identification of species of the genus Xiphinema Cobb, 1913 (Nematoda: Longidoridae) with exclusion of the X. americanum-group: Supplement I. Systematic Parasitology 24:185-189.

Loof, P. A. A., M. Luc, and P. Baujard. 1996. A revised polytomous key for the identification of species of the genus Xiphinema Cobb, 1913 (Nematoda: Longidoridae) with exclusion of the X. americanum-group: Supplement 2.
Systematic Parasitology 33:23-29.

Robbins, R. T., D. J. F. Brown, J. M. Halbrendt, and T. C. Vrain. 1996. Compendium of juvenile stages of Xiphinema species (Nematoda: Longidoridae). Russian Journal of Nematology 4:163-171.

 

Aphelenchida Siddiqi, 1980
Aphelenchina Geraert, 1966
Aphelenchoidoidea Skarbilovich, 1947 (Siddiqi, 1980)
Aphelenchoididae Skarbilovich, 1947 (Paramonov, 1953)
Aphelenchoidinae Skarbilovich, 1947
Aphelenchoides Fischer, 1894
Sanwal, K. C. 1961. A key to the species of the nematode genus Aphelenchoides Fischer, 1894. Canadian Journal of Zoology 39:143-148.

Hunt, D. J. 1993. Aphelenchida, Longidoridae, and Trichodoridae: Their systematics and bionomics. Wallingford, UK: CAB International.

Shahina, F. 1996. A diagnostic compendium of the genus Aphelenchoides Fischer, 1894 (Nematoda: Aphelenchida) with some new records of the group from Pakistan. Pakistan Journal of Nematology 14:1-32.

Aphelenchinae Fuchs, 1937
Aphelenchus Bastian, 1865
Nama, H. S., and Soni, G. R. 1981. Taxonomy of some species of the genus Aphelenchus Bastian, 1865 (Aphelenchoidea) with a key. Proceeding of the Indian Academy of Parasitology 2:107-110.

Anderson, R. V., and D. J. Hooper. 1980. Diagnostic value of vagina structure in the taxonomy of Aphelenchus bastian, 1865 (Nematoda: Aphelenchidae) with a description of A. (Anaphelenchus) isomerus n. subgen., n. sp. Canadian Journal of Zoology 58:924-928.

Bursaphelenchinae Paramonov, 1964
Bursaphelenhus Fuchs, 1937
Tarjan, A. C., and C. B. Aragon. 1982. An analysis of the genus Bursaphelenchus Fuchs, 1937. Nematropica 12:121-144.

Yin, K., Y. Fang, and A. C. Tarjan. 1988. A key to species in the genus Bursaphelenchus with a description of Bursaphelenchus hunanensis sp. n. (Nematoda: Aphelenchoididae) found in pine wood in Hunan Province, China. Proceedings of the Helminthological Society of Washington 55:1-11.

Rhadinaphelenchus J. B. Goodey, 1960
Monotypic: R. cocophilus (Cobb, 1919) J. B. Goodey, 1960

Software - Computerized Keys:

Ditylenchus
Viscardi, T, and M. W. Brzeski. 1993. DITYL: computerized key for species identification of Ditylenchus (Nematoda: Anguinidae). Fundamental and Applied Nematology 16:389-392.

Tylenchorhynchus sensu lato and Geocenamus species Previously available from Dr. Michael W. Brzeski, now deceased.

Helicotylenchus spp.
Fortuner, R., and Y. Wong. 1984. Review of the genus Helicotylenchus Steiner, 1945. 1. A computer program for identification of the species. Revue de Nématologie 7:385-392.

Diederich, J., R. Fortuner, and J. Milton. 1991. Nemisys: Nematode identification system. Pp. 1-42 in R. Fortuner, ed. Advances in computer methods for systematic biology: Artificial intelligence, database systems, computer vision. Baltimore, MD: Johns Hopkins University Press.

Fortuner, R. 1989. A new description of the process of identification of plant-parasitic nematode genera. Pp. 35-44 in R. Fortuner, ed. Nematode identification and expert system technology. New York: Plenum Press.

Pratylenchus and Radopholus
Digital pictorial keys - Dr. Alexander Ryss, Zoological Institute, Russian Academy of Sciences, Universitetskaya, Emb., 1, St. Petersburg 199034, Russia. E-mail: alexryss@AR4280.spb.edu Also, vermes@zin.ru.

Ryss, A. 1997. Computerized identification of species of the genus Radopholus (Tylenchida: Pratylenchidae). Russian Journal of Nematology 5:137-142.

Longidorus
Dr. R. T. Robbins, University of Arkansas, Nematology Laboratory, Fayetteville, Arkansas 72701.

Rey, J. M., M-Fe Andres, and M. Arias. 1988. A computer method for identifying nematode species. 1. Genus Longidorus (Nematoda: Longidoridae). Revue de Nématologie 11:129-135.

Xiphinema
Dr. P. Baujard, Museum National d'Histoire Naturelle, Labratoire de Biologie Parasitaire, Protistologie et Helminthologie, 61, rue Buffon, 75005 Paris, France.

Trichodorus and Paratrichodorus
Dr. W. Decraemer, Department of Invertebrates, Royal Belgian Institute of Natural Sciences, Vautierstraat 29, B-1040 Brussels, Belgium.