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 esophagus 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

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.

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, 1963. *Fergusobia Currie, 1937. Syn. Anguillulina (Fergusobia) Currie, 1937. lotonchiinae T. Goodey, 1953. Syn. lotonchiidae (T. Goodey, 1953) Skarbilovich, 1959. *Iotonchium Cobb, 1920.

* Specimens of these genera were studied for this review.

SUBFAMILY: OTHER SPECIES: Sphaerulariinae (Lubbock, 1861) Pereira, S.dendroctoni (Massey, 1956) Nickle, 1963. 1931. Syn. Sphaerularia dendroctoni Massey, DIAGNOSIS (Emended): Sphaerulariidae. 1956. FL 9 : Stylet present, with or without knobs. S. hastatus (Khan, 1957) Nickle, 1963. Without eggs in uterus. FLc$: With tylen- Syn. Sphaerularia hastata Khan, 1957. choid anterior end. Stylet present, with or Stictylus hastatus (Khan, 1957) without knobs. Caudal alae present or absent. Khan, 1960. Spicules and gubernaculum tylenchoid. AP9: S. pini (Fuchs, 1929) Nickle, 1963. Uterus everted partially or completely. Syn. Tylenchus sulphureus pini Fuchs, Genus Sphaerularia Dufour, 1837 1929. (Figs. 2Z, 3V, 5A) Allantonema sulphureus pini (Fuchs, 1929) Filipjev, 1934. FL 9 : Stylet present, without basal knobs. Parasitylenchus sulphureus f. pini Dorsal gland orifice opens just behind stylet. (Fuchs, 1929) Schneider, 1939. FL $ : Stylet present, not prominent, without Allantonema pini (Fuchs, 1929) knobs. Caudal alae absent. Spicules and Wachek, 1955. gubernaculum tylenehoid. AP9 : Uterus com- Stictylus pini (Fuchs, 1929) Ruhm, pletely everted, greatly enlarged, larger and 1956. not as smooth as Sphaerulariopsis, becomes S. piceae (Fuchs, 1929) n. comb. 15,000-20,000 times the volume of original Syn. Tylenchus sulphureus piceae Fuchs, female body which remains as a minute ap- 1929. pendage to uterus, though often wrinkled and Allantonema sulphureus piceae deformed. Oviparous. (Fuchs, 1929) Filipjev, 1934. HOST INSECTS: Parasitylenchus sulphureus f. piceae Hymenoptera; Bombus, Vespa, "Psithyrus. (Fuchs, 1929) Schneider, 1939. TYPE SPECIES: Stictylus sulphureus piceae (Fuchs, Sphaerularia bombi Dufour, 1837. 1929) Ruhm, 1956. Syn. Tylenchus bombi (Dufour, 1837) Cobb, 1890. S. piniphili (Fuchs, 1929) Nickle, 1963. Syn. Tylenchus sulphureus piniphili Genus: Sphaerulariopsis Wachek, 1955 Fuchs, 1929. (Figs. 2W, 3W, 5A) Allantonema sulphureus piniphili (Fuchs, 1929) Filipjev, 1934. FL 9 : Stylet present, well-developed, with Parasitylenchus sulphureus f. pini- basal knobs which may be irregular in size. phili (Fuchs, 1929) Schneider, Dorsal gland orifice opens anteriorly behind 1939. stylet. FL<4: Stylet present, with knobs. Cau- Stictylus piniphili (Fuchs, 1929) dal alae peloderan. Spicules and gubernaculum Ruhm, 1956. tylenehoid. AP 9 : Uterus completely everted, S. ungiilacaudus (Khan, 1957) Nickle, 1963. greatly enlarged, smoother and not as large as Sphaerularia, original female body wrinkled Syn. Sphaerularia ungulacauda Khan, 1957. and deformed, remains as an appendage to Stictylus ungulacaudus (Khan, 1957) uterus. Oviparous. Khan, 1960. HOST INSECTS: Coleoptera; Ernobius, Pi-ssodes, Dendrocto- This genus may eventually be placed under nus. Hymenoptera; Coeloides. Sphaerularia, but is easily distinguished at this TYPE SPECIES: time by the smaller size of the prolapsed Sphaerulariopsis stammeri Wachek, 1955. uterus, the knobbed stylet, the presence of the Syn. Stictylus stammeri (Wachek, 1955) caudal alae. It is normally parasitic in bark Ruhm, 1956. beetles.

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Genus: Tripins Chitwood, 1935 Genus: Allantonema Leuckart, 1884 Syn. Asconema Leuckart, Syn. Tylenchomorphus 1886. Fuchs, 1915. Atractonema (Leuck- (Figs. 1A, IF, 3A, 3B, 4A) art, 1886) Leuckart, FL 9 : Lips set off. Stylet well-developed. 1887. Ovary with few cells. Excretory pore posterior Proatractonema Bo- to nerve ring. FL $ : Lips weakly set off. Sty- vien, 1944. let present. Caudal alae narrow, peloderan. (Figs. 2BB, 2GC, 3Q, 3Y, 5A) Gubernaculum small. AP 9 : Bean-shaped, FL9: Stylet well-developed, with knobs. only 2-3 times as long as wide. Stylet retracted Two long esophageal glands. Ovary few-celled; into body. Oviparous or ovoviviparous. Vulva walls of uterus with large cells; vulva posterior. terminal. FL $ : Stylet faintly developed, without knobs. HOST INSECTS : Caudal alae absent. Spicules and gubemac- Coleoptera; Hylobius, Hijlastes, Philonthus, ulum small, tylenchoid. AP 9 : Fusiform. Ochthebius, Geotrupes. Diptera; Musca. Length up to 0.9 mm. Uterus partially pro- TYPE SPECIES: lapsed. Mouth cone and tail papilla present. Allantonema mirabile Leuckart, 1884. Stylet present, not retracted. Oviparous. Syn. Tylenchomorphus mirabills (Leuck- HOST INSECTS: art, 1884) Fuchs, 1915. Diptera; Cecidomyia or Dasine-ura, Bradysia, OTHER SPECIES: Sciara. A. matthesi Wachek, 1955. TYPE SPECIES: A. morosa (Fuchs, 1929) Filipjev, 1934. Tripius gibbosus (Leuckart, 1886) Chitwood, Syn. Tylenchus morosus Fuchs, 1929. 1935. Parasitylenchus morosus (Fuchs, Syn. Asconema gibbosiim Leuckart, 1929) Filipjev and Sch. Stek., 1886. 1941. Atractonema gibbosum (Leuckart, A. muscae Roy and Mukherjee, 1937. 1886) Leuckart, 1887. A. philonthi Wachek, 1955. OTHER SPECIES: A. silvaticum von Linstow, 1892. Tripius sciara (Bovien, 1944) Wachek, 1955. A. stricklandi Roy and Mukherjee, 1937. Syn. Proatractonema sciara Bovien, 1944. It is possible that A. muscae and A. strick- Poinar (1965) has shown that the morpho- landi are actually the same species, i.e., two logical basis for the genus Proatractonema may female stages of a Heterotylenchus. be just a stage in the development of a Tripius and he considers Proatractonema as a synonym Genus: Aphelenchulus Cobb, 1920 of Tripius. (Figs. 1C, 3C, 4A) SUBFAMILY : FL9: Not found. FLS: Stylet present. Allantonematinae Pereira, 1931. (Emended Length 1 mm. Caudal alae peloderan. Guber- Chitwood, 1935). naculum small, trough-like. AP 9 : Length 2.6 Syn. Allantoneminae Pereira, 1931. mm. Swollen, worm-shaped; tail not coiled Allantonematidae (Pereira, 1931) when relaxed by heat, ventral side not turned Chitwood and Chitwood, 1937. outward. Tail with diagnostic spike; not dor- Contortylenchidae Ruhm, 1956. sally bent, papilliform. Stylet present. Uterus DIAGNOSIS (Emended): Sphaerulariidae. containing several eggs. Oviparous. FL 9 •. With or -without stylet. Without eggs HOST INSECT: in uterus. FL $ : With tylenchoid anterior end. Coleoptera; Cyllene. Stylet usually present, with or without knobs. TYPE SPECIES: Caudal alae usually peloderan when present, Aphelenchulus mollis Cobb, 1920. not voluminous. Spicules and gubernaculum This nematode needs to be recollected and tylenchoid, when present. AP 9 : Swollen, restuclied to obtain additional information on uterus not everted. its morphological relationships with various

Copyright © 2011, The Helminthological Society of Washington JANUARY, 1967] HELMINTHOLOGICAL SOCIETY 79 groups in the family. The shape of the AP ?, tail drawn in by expansion. Stylet absent. which aids in separating this genus from Con- Vulva with distinct lips; ovary and oviduct tortylenchiis, is shown in Fig. 1G. short; uterus occupies up to 85% of body. Ovoviviparous. Genus: Bovienema Nickle, 1963 HOST INSECTS: (Figs. IB, 3D, 4A) Coleoptera; Aphodius, Spondylis, Throscus. FL?: Small. Stylet well-developed. FLS: Hemiptera; Gerris, Velia, Nepa. Diptera; Stylet present. With peloderan caudal alae. Bibio. Gubernaculum present. AP2: Length 0.5- TYPE SPECIES: 0.75 mm. Body swollen, habit in the form Bradijnema rigidum (von Siebold, 1836) zur of a tight circle, with externally turned ventral Strassen, 1892. side, see Contortylenchiis. Tail tip character- Syn. Filaria rigida von Siebold, 1836. istically peg-like, directed dorsally. Stylet al- Allantonema rigida (von Siebold, ways present, well-developed, not retracted. 1836) Moniez, 1891. Intestine not differentiated. Oviparous. Gonad OTHER SPECIES: with 2 or 3 flexures, one anterior in neck B. bibionis Wachek, 1955. region, one posterior in vicinity of vulva; one B. gerridis Poisson, 1933. to three eggs in uterus at one time. B. nepal Poisson, 1933. HOST INSECT: B. strasseni Wiilker, 1923. Coleoptera; Ptiijogenes. B. trixagi Wachek, 1955. TYPE SPECIES: B. veliae Poisson, 1933. Bovienema tomici (Bovien, 1937) Nickle, 1963. Genus: Chondronema Christie and Syn. Aphelenchulus tomici Bovien, 1937. Chitwood, 1931 Contortylenchiis tomici (Bovien, (Figs. 1C, 4D) 1937) Ruhm, 1956. Head with 4 distinct papillae. Stylet small, OTHER SPECIES: tylenchoid. Dorsal esophageal gland present. Bovienema chalcographi (Fuchs, 1938) Amphidial openings lateral, slightly nearer n. comb. mouth than papillae; amphidial glands large. Syn. Parasitylenchus contortus chalco- Esophagus without bulb-like swelling. A pair graphi Fuchs, 1938. of large lateral pores on the tail. Male with Parasitylenchus contortus f. chalco- small caudal alae; without gubemaculum. graphi (Fuchs, 1938) W. Schnei- Testis reflexed. Body of female degenerat- der, 1939. ing into nearly structureless sac filled with Contortylenchiis chalcographi developing embryos. Body-cavity parasites (Fuchs, 1938) Riihm, 1956. throughout larval development, but free-living throughout adult stage. Genus: Bradijnema zur Strassen, 1892 HOST INSECT: (Figs. 1H, 3E, 4A) Coleoptera; Passalus. FL9 : With smooth cuticle. Lips faintly set TYPE SPECIES: off. Stylet absent. Stomal region faintly cutic- Chondronema passali (Leidy, 1852) Christie ularized. Esophageal glands well-developed, and Chitwood, 1931. extending almost to the genital area. Intestine Syn. Nematoideum passali Leidy, narrow-celled, used for storage. FLcJ: With 1852. smooth cuticle. Lips not distinctly set off. The life history of this nematode differs Stylet absent. Caudal alae absent or faintly from the rest of the genera in the the family developed in B. trixagi. Spicules tylenchoid. because only the larval stages are parasitic in Gubemaculum always present. AP $ : Cuticle the haemocoel of the insect. No AP 9 's could and hypodermis traversed by small narrow be found in the body cavity of the insect. canals. Metabolic products white. Head and Christie and Chitwood described the adult

Copyright © 2011, The Helminthological Society of Washington 80 PROCEEDINGS OF THE [VOL. 34, No. 1 forms as free-living, and found them in the Aphelenchulus contortus typographi frass of the beetles. Larval nematodes prob- (Fuchs, 1915) Micoletzky, 1925. ably leave the host during oviposition. There Anguillulina contortus typographi is no reference to the type of gonad in the (Fuchs, 1915) Baylis and Daub- female, but a small tylenchoid stylet is present. ney, 1926. The morphology and life history of the adult Aphelenchulus diplogaster (Fuchs, forms need more study. 1915) Filipjev, 1934. Tylenchus contortus cembrae Fuchs, Genus: Contortylenchusl(uhm, 1956 1915. (Figs. II, 3F, 4A) Parasitylenchus contortus cembrae FL 9 : Stylet usually well-developed, tip (Fuchs, 1915) Fuchs, 1929. symmetrical, lumen narrow, basal knobs vari- OTHER SPECIES: able, usually prominent. Small postuterine sac. Ovary with fewer cells than in neoparasity- C. acuminati Ruhm, 1956. lenchs; vulva posteriorly located, with small C. amitini Ruhm, 1956. opening, without projecting lips. FL $ : Stylet C. barbems (Massey, 1957) Ruhm, 1960. present, dimorphic. Testis with flexure. Tail Syn. Aphelenchuhis barbems Massey, bulkier, shorter than in neoparasitylenchs; 1957. rounded, always with peloderan caudal alae, C. brevicomi (Massey, 1957) Ruhm, 1960. usually narrow. Spicules small, tylenchoid. Syn. Aphelenchuhis brevicomi Massey, Gubernaculum small. AP 5 : Sulphur to brown- 1957. yellow. Always sausage-shaped or worm- C. cryphali Ruhm, 1956. shaped; often coiled when relaxed by heat, Syn. Contortylenchiis cryphali Ruhm, with ventral side always turned outward; the 1954. nomen nudum. "contortus" group of Fuchs. Body surface smooth, no body swellings; anterior end often C. cunicularii (Fuchs, 1929) Ruhm, 1956. wrinkled, narrower than rest of body in the Syn. Tylenchus contortus cunicularii form of a movable mouth cone with a stylet. Fuchs, 1929. Intestine retained, well defined from rest of Aphelenchulus cunicularii (Fuchs, tissue. Gonad may have 1 or 2 flexures; ovi- 1929) Filipjev, 1934. duct longer than in neoparasitylenchs; uterus Parasitylenchus contortus f. c-uni- narrow, not expanded to hypodermis; vulva cularii (Fuchs, 1929) W. Schnei- subterminal, deeply recessed; vagina heavily der, 1939. cuticularized. Anus functionless. Tail rounded, C. elongatus (Massey, 1960) Nickle, 1963. or tapering to nipple-like papilla. Oviparous. Syn. Aphelenchuhis elongatus Massey, HOST INSECTS: 1960. Coleoptera; Ips, Dendroctonus, Cryphalus, C. grandicolli (Massey, 1957) Ruhm, I960. Hylastes, Orthotomicus. Syn. Aphelenchulus grandicolli Massey, TYPE SPECIES: 1957. Contortylenchiis diplogaster (von Linstow, C. laricis (Fuchs, 1929) Rtihm, 1956. 1890) Ruhm, 1956. Syn. Tylenchus contortus laricis Fuchs, Syn. Allantonema diplogaster von Lin- 1929. stow, 1890. Aphelenchulus laricis (Fuchs, 1929) Tijlenchus diplogaster (von Linstow, Filipjev, 1934. 1890) Fuchs, 1915. Parasitylenchus contortus f. laricis Tylenchus contortus typography (Fuchs, 1929) W. Schneider, Fuchs, 1914. 1939. Parasitylenchus contortus typo- C. reversus (Thorne, 1935) Ruhm, 1956. graphi (Fuchs, 1915) Micoletzky, Syn. Aphelenchulus reversus Thorne, 1922. 1935.

C. spirits (Massey, 1957) Riihm, 1960. Genus: Howardula Cobb, 1921 Syn. Aphelenchulus spirus Massey, 1957. Syn. Tylenchinema T. 'Goodey, 1930. The contortylenchs have the type of ovary Prothallonema Chris- that is typical of the family and do not differ tie, 1938. markedly from the rest of the genera. There- Acarinocola Warren, fore, Contortylenchidae Riihm, 1956, is con- 1941. sidered a synonym of Allantonematinae. (Figs. 1L, 1M, 3J, 3K, 3L, 4A) Genus: Heterotylenchus Bovien, 1937 FL 9 : Stylet present. Lips faintly set off. (Figs. IE, 1J, 2N, 2O, 31, 5B) FL $: Stylet absent. Caudal alae narrow, peloderan. Gubernaculum present. AP 9 : Four distinct forms. With alternation of Length 1.2-7.0 mm. Stylet present. White, gamogenetic and parthenogenetic generations. even in old age. Head and tail retracted. FL 9 : Small, slender, about 0.5 mm long. Lips Oviparous or ovoviviparous. Ovary and ovi- faintly set off. Stylet present, with knobs. duct in oviparous species coiled within body; Esophagus with overlapping glands and indis- in ovoviviparous species, crowded together in tinct orifices. Anus vestigial. Vulva posterior, anterior section of body. Vulva posterior, al- close to anus. FL<5: Slender, about 0.5 mm most terminal, lips protrude slightly. long. Stylet present, small, knobbed. Tail HOST INSECTS AND MITES: conical. Caudal alae absent. Spicules usually Coleoptera; Diabrotica, Phyllotreta. Dip- small, tylenchoicl. Gubernaculum lacking or tera; Oscinella. Thysanoptera; Aptinothrips, very small. Gamogenetic female (AP9): Taeniothrips, Frankliniella. Acarina; Para- Yellow-brown. Body swollen, curved, inert, situs, Poecilochirus, Haemogamasiis, Eury- much larger than the free-living stage (up to parasitus, Cosmolaelaps, Acarinocola. 3.00 mm); both ends rounded; or, tail with TYPE SPECIES: spike-like papilla. Stylet present, may be Howardula benigna Cobb, 1921. retracted into body. Intestine a syncytium. OTHER SPECIES: Anus vestigial. Ovary, oviduct and uterus of H. acarinorum Wachek, 1955. only moderate dimensions, outstretched; or, H. aoronymphium Welch, 1959. ovary and oviduct may lie in many convolu- H. aptini. (Sharga, 1932) Wachek, 1955. tions. Oviparous. Eggs larger than those of Syn. Tylenchus aptini Sharga, 1932. parthenogenetic female. Parthenogenetic fe- AngiiilluUna aptini (Sharga, 1932) male: More slender (up to 1.00 mm long). Lysaght, 1936. Anterior end drawn in by expansion. Tail H. clamger (Warren, 1941) Wachek, 1955. conical or dome-shaped. Stylet weakly de- Syn. Acarinocola claviger Warren, 1941. veloped. Esophagus degenerate. Ovary greatly H. cuneifer (Warren, 1941) Wachek, 1955. developed, reaching into head region. Ovip- Syn. Acarinocola cuneifer Warren, 1941. arous or ovoviviparous; when ovoviviparous, H. dubhim (Christie, 1938) Nickle, 1965. ovary and oviduct displaced anteriorly by Syn. Prothallonema dubhim Christie, uterus. 1938. HOST INSECTS: H. hirsuta (Warren, 1941) Wachek, 1955. Diptera; Hylemya, Musca. Coleoptera; Bem- Syn. Acarinocola hirsutus Warren, 1941. bidion, Clivina. Siphonaptera; Coptopsylla, H. oscinellae (Goodey, 1930) Wachek, Ceratophyllus. 1955. TYPE SPECIES: Syn. Tylenchinema oscinellae Goodey, Heterotylenchiis aberrans Bovien, 1937. 1930. OTHER SPECIES: H. phyllotretae Oldham, 1933. H. bovieni Wachek, 1955. H. terribilis (Warren, 1941) Wachek, 1955. H. stammeri Wachek, 1955. Syn. Acarinocola terribilis Warren, 1941. H. ivillkeri Wachek, 1955. This genus contains several diverse forms, PI. pawloivskyiKumchkin, 1960. and after more study it may be separated into

Copyright © 2011, The Helminthological Society of Washington 84 PROCEEDINGS OF THE [VOL. 34, No. 1 valid genera or subgenera. The distinguishing well-developed, with basal swellings. Ovary feature is the lack of stylet in the male. with more cells than Contortylenchus and Allantonema spp. FL 6 : Stylet present, not Genus: Metaparasitylenchus prominent. Caudal alae peloderan or lepto- (Wachek, 1955) n. grad. deran. Spicules and gubernaculum small, (Figs. 2S, 2T, 3N, 4A) tylenchoicl. AP 5 : Swollen, slightly mobile, Larvae leave host after second moult, third usually sausage- or worm-shaped. Stylet not moult occurs in free-life. After copulation, im- retracted into inner body. Tail terminal with pregnated young females infect host larva or or without papilla. Oviparous or ovoviviparous. pupa, moult once in new host for fourth and Ovary and oviduct intertwined irregularly, last time (after Wachek, 1955). FL?: Stylet crowded together anteriorly; uterus long, wide, strongly knobbed. FL $ : Stylet present. Cau- filled with eggs or larvae. dal alae pelocleran, frequently broad. Guber- HOST INSECTS: naculum present. AP 9 : White, seldom with Coleoptera (bark beetles); Hylurgus, Hy- yellow-brown deposits. Oviparous or ovovivip- lastes, Dendroctonus, Scolytus, Cryphalus, arous. Crypturgus, Pityophthorus, Pityogenes, Ips. HOST INSECTS: TYPE SPECIES: Coleoptera (not bark beetles); Cryptophagus, Neoparasitylenchus cnjphali (Fuchs, 1914) Mijcetophagus, Telmatophilus, Cossonus, Te- 11. comb. tropium, Strangalia, Rhizophagus, Plelmis, Syn. Tylenchus dispar cnjphali Fuchs, Latelmis, Riohis. 1914. TYPE SPECIES: Parasitylenchus dispar var. cnjphali Metaparasitylenchus telmatophili (Wachek, (Fuchs, 1914) Micoletzky, 1922. 1955) n. grad. Aphelenchuhis cnjphali (Fuchs, Syn. Parasitylenchus (Metaparasitylen- 1914) Filipjev, 1934. chus} telmatophili Wachek, Parasitylenchus dispar f. cnjphali 1955. (Fuchs, 1914) Schneider, 1939. OTHER SPECIES: Parasitylenchus (Parasitylenchus) M. cossoni (Wiilker, 1929) n. comb. cnjphali (Fuchs, 1914) Ruhm, Syn. P. cossoni Walker, 1929. 1956. P. (M.) cossoni (Wiilker, 1929) OTHER SPECIES: Wachek, 1955. N. avulsi (Massey, 1958) n. comb. M. cryptophagi (Wachek, 1955) n. comb. Syn. P. (M.) cryptophagi Wachek, 1955. Syn. Parasitylenchus avulsi Massey, 1958. M. helmidi-s (Wachek, 1955) n. comb. P. (P.) avulsi (Massey, 1958) Syn. P. (M.) helmidis Wachek, 1955. Ruhm, 1960. M. mycetophagi (Wachek, 1955) n. comb. N. betulae (Rtihm, 1956) n. comb. Syn. P. (M.) mycetophagi Wachek, 1955. Syn. P. (P.) betulae EiT^m, 1956. N. chalcographi (Fuchs, 1938) n. comb. M. oschei (Rtihm, 1956) n. comb. Syn. P. dispar chalcographi Fuchs, 1938. Syn. P. (M.) oschei Rtihm, 1956. P. dispar f. chalcographi (Fuchs, M. rhizophagi (Wachek, 1955) n. comb. 1938) Schneider, 1939. Syn. P. (M.) rhizophagi Wachek, 1955. P. chalcographi (Fuchs, 1938) M. strangalioe (Wachek, 1955) n. comb. Filipjev & Sch. Stek., 1941. Syn. P. (M.) strangaliae Wachek, 1955. P. (P.) chalcographi (Fuchs, 1938) M. tetropii (Wachek, 1955) n. comb. Ruhm, 1956. Syn. P. (M.) tetropii Wachek, 1955. N. cinerei (Fuchs, 1929) n. comb. Syn. Tylenchus dispar cinerei Fuchs, Genus: NeoparosUylenchusn. gen. 1929. (Figs. 2Q, 3P, 4A) Aphelenchuhis cinerei (Fuchs, 1929) With 3 distinct forms; one free-living sexual Filipjev, 1934. generation, one swollen female developing from P. dispar f. cinerei (Fuchs, 1929) the mated free-living female. FL9: Stvlet Schneider, 1939.

P. (P.) cinerei (Fuchs, 1929) N. louelkeri (Ruhm, 1956) n. comb. Ruhm, 1956. Syn. P. (P.) wuelkeri'Ruhm, 1956. P. dispar pusilli Fuchs, 1938. N. xylebori (Schvester, 1950) n. comb. P. dispart, pusilli (Fuchs, 1938) Syn. P. dispar xylebori Schvester, 1950. Schneider, 1939. P. (P.) xylebori (Schvester, 1950) P. pusilli (Fuchs, 1938) Filipjev Ruhm, 1956. and Stek., 1941. N. hi/last is (Wiilker, 1923) n. comb. Genus: ParasitylenchoidesWachek, 1955 Syn. Tylenchus hylastis Wiilker, 1923. (Figs. 2U, 3S, 4A) Tylenchus dispar ateri Fuchs, 1929. FL5 : Excretory pore 0.105-0.125 mm from Tylenchus dispar cunicularii Fuchs, anterior end, against 0.038-0.072 mm in Neo- 1929. para.sitylenchus (Diagnostic). Stylet present, P. hylurgi Filipjev, 1934. with knobs. FL $ : Stylet present. Caudal P. dispar f. cunicularii (Fuchs, 1929) alae peloderan or absent. Spicules tylenchoid. Schneider, 1939. Gubernaculum small. APS: Swollen, worm- P. dispar L ateri (Fuchs, 1929) shaped; head and tail not drawn in by expan- Schneider, 1939. sion; with mouth-cone and tail papilla. Stylet P. hylastis (Wiilker, 1923) Filipjev, in normal position or retracted. Oviparous or 1934. ovoviviparous. With oviparity, ovary and ovi- P. (P.) hylastis (Wiilker, 1923) duct show few windings. With ovoviviparity, Ruhm, 1956. ovary displaced anteriorly or to the side as N. ligniperdae (Fuchs, 1929) n. comb. uterus fills 80% of body. Metabolic products Syn. Tylenchus ligniperdae Fuchs, 1929. yellow-brown. P. ligniperdae (Fuchs, 1929) HOST INSECTS: Filipjev, 1934. Coleoptera; Anobium, Plegaderus, Micro- P. (P.) ligniperdae (Fuchs, 1929) malthus, Ditoma, Paederus, Sciodrepa, Aleo- Ruhm, 1956. chara, Oxytelus, Stenus, Paromalus. N. orthotomici (Ruhm, 1960) n. comb. TYPE SPECIES: Syn. P. (P.) orthotomici Ruhm, 1960. P. steni Wachek, 1955. N. ovaritis (Massey, 1958) n. comb. OTHER SPECIES : Syn. P. ovarius Massey, 1958. P. anobii, Wachek, 1955. N. pessoni (Ruhm, 1957) n. comb. P. diatomae Wachek, 1955. Syn. P. (P.) pessoni Ruhm, 1957. P. korneri Wachek, 1955. N. pityophthori (Ruhm, 1956) n. comb. P. paederi Wachek, 1955. Syn. P. (P.) pttyophthoriJtijhm, 1956. P. paromali Wachek, 1955. N. poligraphi (Fuchs, 1938) n. comb. P. rheocharae Wachek, 1955. Syn. P. dispar poligraphi Fuchs, 1938. P. sciodrepae Wachek, 1955. P. dispar f. poligraphi (Fuchs, 1938) P. wichmanni Wachek, 1955. Schneider, 1939. P. poligraphi (Fuchs, 1938) Filipjev Genus: Parasitylenchus Micoletzky, 1922 andSch. Stek., 1941. Syn. Polymorphotylenchus P. (P.) poligraphi (Fuchs, 1938) Ruhm, 1956. Ruhm, 1956. (Figs. 2V, 2X, 2Y, 30, 4C) N. rugulosi (Schvester, 1957) 11. comb. With 5 distinct forms, 2 sexual generations, Syn. P. dispar rugulosi Schvester, 1957. one swollen female developing from mated P. (P.) rugulosi (Schvester, 1957) free-living female. Small parasitic female: Ac- Ruhm, 1960. tive, narrow, sausage-shaped, present in large N.scolyti (Oldham, 1930) n. comb. numbers; or motionless, yellow-brown, raisin- Syn. P. scolyti Oldham, 1930. shaped. Stylet present, may be retracted, P. secundus Fuchs, 1933. smaller than in large female; basal knobs P. (P.) scolyti (Oldham, 1930) small, flat. Gonad with a short flexure, uterus Ruhm, 1956. with few eggs; or greatly developed with nu-

Copyright © 2011, The Helminthological Society of Washington 86 PROCEEDINGS OF THE [VOL. 34, No. 1 merous flexures of ovary. Small parasitic male: tis (Fuchs, 1914) Micoletzky, Active, narrow. Lips not set off. Excretory 1925. pore opens near stylet. Stylet present, weakly Anguillulina dispar var. curvidentis developed. Esophagus and intestine weak. (Fuchs, 1914) Baylis and Daub- Gonad outstretched, extends almost to stylet. ney, 1926. Tail plump, rounded. Caudal alae present or Aphelenchulus curvidentis (Fuchs, absent. Spicules and gubernaculum small. Free- 1914) Filipjev, 1934. living stages not described. AP 9 : Swollen, Parasitylenchus dispar f. curvidentis sausage-shaped, inwardly curved, milky white (Fuchs, 1914) Schneider, 1939. with red-brown spots; may be transparent or Polyrnorphotylenchus (Thylakolen- partly dark brown. Stylet present, knobs very chus) curvidentis (Fuchs, 1915) small, flat. Gonad well-developed; ovary and Riihm, 1956. oviduct with numerous crowded loops in an- Parasitylenchus diplogenus Welch, 1959. terior end; uterus large; or gonad weakly de- Syn. Polyrnorphotylenchus diplogenus veloped with more or less long flexure, not (Welch, 1959) Baker, 1962. pushed anteriorly, uterus narrow with few Welch (1959) pointed out that Riihm im- eggs. properly used the type species of the genus HOST INSECTS: Parasitylenchus Micoletzky, 1922, as the type Coleoptera; Ips, Pityokteines. Diptera; Dro- species of his new genus Polyrnorphotylenchus. sophila. Filipjev's 1934 designation of Tylenchus dispar TYPE SPECIES: typographi Fuchs, 1915 as the only synonym Parasitylenchus dispar (Fuchs, 1915) Filip- of his type species, Parasitylenchus dispar jev, 1934. (Fuchs, 1915) may not have been the best Syn. Tylenchus dispar typographi Fuchs, choice; however, it is valid according to the 1914. nomen nudum. International Rules of Zoological Nomencla- Tylenchus dispar typographi Fuchs, ture. Therefore, most of the other species have 1915. been removed from Parasitylenchus and placed Parasitylenchus dispar var. typo- in a new genus Neoparasitylenchus. The graphi (Fuchs, 1915) Micoletzky, subgenera Sulphuretylenchus, Metaparasitylen- 1922. chus, and Proparasitylenchus have been raised Aphelenchulus dispar var. tijpo- to generic level. Polymorphotylenchus has graphi (Fuchs, 1915) Micoletzky, been synonymized with Parasitylenchus. 1925. Genus: Proparasitylenchus Wachek, Anguillulina dispar var. tijpographi 1955n. grad. (Fuchs, 1915) Baylis and Daub- ney, 1926. (Figs. 2P, 3R, 4A) Parasitylenchus dispar f. typographi FL 9 : Copulated females penetrate into (Fuchs, 1915) Schneider, 1939. host pupa. Stylet present, usually with strong Polymorphotylenchus (Polymorpho- knobs, never bare or cleft. FL $ : Stylet pres- tylenchus) typographi (Fuchs, ent. Caudal alae peloderan or leptoderan, 1915) Riihm, 1956. always present. Gubernaculum present. AP 9 : OTHER SPECIES: White, or faintly citron-yellow, never brown. Parasitylenchus curvidentis (Fuchs, 1914) Ovoviviparous. Larvae leave host after second Micoletzky, 1922. moult, two moults in free-life. Syn. Tylenchus dispar curvidentis Fuchs, HOST INSECT: 1914. Coleoptera (staphylinids); Platystethus, Me- Parasitylenchus dispar var. curvi- don, Trogophloeus, Zyras, Atheta, Oxytelus. dentis (Fuchs, 1914) Micoletzky, TYPE SPECIES: 1922. Proparasitylenchus platystethi (Wachek, Aphelenchulus dispar var. curviden- 1955) n. grad. '

Syn. Parasitylenchus (Proparasitylenchus) basal swellings flat, not knobbed. Esophagus platystethi Wachek, 1955. and intestine degenerate. Ovoviviparous. Vulva OTHER SPECIES: and anus difficult to see in mature parasites. P. athetae (Wachek, 1955) n. grad. Gonad almost fills body, flexed about 3 times. Syn. P. (P.) athetae Wachek, 1955. Papillate tail tip absent. P. boopini (Wachek, 1955) n. grad. HOST INSECTS: Syn. P. (P.) boopini Wachek, 1955. Coleoptera (bark beetles); Pityogenes, Ips, P. medonis (Wachek, 1955) n. grad. Polygraphus, Dryocoetes, Hylastes, Scolytus. Syn. P. (P.) medonis Wachek, 1955. TYPE SPECIES: P. myrmedoniae (Wachek, 1955) n. grad. Sulphuretylenchus sulphureus (Fuchs, 1938) Syn. P. (P.) myrmedoniae Wachek, n. grad. 1955. Syn. Parasitylenchus sulphureus chalco- P. oxyteli (Wachek, 1955) n. grad. graphi Fuchs, 1938. Parasitylenchus (Sulphuretylenchus) Syn. P. (P.) oxyteli Wachek, 1955. sulphureus (Fuchs, 1938) Riihm, P. trogophloei (Wachek, 1955) n. grad. 1956. Syn. P. (P.) trogophloei Wachek, 1955. OTHER SPECIES: Genus: Protylenchus Wachek, 1955 S. elongatns (Massey, 1958) n. comb. (Figs. IK, 3T, 4A) Syn. P. elongatus Massey, 1958. S. escherichi (Riihm, 1956) n. grad. FL 9 : Lips not set off. Stylet present. Syn. P. (S.) escherichi Riihm, 1956. FL $ : Lips not set off. Stylet well-developed S. fuchsi (Fuchs, 1938) n. grad. with three cleft knobs. Caudal alae absent. Syn. P. sulphureus poligraphi Fuchs, Spicules tylenchoid. Gubernaculum proximally 1938. thickened. AP 9 : Cuticle bare, hypodermis P. (S.) fuchsi (Fuchs, 1938) Riihm, thick. Stylet present, retracted within body. 1956. Tail papilla present. Ovary and oviduct long, S. grossmannae (Riihm, 1954) n. grad. coiled profusely throughout body; vulval lips Syn. Paratylenchus (Sulphuretylenchus) slightly arched forward; uterus short. Ovip- grossmannae Riihm, 1954. Lapsus arous. Metabolic products white. No alterna- calami. tion of generations. S. kleinei (Riihm, 1956) n. grad. HOST INSECT: Syn. Parasitylenchus (Sulphuretylenchus) Coleoptera; Heterocerus. TYPE SPECIES: kleinei Riihm, 1956. P. heteroceri Wachek, 1955. S. pilifronis (Massey, 1958) n. comb. Syn. P. pilifronis Massey, 1958. Genus: Sulphuretylenchus Riihm, The significance of a group of genera with 1956 n. grad. a long stylet, without knobs, and with a dorsal (Figs. 2R, 3X, 4A) gland orifice which opens over a stylet length FL9: Longer than most sphaerulariids. behind the stylet base, is not known to me at V—A distance longer than neoparasitylenchs. this time. It includes some species of Sul- Excretory pore opens below nerve ring. Stylet phuretylenchus, Howardula, Heterotylenchiis large; lumen wide. FL $ : Stylet present. Tail and others. Future taxonomic work \vill un- longer than typical neoparasitylench. Caudal doubtedly clarify this relationship. alae peloderan. Spicules and gubemaculum small. AP9: Shape variable, usually long, Genus: Scatonema Bovien, 1932 with swollen spot on tube-shaped body, swell- (Figs. 2AA, 3U, 4A) ing caused by congestion of larvae and eggs FL9 : Small (0.33 mm long), slender. Sty- (Endotokie matricide). Body surface char- let well-developed, with basal knobs; tip with acteristically wavy, with constrictions. Sulphur oblique cut. First part of esophagus with dis- to yellow-brown or dark brown. Stylet re- tinct cuticular lining; two large ventral esoph- tracted into inner body, spear tip asymmetrical, ageal glands open separately into esophagus in

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Copyright © 2011, The Helminthological Society of Washington 90 PROCEEDINGS OF THE [VOL. 34, No. 1 front of nerve ring. No anal opening found. DIAGNOSIS (Emended): Sphaerulariidae. Oocytes few. Vagina surrounded by circle of FL9: Found in Eucalyptus gall. Eggs in large perivaginal cells. Tail tapering poste- uterus. Stylet well-developed. FL $ : With ty- riorly. FL6: Small (0.33 mm long), slender. lenchoid anterior end. Stylet present, knobbed. Stylet absent. Esophageal glands not seen. Caudal alae peloderan, not voluminous. Spic- Tail tapering behind anus. Spicules and guber- ules tylenchoid. Gubernaculum absent. AP 9 : naculum tylenchoid. Caudal alae present, nar- Swollen, uterus not everted. row. AP9 : Cigar- or sausage-shaped. Often coiled. Ovary greatly developed; swelling at Genus: Fergitsobia Currie, 1937 end of oviduct serves as receptaculum seminis; Syn. Angutttulma (Fergits- uterus occupies bulk of body; perivaginal cells obia) Currie, 1937. strikingly developed. Ovoviviparous. (Figs. ID, 3H, 4B) HOST INSECT: Three stages found: FL9: In gall. Head Diptera; Scatopse. offset, narrower than body, which is quite TYPE SPECIES: plump and faintly annulated. Spear tylenchoid Scatonema iviilkeri Bovien, 1932. with well-developed basal knobs. Anterior part of esophagus distended by ampullae filled with Genus: Dotylaphus Andrassy, 1958 esophageal secretions and appearing valve-like (Fig.3G) at times; esophagus narrows considerably at Based on one specimen. FL9 : Cuticle an- isthmus surrounded by nerve ring; glands nulated. Head not set-off, with small lips and overlap the intestine; dorsal gland orifice papillae. Stylet short, well-developed, similar typically tylenchoid, just behind spear base; to species of Dorylaimus, however, tip slanting subventral esophageal glands empty into swol- ventrally; guiding ring asymmetrical; stylet len median bulb-like area. Excretory pore pos- knobs are lacking; stylet continuation sclero- terior to nerve ring. Female gonad prodelphic; tized, relatively long; lumen wide (similar to no postuterine sac; vulva with distinct lips. Dorylaimus). Esophagus without muscles, Eggs present. Rectum and anus visible. FL 6 : without bulb, opening gradually into intes- Spicules paired, robust, knobbed, arcuate. tine; esophageal glands well-developed, long, Gubernaculum absent. Goodey illustrates a lie free in body cavity, outstretched, with en- narrow, peloderan caudal alae. Stylet present. larged ampulla-like openings; subventral gland Gonad single, outstretched. Tails of both sexes orifices in the middle of esophagus; dorsal short, conoid, rounded. AP 9 : Similar in shape gland orifice between anterior end and middle to Heterotijlenchus from carabid beetles. Ovip- of esophagus, far behind stylet. Lateral organs arous. Ovary winds several times within body. not seen. Lateral field well-developed, with Stylet retracted. Numerous lateral somatic several lines. Gonad single, prodelphic; vulva setae present. is posteriorly located. Tail short, rounded. HOST INSECT: FL5: Unknown. AP9: Unknown. Diptera; Fergusonina. HOST INSECT: TYPE SPECIES: Unknown. Fergusobia tumifacien.s (Currie, 1937) TYPE SPECIES: Chitwood and Chitwood, 1950. Dotylaphus riihmi Andrassy, 1958. Syn. Anguillulina (Fergusobia) tumi- This specimen belongs in this family. Un- faciens Currie, 1937. fortunately, it can fit into a few existing genera Anguillulina (Fergusobia) curriei and without further information it is difficult (Currie, 1937) Johnston, 1938. to place at this time. Therefore, these taxa I have studied males and females from the should be treated as genus and species in- gall and also the adult parasitic females from quirendae. the body cavity of the insect. Though the sub- SUBFAMILY: ventral gland orifices are swollen in the middle Fergusobiinae J. B. Goodey, 1963. of the esophagus, no criconematid valve could Syn. Fergusobiidae Siddiqi and J. B. be found. This fact dismisses any possible Goodey, 1963. association with the Criconematidae and dis-

Copyright © 2011, The Helminthological Society of Washington JANUARY, 1967] HELMINTHOLOGICAL SOCIETY 91 allows the family Fergusobiidae Siddiqi and OTHER SPECIES: J. B. Goodey, 1963. Fisher (1965) has re- I. fungorum (Biitschli, 1873) Fil. and Stek., cently found that the female in the gall pro- 1941. duces eggs. This separates this genus from all Syn. Tylenchus fungo-rum Biitschli, 1873. other members of this family. Hexatylus fungomm (Biitschli, SUBFAMILY: 1873) T. Goodey, 1932. lotonchiinae T. Goodey, 1953. Tylenchus (Anguillulina) fungorum Syn. lotonchiidae (T. Goodey, 1953) (Biitschli, 1873) Filipjev, 1934. Skarbilovich, 1959. Neotylenchus fungorum (Biitschli, DIAGNOSIS (Emended): Sphaerulariidae. 1873) Filipjev, 1936. FL 9 : With post-vulval and mid-ventral sup- Anguillulina fungorum (Biitschli, plement. Without eggs. Stylet well-developed. 1873) W. Schneider, 1939. FL $ : Head bilaterally symmetrical, tri-lobed, I. bifurcatum T. Goodey, 1953. flattened dorso-ventrally. Stylet not well- /. cephalostrictum Meyl, 1954. developed. Spicules large, L-shaped, elaborate. I. macrospiculatum (Meyl, 1954) J. B. Caudal alae voluminous, envelop tail tip. Post Goodey, 1956. anal papillae sometimes present. AP 5 : Un- Syn. Hexatylus macrospiculatus Meyl, known. 1954. I. mycophilum Meyl, 1954. Genus: lotonchium Cobb, 1920 T. Goodey (1953) and J. B. Goodey (1956, (Fig.3M) 1963) showed the correct sphaerulariicl status Esophagus ill-defined; dorsal gland orifice of lotonchium. I also agree with the synonymy joining esophageal lumen 4 head widths from of lotonchiidae (T. Goodey, 1953) Skarbilo- anterior end of body; subventral esophageal vich, 1959, with lotonchiinae T. Goodey, 1953. glands opening about 4—5 head-widths further In so doing we may discount Biitschli's female back; anterior part of esophagus cylindrical, of Tylenchus imperfectus (His Fig. 7d) which nerve ring crossing slightly constricted isthmus; showed an egg in the female, and accept the posterior part not distinguishable from intes- sphaerulariid ovary as described by T. Goodey. tine. Lateral fields, deirids and hemizonid However, the female with the egg may be present in both sexes. Female: Spear well- valid and could represent a situation similar to developed, probably changing shape at final that of Fergusobia. The life cycles of these moult. Male: Head offset by expansion, stylet nematodes need more study, especially the not well-developed. Spicules large, markedly identity of the host insects, if any, and the angular and L-shaped, sometimes with pos- descriptions of AP 2 's. terior extensions protruding through cloaca. Gubernaculum absent. Pre-adult male larva SUMMARY precocious; having fully developed spicules A taxonomic revision of the insect parasitic and gonad but no caudal alae. Head of larval nematode family Sphaerulariidae Lubbock, male radially symmetrical, changing to adult 1861 is presented and includes 4 subfamilies, shape at final moult. AP9: Unknown. 21 genera, and 116 species. The family group HOST INSECT: diagnoses are emended, and generic diagnoses Unknown. Found in basidiomycetous fungi are given along with a listing of species and Entoloma, Pleurotiis, Hygrophorus, and Tri- synonymies. The dorsal esophageal gland ori- choloma. fice of Sphaerularia bombi was found for the TYPE SPECIES: first time and was located in the normal lotonchium imperfect-urn (Biitschli, 1876) tylenchoid position. This discovery required Cobb, 1920. the shifting of the Sphaerulariidae from the Syn. Tylenchus imperfectus Btitschli, Aphelenchoidea to the Tylenchoidea, and be- 1876. cause the Sphaerulariidae is an older family Anguillulina (lotonchium) imper- group name than the Allantonematidae the fecta (Btitschli, 1876) W. Schnei- latter drops to subfamily status. Polymorpho- der, 1939. tylenchus Riihm, 1956 is synonymized with

Parasitylenchus Micoletzkv, 1922 and P. dispar , AND M. B. CHITWOOD. 1937. An Intro- (Fuchs, 1915) Filipjev, 1934 is considered as duction to Nematology. Baltimore, Maryland. the type species of Parasitylenchus. The re- CHRISTIE, J. R. 1938. Two nematodes associated with decaying citrus fruit. Proc. Helminth. maining species of the old genus Parasitylen- Soc. of Wash. 5(1): 29-33. chus are placed in a new genus Neoparasity- , AND B. G. CHITWOOD. 1931. Chondro- lenchns, requiring 16 new combinations. The nema passali (Leidy, 1852) n. g. (Nematoda) families Contortylenchidae and Allantonemati- with notes on its life history. Jour. Wash. dae are considered at this time to be synonyms Acad. Sci. 21: 356-364. of the Allantonematinae. Fergusobiidae is COBB, N. A. 1920. One hundred new nemas. synonymizecl with Fergusobiinae which is Contrib. Sci. Nematol. 9: 217-343. placed as a subfamily of the Sphaerulariidae. . 1921. Howardula benigna, a nemic parasite of the cucumber beetle (Diabrotica). The writer agrees with Goodey's synonymy of Contrib. Sci. Nematology 10: 345-352. lotonchiidae with lotonchiinae and its place- CURRIE, G. A. 1937. Galls on Eucalyptus trees. ment with these insect parasites. Dotylaphus A new type of association between flies and ruhmi Andrassy, 1958 is treated as genus and nematodes. Proc. Linn. Soc. N.S. Wales 62 species inquirendae. The subgenera Sulphur- (3/4): 147-174. etylenchus Riihm, 1956, Metaparasitylenchus DUFOUR, L. 1837. Recherches sur quelques en- Wachek, 1955, and Proparasitylenchus Wachek, tozoaires et larves parasites des insectes or- 1955 are raised to generic rank. Host insects thopteres et hymenopteres. Ann. Sci. Natur. are given whenever possible because they are Zool. 7: 5-20. FILIPJEV, I. N. 1934. Harmful and useful nema- useful in identification. Typical life cycles are todes in rural economy. 440 pp. Leningrad, illustrated along with plates of representative Moskva. adult parasitic females, and free-living male , AND J. H. SCHUURMANS-STEKHOVEN, JR. and female stages. 1941. A manual of agricultural helminthol- ogy. E. J. Brill, Leiden, 878 pp. LITERATURE CITED FISHER, J. M. 1965. (Personal Communication). FUCHS, G. 1914. Tijlenchus dispar cnrvidentis ANDRASSY, I. 1958. Erd-und Siisswassernemato- m. und Tijlenchus dispar cryphali m. Zool. den aus Bulgarian. Acta Zoologica, Acad. Sci. Anz. 45(5): 195-207. Hung. IV (1/2): 1-88. . 1915. Die Naturgeschichte der Nema- BAKER, A. D. 1962. Check lists of the nematode toden und einiger anderer Parasiten des Zp.s superfamilies Dorylaimoidea, Rhabditoidea, typographies L. und Hylobius abietif, L. Zool. Tylenchoidea, and Aphelenchoidea. E. J. Brill, Jahrb. (Syst.) 38(3/4)': 109-222. Leiden, 261 pp. . 1929. Die Parasiten einiger Riissel- und BOVIEN, P. 1932. On a new nematode, Sca- Borkenkjifer. Ztschr. Parasitenk. 2(2): 248- tonema ivi'dkeri gen. et spec. n. parasitic in the 285. body-cavity of Scatopse fuscipes Meig. (Dip- . 1933. Einige Nematoden bei Scolytus tera Nematocera). Vid. Meddel. fra Dansk scoltjtus. F. Capita Zoologica 4(1): 1—45. naturh. Forening. K0benhavn. 94: 13-32. 1938. Neue Parasiten und Halbpara- •. 1937. Some types of association be- siten bei Borkenkafern und einige andere tween nematodes and insects. Vid. Meddel. Nematoden. II, III, IV. Zool. Jahrb. (Syst.) fra Dansk naturh. Forening. K0benhavn 101: 71(1/2): 123-190. 1-114. GOODEY, J. B. 1956. Observations on species of 1944. Proatractonema sciarae n. g., n. the genus, lotonchium Cobb, 1920. Nema- sp., a parasitic nematode from the body cavity tologica 1(3): 239-248. . 1960. The classification of the Aphelen- of a dipterous larva. Vid. Meddel. fra Dansk choidea Fuchs, 1937. Nematologica 5: 111- naturh. Forening. K0benhavn. 108: 1-14. 126. BUTSCHLI, O. 1873. Beitrage zur Kenntnis der 1963. Soil and freshwater nematodes. freilebenden Nematoden. Nova Acta (Acad. (Revision of T. Goodey, 1951) Methuen, Nat. Curios) 36: 144 pp. London, 544 pp. . 1876. Untersuchungen iiber freilebende GOODEY, T. 1930. On a remarkable new nema- Nematoden und die Gattung Chaetonotus. tode, Tylenchinema oscinellae gen. et sp. n., Ztschr. Wissensch. Zool. 26: 363-413. parasitic in the frit fly, Oscinella frit L., at- CHITWOOD, B. C. 1935. Nomenclatorial notes, 1. tacking oats. Philos. Tr. Roy. Soc. London Proc. Helminthol. Soc. of Wash. 2: 51-54. 218: 315-343.

. 1953. On certain eelworms, including . 1963b. Bovienema (Nematoda: Allan- Biitschli's Tylenchus fungorum, obtained from tonematidae), a new genus parasitizing bark toadstools. Jour. Helminth. XXVII (1/2): 81- beetles of the genus Pityogenes Bidel, with 94. notes on other endoparasitic nematod.es of KHAN, M. A. 1957a. Sphaerularia bombi Duf. scolytids. Proc. Helminthol. Soc. Wash. 30 {Nematoda: Allantonematiclae) infesting bum- (2): 256-262. blebees and Sphaerularia hastata sp. nov. in- -, AND G. W. WOOD. 1964. Howardula festing bark beetles in Canada. Canad. Jour. aptini (Sharga, 1932) parasitic in blueberry Zool. 35(4): 519-523. thrips in New Brunswick. Can. Jour. Zool. . 1957b. Sphaerularia ungulacauda sp. 42(5): 843-846. nov. (Nematoda: Allantonematidae) from the PEREIRA, C. 1931. Myenchus botelhoi n. sp., Douglas fir beetle, Dendroctonus pseudotsugae curiosa nematoide parasite de Limnobdella Hopk., with key to Sphaerularia species brasiliensis Pinto (Hirudinea). Thesis, Fac. (Emended). Canad. Jour. Zool. 35(6): 635- Med., Sao Paulo, 30 pp. 639. POINAR, G. O. 1965. The bionomics and para- 1960. Stictylus hastatus (Khan, 1957) sitic development of Tripins sciarae (Bovien) n. comb., and Stictylus ungidacaudus (Khan, (Sphaerulariidae: Aphelenchoidea), a nema- 1957) n. comb. (Nematoda: Neotylenchidae). tode parasite of sciarid flies (Sciaridae: Dip- Canad. Jour. Zool. 38(1): 225-226. tera). Parasitology 55: 559-569. KUROCHKIN, Y. V. 1960. The nematode Hetero- POISSON, R. 1933. Trois nouvelles especes de tylenchus pawlowskyi n. sp. castrating flea nematodes cle la cavite generale d'hemipteres carriers of plague. Dokladi Academii Nauk aquatiques. Ann. Parasit. Humaine et Com- U.S.S.R. 135(5): 1281-1284. pare. 11(6): 463-466. LEUCKART, R. 1884. Ueber einen neuen hetero- ROY, D. N., AND P. K. MUKHERJEE. 1937. Al- genen Nematoden. Tagebl. 57. Vers. Deut- lantonema stricklandi n. sp., a parasitic nema- scher Naturf. u. Aerzte. p. 320. tode of houseflies, Mnsca vicina. Ann. Trop. LINSTOW, O. VON. 1890. Uber Allantonema und Med. and Parasit. 31(4): 453-456. Diplogaster. Zentralbl. Bakt. und Paraskde RiiHM, W. 1956. Die Nematoden der Ipiden. 8(16): 489-493. Parasitologische Schriftenreihe 6: 1-437. VEB LUBBOCK, J. 1861. On Sphaerularia bombi. Nat. Fischer Verlag, Jena. Hist. Rev. 1(1): 44-57. . 1960. Ein Beitrag zur Nomenklatur und MASSEY, C. L. 1956. Nematode parasites and Systematik einiger mit Scolytiden vergesell- associates of the Engelmann spruce beetle schafteter Nematodenarten. Zool. Anz. 164 (Dendroctonus engelmanni). Proc. Helmin- (5/6): 201-213. thol. Soc. Wash. 23(1): 14-24. . 1957. Four new species of Aphelen- SCHNEIDER, W. 1939. Wurmer oder Vermes. II. chulus (Nematoda) parasitic in bark beetles Fadenwiirmer oder nematoden. I. Freile- in the United States. Proc. Helminthol. Soc. bende und pflanzenparasitische Nematoden. Wash. 24(1): 29-34. Tierwelt Deutschlands (Dahl) 36: 1-260. . 1958. Four new species of Parasity- SIDDIQI, M. R., AND J. B. GOODEY. 1963. The lenchus (Nematoda) from scolytid beetles. status of the genera and subfamilies of the Proc. Helminthol. Soc. Wash. 25(1): 26-30. Criconematiclae (Nematoda); with a comment . 1960. Nematode parasites and associ- on the position of Fergusobia. Nematologica ates of the California five-spined engraver, 9(3): 363-377. Ip* confusus (Lee.). Proc. Helminthol. Soc. SKARBILOVICH, T. S. 1947. Revision of the sys- Wash. 27(1): 14-22. tematics of the nematode family Anguillulin- 1962. Life history of Aphelenchulus ,idae Baylis and Daubney, 1926. Akademia elongatus Massey ( Nematoda), an enclopara- Nauk USSR 57(3): 307-308. site of Ips confusus LeConte, with a descrip- . 1959. On the structure of systematics of tion of the male. Jour. Insect Pathol. 4(1): nematodes, order Tylenchida Thorne, 1949. 95-103. MICOLETZKY, H. 1922. Die freilebenclen Erd- Acta Parasitol. Polonica 7(4): 117-132. Nematoden. Arch. Naturg. Abt. A. 87: 1-650. STRASSEN, OTTO K. L. ZUR. 1892. Bradijnema NICKLE, W. R. 1963a. Notes on the genus rigidiim v. Sieb. Ztschr. f. wissensch. Zool. Contortylenchus Riihm, 1956, with observa- 54(4): 655-747. tions on the biology and life history of C. THORNE, G. 1935. Nemic parasites and asso- elongatus (Massey, 1960) n. comb., a para- ciates of the mountain pine beetle (Dendroc- site of a bark beetle. Proc. Helminthol. Soc. tonus monticolae) in Utah. Jour. Agr. Res. Wash. 30(2): 218-223. 51(2): 131-144.

WACHEK, F. 1955. Die entoparasitischen Tylen- clrosophilid flies. Parasitology 49(1/2): 83- chiden. Parasitolog. Schriftenreihe, 3: 1—119 103. Jena, Fischer. WULKER, G. .1923. Uber Fortpflanzung und En- WARREN, E. 1941. On the occurrence of nema- twicklung von Allantonema und verwandten todes in the haemocoel of certain gamasid Nematoden. Ergebn. Fortschr. d. Zool. 5: mites. Ann. Natal Mus., 10(1): 79-94. 389-507. WELCH, H. E. 1959. Taxonomy, life cycle, de- . 1929. Bemerkungen zur Arbeit von G. velopment, and habits of two new species of Fuchs: Die Parasiten einiger Riissel-und Bork- Allantonematidae (Nematoda) parasitic in enkiifer. Z. Parasitenkcle 2(2): 286-290.

Encholaimoidea (Nematoda: Dorylaimida), A New Superfamily Representing Dorylaimid Specimens with Cephalic Setae

A. MORGAN GOLDENTI AND D. G. MuRPHY1

In the fall of 1963 nematodes obtained from as in other dorylaims. It is of the masculine a soil sample around the roots of coconut gender and is the latinized combination formed (Cocos nucifera L.) in Trinidad were sub- from the Greek "enchos," meaning spear, and mitted for identification by Dr. Karl Mara- "laimos," meaning throat. The specific Latin morosch, Boyce Thompson Institute for Plant name "taunts" means "bull," and refers here Research, Yonkers, New York. Among the to the presence of horn-like cephalic setae. nematodes found in this and other samples col- lected shortly thereafter from the same area Order Dorylaimida (De Man, 1876) were a total of about 30 specimens of a very Pearse, 1942 emended remarkable nematode. The amphids, spear, DIAGNOSIS: Nematoda, Adenophorea. Setae esophagus, and certain other structures were usually absent. Cuticle generally smooth; but clearly dorylaimid. Furthermore, the develop- sometimes heavily annulated, in some cases ment of the spear as seen in immature specimens, is the same as in other species of spear- longitudinal striae forming plate-like cuticle. bearing dorylaims. Yet, these specimens pos- Esophagus cylindrical, in 1 or 2 parts; when sess four small, and six prominent horn-like in 2, the posterior part conoid cylindrical or cephalic setae previously unknown in the pyriform. Glands uninucleate, and 3, 5, or 7 Dorylaimida. Also, the heavily annulated cuti- in number contained within the confines of cle divided into plates by numerous longi- the esophagus; all gland openings posterior to tudinal striae is unique in the dorylaims. the nerve ring. Head with 6—18 inner labial This interesting nematode is described here- papillae and 6, 10, or 14 outer papillae or in as the representative of a new genus, family, setae. Amphids cyathiform, pouch-like or tu- and superfamily. The Order Dorylaimida is bular. Stoma with mural tooth or teeth; with emended as specified below, so that this an axial stylet, or vestigial, or unarmed. Pre- nematode could be placed in this major group anal supplements usually present. to which it apparently belongs. Encholaimoidea, new superfamily The new generic name, Encholaimus, sug- DIAGNOSIS: Dorylaimida, sub-order Dory- gests "spear in the throat," referring to the laimina. development of the spear in the esophagus Head with setae. Stoma armed. Amphids pouch-like, with slit-like apertures. Esoph- 1 Nematologists, Nematology Investigations, Crops Re- agus composed of a slender anterior part search Division, Agricultural Research Service, U. S. De- partment of Agriculture, Beltsville, Maryland. and an expanded posterior portion. Excretory