132 Journal o[ Nematology, Volume 10, No. 2, April 1978 material is a residue of secretions present chodoridae. Pages 103-127. in F. Lamberti, in the spicules. Because of no visible vas C. E. Taylor, and J. w. Seinhorst, eds. Nematode vectors of plant viruses. Plenum deferens, and because spicules appear hol- Press, London and New York. low with a terminal or ventro-terminal 6. RODRIGUEZ-M, R., and A. H. BELL. 1978. opening or pore, it is possible to speculate Three new species of Trichodoridae that sperms pass through the spicule body. (Nematoda: Diphtherophorina) with obser- vations on the vulva in Paratriehodorus LITERATURE CITED Siddiqi, 1973. J. Nematol. 10:132-141. 7. RODRIGUEZ-M, R., S. A. SHER, and M. R. 1. ALLEN, M. W. 1975. A review of the nematode SIDD1QI. 1978. Systematics of the monodel- genus Trichodorus with descriptions of ten phic species of Trichodoridae (Ncmatoda: new species. Nematologica 2:32-62. Diphtherophorina) with descriptions of a new 2. HOOPER, D. J. 1962. Three new species of genus and four new species. J. Nematol. 10: Trichodorus (Nematoda: Dorylaimoidea) and 141-152. observations on T. mi,nor Colbran, 1956. 8. SHER, S. A. and A. H. BELL. 1975. Scanning Nematologica 7:273-280. electron tnicrographs of the anterior region 3. HOOPER, D. J. 1975. Morphology of Tri- of some species of Tylenchoidea (Tylenchida: chodorid Nematodes. Pages 91-101. in Netnatoda). J. Nematol. 7:69-83. F. Lamherti, C. E. Taylor, and J. w. Sein- 9. SIDDIQI, M. R. 1973. Systematics of the horst, eds. Nematode vectors o1 plant viruses. genus Trichodorus Cobb, 1913: (Nematoda: Plenum Press, London and New York. Doylaimida) with descriptions of three new 4. LOOF, P. A. A. 1973. Taxonomy of Trichodorus species. Nematologica 19:259-278. aequalis-complex (Diphtherophorina). Nema- 10. SCZYGIEL, A. 1968. Trichodorus sparsus sp. n. tologica 19:49-61. (Nematoda: Trichodoridae). Bull. Ac. Pol. 5. LOOF, P. A. A. 1975. Taxonomy of Tri- Sci. 16:695-698. Three New Species of Trichodoridae (Nematoda: Diphtherophorina) with Observations on the Vulva in Paratrichodorus R. RODRIGUEZ-M. and A. H, BELL~ Abstract: Trichodorus intermedius n. sp., Trichodorus dilatatus n. sp., and Paratrichodorus (dtlantadorus) grandis n. sp. are described from soil around native plants of Southern California. The females of the three subgenera of Paratrichodorus are further characterized on the basis of the vulva, which is pore-like in Atlantadorus, a longitudinal slit in Paratrichodorus, and a transverse slit in Nanidorus. Key Words: Taxonomy, morphology, Trichodorus intermedius, Trichodorus dilatatus, Paratrichodorus (Atlantadorus) grandis, supplementary papillae. In 1913 Cobb described the genus Tri- taxonomic study of the genus (1). Siddiqi chodorus on the basis of T. obtusus collected discussed the systematics and morphology around roots of grasses at Arlington, Vir- of the genus, proposed the genus Paratri- ginia (3). Interest in the genus was increased chodorus, and erected subgenera (8). He by the discovery by Christie and Perry of also constructed a key to the species of the pathogenicity of a species of Tri- Trichodorus on the basis of male char- chodorus to beets and corn. This species acters. Because males are rare in some was later described as T. christei by Allen species and unknown in others, we under- in a review of the germs that for the first took a search for characters that would time clearly identified the morphological facilitate identification of individual fe- characters which could be used for species males. identification, laying a foundation for the During a study of trichodorids in the nematode collection at the University of California, Riverside, three new species Received for publication 8 June 1977. aDepartment of Nematology. University of California, River- associated with native plants of Southern side, CA 92521. Present Address (Senior Author): Colegio California were found and are described de Postgraduados, Escuela National de Agricultura, Cha- pingo, Mexico. herein. Additional species were included in Trichodoridae Taxonomy: Rodriguez-M., Bell 133 a study of variation of the morphology of arcuate, slightly cephalated, distal one-third the vulva. slightly expanded. Proximal two-thirds of retracted spicules appear serrated; when METHODS AND MATERIALS extended, bristle-like structures occur along proximal two-thirds of spicule. Guber- Specimens used in the description of naculum linear, 19 tzm, with ventral end new species were fixed in formalin, trans- expanded. Protractor muscles prominent. ferred to dilute glycerin, and allowed to Three ventromedian supplementary papil- dehydrate slowly. Processed specimens were lae, 72, 24, and I0 /zm from anus. Paired mounted on permanent slides either in subventral postcloacal papillae. Tail dehydrated glycerin or in glycerin jelly rounded, with slight thickening of cuticle (dorsal-ventral mounts). Type materials around almost terminal caudal pores. were used for all species. Measurements Allotype 9: L = 0.73 mm; a -- 18.0; were of permanent mounts by means of b = 4.9; V = 55; onchiostyle = 64 ~m. camera lucida and are expressed as the Body straight to slightly curved ventrally. mean ± 1 standard deviation followed by Cuticle, lip region, and esophagus as in the range. male. Excretory pore 110 (78.4-138 in paratypes) t~m from anterior end. Gonads Trichodorus intermedius n. sp. paired, reflexed. Spermatheca (present or absent in paratypes) filled with round Fig. 1 sperm. Vagina more than one-half vulvar- Measurements: Paratypes 11 ~ ~ : L = body-width in length, with cuticularization 0.83 -+- 0.07 (0.72-0.95) ram; a = 21.0 ± and musculature prominent. Vaginal 2.1 (18-24); b = 5.0 +__ 0.59 (3.9-5.9); c = cuticularization appearing as two triangles 68 -+- 10.2 (59-91); T -- 66.8 -+- 3.6 (63- with long bases in lateral view oblique to 72.8); onchiostyle = 69.0 ~ 2.5 (62-71.5) body axis, with tips characteristically bent ~m; spicules = 45.0 ± 3.0 (42-50) #m; toward interior of vagina (Fig. 4-D). Vulva gubernaculum = 20.0 -4- 1.3 (18-22) ttm. a large transverse slit. Paired lateral body Paratypes 23 9 9: L = 0.85 -+- 0.1 (0.68- pores about 20 tam posterior to vulva. Tail 1.13) ram; a -- 20.0 ± 1.1 (17.2-22); b = rounded, with terminal caudal pores. Anus 5.0 -+- 0.5 (4.3-6.4); anus subterminal; V = subterminal. 57.0 -+- 2.7 (53-64); onchiostyle -- 69.0 ± Holotype ~: Collected by A. Bell on 3.4 (62-75.2) ttm. April 22, 1967. Department of Nematology Holotype 5: L = 0.72 ram; a -- 20.0; Nematode Collection, University of Cali- b = 4.6; ¢ = 91; T = 63. Body straight fornia, Riverside. Catalog No. 28. anteriorly, strongly curved ventrad at pos- Allotype 9: Same data as holotype. terior end. Cuticle not abnormally swollen, Catalog No. 29. subcuticle finely striated. Lip region promi- Type locality and habitat: Soil around nent, somewhat conical. Sixteen papillae roots of live oak (Quercus virginiana Mill) arranged in an inner circle of 6 around oral at Oak Grove Campground, San Diego Co., aperture and an outer circle of 10 com- California. Additional specimens have been posed of 2 pairs of subdorsal papillae, 2 found associated with the following hosts pairs of subventral papillae, and 2 single in Southern California: grass and wil- lateral papillae (7). Amphid aperture near low (Salix sp.), Dripping Springs near level of outer circle of papillae. One Temecula; Chenopodium sp. and Ambrosia ventral cervical papilla anterior to ex- sp., Santa Ana River, Riverside; Pinus sp., cretory pore, 100 ~m from anterior end. Mt. Palomar; Salvia leucophyUa, south of Excretory pore 122 ttm from anterior end. Camarillo; Quercus sp., Fallbrook; Quercus Onchiostyle 62 tzm, tip 30 /~m. Collar or sp. and grass, Live Oak Park; also in red- "guide ring" 27 ~m from anterior end. wood soil at Big Sur. Nerve ring midway between onchiostyle T. intermedius was sometimes found base and ventral cervical papilla. Esophagus mixed with Paratrichodorus (P.) aUius, and esophageal gland nuclei position typ- Paratrichodorus (N.) minor, or Trichodorus ical of genus. Testis single, outstretched, aequalis. with large round sperm. Spicules 42 t~m, Paratypes: 15 ~ ~, 30 9 9, 20 jj, same 134 Journal of Nematolog),, Volume 10, No. 2, April 1978 ['~' , .: . .:. ~i Q .............~:. .... , • ~'. .>,~:~ ~ :.,,,~.. ~.@ @ ".~.~......':,.'.'..:~, • .~ ., • ~....!~..;::.~:.;~.. z;~ o i ; '-.. ~!~.-- .?.'; ~:-~. I .A" F 0.03 mm , FIG. I-(A-F). Trichodorus intermedius n. sp.: ,4) male anterior region (holotype); B) female posterior region (paratype); C) vagina and related structures in lateral view; D) vulva in ventral view; E) male posterior region (holotype); F) male spicule (paratype), Trichodoridae Taxonomy: Rodriguez-M., Bell 135 data as holotype, distributed as follows: disappear near esophagus. Onchiostyle 65 6 ~ ~, 15 9 9, 13 jj, Department of Nema- /~m, tip 30 /~m. Collar or "guide ring" 29 tology, University of California, Riverside; ixm from anterior end. Nerve ring 18 /~m 2 ~ ~, 3 9 2, Nematode Collection, Uni- posterior to onchiostyle base, midway be- versity of California, Davis; 2 ~ ~, 3 9 9, tween onchiostyle base and ventral cervical USDA Nematode Collection, Nematology papilla. Esophagus typical of genus; an- Investigations, Behsville, Maryland; 1 ~, terior subventral gland nuclei anterior to 2 2 2, Rothamsted Experimental Station, excretory pore; posterior subventral gland Harpenden, England; 2 ~ ~, 4 9 2, 7 j j, nuclei 20 /~m anterior to the esophago- Commonwealth Institute of Helminthology, intestinal junction; dorsal esophageal gland St. Albans, England; 2 ~ ~, 3 9 9, Nema- nucleus larger than nuclei of subventral tology Department, Landbouwhogeschool, glands, and located near same level as Wageningen, The Netherlands. posterior subventral gland nuclei. Gonad Diagnosis: By having males with only single, outstretched, containing large round one ventral cervical papilla, T. intermedius sperm. Spicules curved ventrally, expanded is similar to T. californicus Allen, 1957 (1) at proximal end, then uniformly tapered and T. cottieri Clark, 1963 (2), but differs to distal end.