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VOL. 34, No. 1 One of the First Obligate Tylenchid Parasites Of 72 PROCEEDINGS OF THE [VOL. 34, No. 1 Thome, 1939 and Hellcotylenchus nanus Stei- RASKI, D. J., AND J. D. RADEWALD. 1958. Repro- ner, 1945. Bull. Md. Agric. Exp. St. A. 106, duction and symptomatology of certain ecto- 16 pp. parasitic nematodes on roots of Thompson PITCHER, R. S., AND A. F. POSNETTE. 1963. Vas- seedless grape. Pi. Dis. Reptr. 42: 941-943. cular feeding by Xiphinema diversicaudatum SCHINDLER, A. F. 1957. Parasitism and patho- (Micol.). Nematologica 9: 301-302. genicity of Xiphinema diversicaudatum, an RADEWALD, J. D. 1962. The biology of Xiphi- eetoparasitic nematode. Nematologica 2: 25- nema index and the pathological effect of the 31. species on grape. Ph.D. Thesis, University of SCHINDLEH, A. F., AND A. J. BRAUN. 1957. California, Davis, California. Pathogenicity of an eetoparasitic nematode, RADEWALD, J. D., AND D. J. RASKI. 1962. A Xiphinema diversicaudatum, on strawberries. study of the life cycle of Xiphinema index. Nematologica 2: 91-93. Phytopathology 52: 748 (abstr.). VAN GUNDY, S. D., AND I. J. THOMASON. 1962. RADEWALD, J. D., AND D. J. RASKI. 1962. Studies Factors influencing storage life of Hemi- on the host range and pathogenicity of Xiphi- cycliophora arenaria, Pratylenchus scribneri nema index. Phytopathology 52: 748-749 and Trichodorus christiei. Phytopathology 52: (abstr.). 366-367. On the Classification of the Insect Parasitic Nematodes of the Sphaerulariidae Luhhock, 1861 (Tylenchoidea: Nematoda) W. R. NlCKLE1 One of the first obligate tylenchid parasites 1963b). Fuchs, Wachek, and Riihm each pub- of insects was described by Dufour in 1837 lished comprehensive works, and did much to as Sphaerularia bombi from the bumblebees, clarify the systematic^ of this family. Bovien's Bom-bus terrestris and Bombus hortorus. In publications on insect nematodes were out- 1861, Lubbock proposed the family Sphaeru- standing and are classics today. lariaceae to contain the genus Sphaerularia. At present, 4 subfamilies, 21 genera, and Other significant early contributions to the 116 species are included in the Sphaerulariidae. taxonomy of these forms include: Leuckart Generally the classification is strong at the (1884), von Linstow (1890), zur Strassen generic level, but at the specific level it is (1892), Cobb (1920, 1921), Micoletzky (1922), rather weak because detailed descriptions of T. Goodey (1930), Filipjev (1934), Thorne free-living males and females are lacking. A (1935), Fuchs (1915, 1929, 1933, 1938), Bovien (1937), Currie (1937), Chitwood and tylenchid species can generally parasitize hosts Chitwood (1937), Christie (1938) and Schnei- in one genus, and unlike the mermithids, der (1939). More recent contributions in- adults are found in adult hosts. Therefore, clude: T. Goodey (1953), Wachek (1955), knowledge of the host aids in identification. Riihm (1956), Massey (1956, 1957, 1958, In several of the nematode genera the dorsal 1960, 1962), J. B. Goodey (1956, 1963), esophageal gland orifice is located at least one Khan (1957a, 1957b, 1960), Skarbilovich (1947, stylet length behind the base of the stylet or 1959), Welch (1959), and Nickle (1963a, could not be found. The subventral esophageal glands may empty into the lumen of the esoph- agus a short distance behind the dorsal gland 1 Research Nematologist, Crops Research Division, Agri- cultural Research Service, U. S. Department of Agriculture, orifice. This led Wachek (1955) to place Beltsville, Maryland. Some studies were made while the author was Research Assistant, University of California, Sphaerularia, Tripius, and Scatonema in the Davis, California, and also Research Officer, Canada De- partment of Agriculture, Belleville, Ontario. Aphelenchoidea, though they lack the typical Copyright © 2011, The Helminthological Society of Washington JANUARY, 1967] HELMINTHOLOGICAL SOCIETY 73 aphelenchoid median bulb and in all other Because of the brevity of the original de- respects appear to be tylenchoid. scription, the family diagnosis is emended. Recently, I observed the dorsal esophageal Brief diagnoses of the sub-families and genera gland orifice in the fourth stage female larva are given and a new genus is established. of Sphaerularia bombi and found that it was Synonymies are presented along with my com- located behind the stylet in the normal tylen- ments on the status of controversial groups. choid position. Further, the presence of tylen- The characters that typify the family in- choid spicules and a gubernaculum in these clude: absence of a valved median bulb; forms also, supports their tylenchoid status. unique ovary of the free-living female; en- For these reasons I question the aphelenchoid largement of female nematode in the body status of these genera, and I now consider cavity of host, resulting in the nematode be- some of the genera placed in the Aphelen- coming a reproductive sac (heteromorphism), choidea as belonging to the Tylenchoidea. or female nematode with a prolapsed swollen The problem inherent in making this change uterus; obligate insect or mite parasitism. is that Sphaerulariaceae Lubbock, 1861 is an FAMILY : older family name than Allantonematidae Sphaerulariidae Lubbock, 1861. (Pereira, 1931) Chitwood and Chitwood, 1937. Syn. Sphaerulariaceae Lubbock, 1861. In my opinion the insect parasitic Tylenchoidea DIAGNOSIS (Emended): Tylenchoidea. Usu- merit only family rank along with the plant ally with three distinct forms. Two free-living, parasitic families; Tylenchiclae, Heteroderidae, slender, 0.5-1.0 mm long, often found in hab- Criconematidae, Neotylenchidae, Hoplolaimi- itat of young host and one adult parasitic form dae, and Tylenchulidae. Future work in the found in haemocoel of insect or mite. (1) group may necessitate the creation of a super- FL$*: Stylet or pseudostylet rarely absent. family but at present, all insect-parasitic Tylen- Esophagus without median valvular bulb; am- choidea should be placed in the family Sphaer- pullae near gland orifices often strongly de- ulariidae Lubbock, 1861; the Allantonematidae veloped; dorsal gland orifice not always found (Pereira, 1931) Chitwood and Chitwood, 1937 immediately behind spear, may be midway should be given the rank of subfamily. between spear and subventral gland orifices; Entaphelenchus Wachek, 1955 and Pera- esophageal glands long, overlap intestine. Go- phelenchus Wachek, 1955 are endoparasitic nad diagnostic; prodelphic; ovary small, finger- tylenchids, having the typical aphelenchoid like with few oocytes; oviduct short; uterus median bulb and spicules which are shaped prominent, temporarily packed with individual like rose thorns. The gubernaculum is lacking, sperm after copulation. Becomes infective and in the writer's opinion, only these two stage. Eggs absent. (2)FL3: Usually slightly genera are typical of the Aphelenchoidea. longer than female, not infective. Stylet may During the last 8 years, I have studied rep- be absent. Esophagus weakly developed. Spic- resentative specimens from 15 of the 22 genera ules usually tylenchoid, may be elaborate. belonging to the families Sphaerulariidae and Gubernaculum usually present. Caudal alae Allantonematidae before making the revisions often present and peloderan. (3) AP?: Ob- contained herein. It is hoped that this com- ligate parasite in haemocoel of insects or mites. pilation will stimulate work in this interesting Large swollen reproductive sac, produced by but neglected group. expansion of free-living female; or, with uterus Economically, insect parasitic nematodes of prolapsed, swollen. Stylet present or rarely the Sphaerulariidae annually reduce insect absent, may be retracted. Esophagus and in- populations by untold millions, and are indeed testine usually degenerate at expense of repro- important self-perpetuating biological control ductive system which often fills 80% of body. agents. However, little effort has been made Oviparous or ovoviviparous; ovary usually in manipulating, or exploiting these parasites flexed one or more times; rachis common. for biological control, but the time is ripe for * FL$ : Free-living female. FLcT: Free-living male. this type of endeavor. AP9= Body cavity adult parasitic female. Copyright © 2011, The Helminthological Society of Washington 74 PROCEEDINGS OF THE [VOL. 34, No. 1 PROPOSED CLASSIFICATION OF THE SPHAERULARIIDAE Sphaerulariidae Lubbock, 1861. Syn. Sphaerulariaceae Lubbock, 1861. Sphaerulariinae (Lubbock, 1861) Pereira, 1931. *Sphaerularia Dufour, 1837. *Sphaerulariopsis Wachek, 1955. *Tripius Chitwood, 1935. Syn. Asconema Leuckart, 1886. Atractonema (Leuckart, 1886) Leuckart, 1887. Proatractonema Bovien, 1944. Allantonematinae Pereira, 1931. (Emended Chitwood, 1935). Syn. Allantoneminae Pereira, 1931. Allantonematidae (Pereira, 1931) Chitwood and Chitwood, 1937. Contortylenchidae Ruhm, 1956. Allantonema Leuckart, 1884. Syn. Tylenchomorphiifi Fuchs, 1915. * Aphelenchulus Cobb, 1920. *Bovienema Nickle, 1963. *Bradynema zur Strassen, 1892. *Chondronema Christie and Chitwood, 1931. *Contortylenchus Ruhm, 1956. *Heterotijlenchus Bovien, 1937. *Howardula Cobb, 1921. Syn. Tylenchinema T. Goocley, 1930. *Prothallonema Christie, 1938. Metaparasitylenchns Wachek, 1955. n. grad. *Neoparasitylenchus n. gen. Parasitylenchoid.es Wachek, 1955. *Parasitylenchus Micoletzky, 1922. Syn. Polymorphotylenchus Riihm, 1956. Proparasitijlenchus Wachek, 1955. n. grad. Protylenchus Wachek, 1955. *Sulphuretylenchiis Riihm, 1956. n. grad. Scatonema Bovien, 1932. Dotylaphus Andrassy, 1958. (genus inquirenda). Fergusobiinae J. B. Goodey, 1963. Syn. Fergusobiidae Siddiqi and J. B. Goodey,
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