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A Revision of the Genus Rotylenchulus Liiiford and Oliveira, 1940 (Nematoda: Tylenchiclae)1 D

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A Revision of the Genus Rotylenchulus Liiiford and Oliveira, 1940 (Nematoda: Tylenchiclae)1 D OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 169 Hagan, H., and R. Hoopingarner. In press. vien) (Sphaerulariidae: Aphelenchoidea), a Mermithid nematode parasitism of Aedes nematode parasite of sciarid flies (Sciariclae: stimulans (Walker) (Diptera: Culicidae) Diptera). Parasitology 55: 559-569. from Ingham County, Michigan. Ann. Ent. In press. Arthropod immunity to worms. Soc. Amer. In: Jackson, G., and R. Herman, Immunity Hyinan, L. H. 1951. The Invertebrates: Acan- to parasitic animals. Appleton-Century-Crofts, thocephala, Aschelminthes, and Entoprocta, New York. The pseudocoelomate bilateria. Vol. III. Mc- -, and H. E. Welch. In press. A new Graw-Hill, New York. 572 pp. nematode, Filipjevimermis leipsandra sp. nov. Johnson, A. A. 1955. Life history studies on (Mermithidae) parasitic in chrysomelid larvae Hydromermis contorta (Kohn), a nematode (Coleoptera). J. Invert. Path. parasite of Chironomiis plumostis (L.). Ph.D. Rennie, J. 1925. A mermithid parasite of Tipula thesis Univ. 111. 92 pp. paludosa Meigen. Proc. Royal Phys. Soc. Lavoipierre, M. J. J. 1958. Studies on the host- Edinb., 21: 1-3. parasite relationships of filarial nematodes and Rimando, L. C., R. A. Corey, and Yun-Pei Sun. their arthropod hosts. 1. The sites of develop- 1965. Mass rearing of the western spotted ment and the migration of Loa loa in Chnj- cucumber beetle. J. Econ. Ent. 59: 230-231. sops silacea, the escape of the infective forms Steiner, G. 1933. Some morphological and phys- from the head of the fly, and the effect of the iological characters of the mermithids in their worm on its insect host. Ann. Trop. Med. relationship to parasitism. J. Parasit. 19: Parasit. 52: 103-121. 249-250. Poinar, Jr., G. O. 1965. The bionomics and Strickland, E. H. 1930. Phagocytosis of inter- parasitic development of Tripius sciarae (Bo- nal insect parasites. Nature. London, 126: 95. A Revision of the Genus Rotylenchulus Liiiford and Oliveira, 1940 (Nematoda: Tylenchiclae)1 D. R. DASGUPTA, D. J. RASKI, AND S. A. SHER Department of Nematology, University of California Species of the genus Rotylenchulus have The genus Rotylenchulus was proposed by been one of the most misidentified groups of Linford and Oliveira in 1940 when they de- all the tylenchs. As evidence, nematodes scribed R. reniformis. Yokoo and Tanaka belonging to this genus have been described (1954) described Tetylenchus nicotiana from in at least four different genera. Also this Japan which was subsequently transferred to genus has been variously assigned to three the genus Rotylenchulus by Baker (1962). families: Tylenchidae (Linford and Oliveira, Three other species (Helicotylenchus eli- 1940; Thome, 1949, 1961; Allen and Sher, sensis Carvalho, 1957, 1959; Spirotylenchus 1967); Heteroderidae (Chitwood and Chit- queirozi Lordello and Cesnik, 1958, and Heli- wood, 1950; Skarbilovich, 1960); Hoplolaim- cotylenchus parvus Williams, 1960) were idae (Hopper and Cairns, 1959; Goodey, transferred to the genus Rotylenchulus by Sher 1963; Husain and Khan, 1967) and five dif- (1961). In 1960 Das proposed a new genus, ferent subfamilies: Nacobbinae (Hopper and Leiperotylenchus, which he considered to be Cairns, 1959; Goodey, 1963); Pratylenchinae closely related to Tylenchus and Ditylenchus. (Thome, 1949, 1961; Baker, 1962); Tylen- However, the position of the dorsal gland chulinae (Skarbilovich, 1960); Hoplolaiminae orifice and characters of male tail indicated a (Loof and Oostenbrink, 1962); Rotylenchul- close relationship with Rotylenchulus. Indeed, inae (Husain and Khan, 1967; Allen and Sher, Loof and Oostenbrink (1962) transferred 1967). Leiperotylenchus leiperi to the genus Roty- lenchulus. Goodey (1963) synonymized eli- i A part of the thesis submitted by senior author in par- sensis, parvus, leiperi and queirozi with R. tial fulfillment of the requirements for the Ph.D. degree, University of California, Davis. reniformis. Husain and Khan (1965) described Copyright © 2011, The Helminthological Society of Washington 170 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY R. stakmani from India. Swarup, et al. (1967) land (USDANC, Beltsville); Nematology De- considered it a synonym of R. reniformis. partment, Rothamsted Experimental Station, The kidney-shaped mature females from Harpenden, England (NDRES, England); which the common name, reniform nematodes, Plantenziektenkundige Dienst, Wageningen, was derived, are described only in two species, The Netherlands (PD, The Netherlands). R. reniformis and R. borcalis. The descriptions of other species were based on immature Morphology of Rotylenchulus females and males. Rotylenchulus species are characterized by Representatives of the genus Rotylenchulus sexual dimorphism, mature females being have world-wide distribution and their host swollen to kidney shape and males vermiform records are numerous and diverse (Linford with not as well developed stylet and esoph- and Yap, 1940; Peacock, 1956; Martin, 1955). agus. Although detailed morphology of Their general occurrence and economically mature females is obscure, the overall shape great potential as plant pathogens make it and that part of the body posterior to the especially important that the taxonomy of this vulva provide useful taxonomic characters. group be soundly established to assure accu- Fixed specimens of males, immature females rate identification of species. For this purpose and larvae assume a spiral to open C shape this study has been carried out. upon fixation. INCISURES: Immature females, males and Materials and Methods larvae all have four incisures in the lateral Over 3,500 permanently mounted specimens field. Posteriorly on males these incisures of Rotylenchulus from more than thirty coun- extend varying distances from slightly anterior tries covering most parts of the world have to the cloacal opening to about midway on been assembled for this generic revision. Type the tail. The most ventral line always ends at specimens for most of the nominal species of the level of or anterior to the cloaca and on this genus were obtained. the caudal alae except in R. macrodoratus For immature females two measurements of where that ventral line appears to merge with the esophagus, b and b' (Sher, 1963), are the edge of the caudal alae. The lateral field included in the de Man formula. Also included of mature females was not visible. in these measurements are: o (Perry, Darling PHASMIDS: These are porelike and located and Thome, 1959); c' (Sher, 1986) and h slightly anterior to the middle of the tail. (length of hyaline portion of tail in microns). EXCRETORY PORE: The excretory pore is The measurements given in parenthesis in located posterior to the median bulb in all descriptions of the holotype, allotype and neo- species. type refer to population range. HEMIZONID: The hemizonid is two to three An immature female was designated as annules long and immediately anterior to the holotype for R. parvus and mature females as excretory pore. holotype for R. borealis and neotype for R. LIP REGION: The lip region is continuous, nicotiana. The immature female is used as not set off and varies in shape and annulation. the holotype for the new species described Most species have either low and rounded lip here because many morphological characters region (R. parvus) or higher and conoid in are more easily seen in these than in swollen shape (R. macrodoratus n. sp.). Some species females. The swollen female is more difficult appear to be intermediate between these two to collect, indeed is unknown for most species categories (R. reniformis1). despite diligent search for them. Immature females of some species have Type specimens are being deposited in five three or more distinct annules (R. reniformis). permanent institutional collections -which are In other species there is evidence of fine indicated by the following abbreviations: ampliations, but these are difficult to count University of California Nematode Collection, (R. parvus). There is still another group Davis (UCNC, Davis); University of Califor- where annulation is entirely lacking or the nia Nematode Collection, Riverside (UCNC, annules are too fine to resolve (R. leptus n. Riverside); United States Department of Agri- sp.). culture Nematode Collection, Beltsville, Mary- The shape and annulation of the lip region Copyright © 2011, The Helminthological Society of Washington OF WASHINGTON, VOLUME 35, NUMBER 2, JULY 1968 171 of larvae in all species are similar to the im- agus in males is much reduced with almost mature females except R. anamictus n. sp. no evidence of the median bulb, valve and where the larvae have a conoid lip region in lumen. The posterior part of the male esoph- contrast to the low and rounded lip region of agus overlaps the intestine laterally and ven- immature females. Males in general have a trally, usually more ventrally. rounded lip region. The esophagus of the second-stage larva is The labial framework of immature females not reduced like that of males. Often it is is conspicuous. The cheilorhabdions are obscure in fixed specimens but when seen it thickened anteriorly, extend laterally and in is usually similar to the immature female. some species are thickened at both ends. The However, the posterior part of the esophagus basal ring of the cephalic framework has the of larvae is often asymmetrical. same thickness as that of the cheilorhabdions REPRODUCTIVE SYSTEM: Rotijlenchulus fe- and arches posteriorly in a characteristic man- males have didelphic, amphidelphic ovaries. ner. The cephalic framework in males is The ovaries
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