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A Reappraisal of Tylenchina (Nemata) : 8. the Family Hoplolaimidae Filip

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A Reappraisal of Tylenchina (Nemata) : 8. the Family Hoplolaimidae Filip A reappraisal of Tylenchina (Nemata). 8. The family Hoplolaimidae Filip'ev, 1934(l) Renaud FORTUNER" California Department of Food and Agriculture, Analysis and Identification (Nematology), 1220 N Street, Sacramento, CA 95814, USA. SUMMARY The classification of the family Hoplolaimidae is revised.The familes Nemonchidae, Aphasmatylenchidae, and Rotylenchulidae are considered synonyms of Hoplolaimidae. Only two subfamilies are accepted within Hoplolaimidae : Hoplolaiminae (syn. : Nemonchinae, Rotylenchoidinae, Aphasmatylenchinae, Rotylenchinae, and Pararotylenchinae) with eight valid genera: Hoplolai- mus, Rotylenchus, Helicotylenchus, Scutellonema, Aorolaimus, Aphasmatylenchus, Antarctylus,and Pararoty1enchus;and Rotylenchu- linae (syn. : Acontylinae) with Rotylenchulus, Acontylus, and Senegalonema. Peltamigratus and Nectopelta are proposed as new synonyms of Aorolaimus; Hoplolaimoides asa new synonym of Hoplolaimus; Orientylus and Varotylus asnew synonyms of Rotylenchus. A tabular key for the identification of the valid genera is presented. New combinations proposed for each accepted genus are listed. RESUME Réévaluation des Tylenchina (Nemata). 8. La fàmille des Hoplolaimidae Filip'ev, 1934 (Tylenchoidea) La classification de la famille Hoplolaimidae est révisée. Les familles des Nemonchidae, Aphasmatylenchidae et Rotylenchulidae sont considérées comme synonymes mineurs des Hoplolaimidae. Deux sous-familles sont acceptéesà l'intérieur des Hoplolaimidae: Hoplolaiminae (= Nemonchinae, Rotylenchoidinae, Aphasmatylenchinae, Rotylenchinae et Pararotylenchinae) avec huit genres valides : Hoplolaimus, Rotylenchus, Helicotylenchus, Scutellonema, Aorolaimus, Aphasmatylenchus, Antarctylus, Pararotylenchus;et Rotylenchulinae (= Acontylinae) avecRotylenchulus, Acontylus et Senegalonema. Peltamigratuset Nectopelta sont proposés comme nouveaux synonymes mineurs d'dorolainzus; Hoplolaimoides comme nouveau synonyme mineur d'Hoplolaimus, et Orientglus et Varotylus comme nouveaux synonymes mineurs de Rotylenchus. Une clé tabulaire pour l'identification des genres valides est présentée. .Une liste des combinaisons nouvelles est présentée pour chaque genre valide. The hoplolaimids were fïrst proposed as a separate accepted todayas the mosttypical representatives of the taxon byFilip'ev (1934).At thetime, the nominal genus, family. Hoplolaimus, included somespecies that now belong to The Thorneianconcept of hoplolaimidshas been the criconematids (H. annulifer, H. informis, H. menzeli, accepted by al1 later authors. The contribution of the etc., even Criconemaguernei and C. morgeme had more recent classifications consisted mostly in the ad- belonged for a time to Hoplolaimus). Not surprisingly, dition of newly described genera and the arrangement Filip'ev included a number of criconematidgenera of recognized genera under various subfamilies. The [Paratylenchus,Iota (= Ogma),Criconema, Procrico- validityof some of these genera rests on criteria of nenza (= Hemicycliophorajl in his newsubfamily Hoplo- dubious taxonomic value. The increase in the number laiminae. of subfamilies (seven subfamilies for eleven valid gen- After the organization of the criconematids as a era) raise some doubtson the necessity of creating separate subfamily by Taylor (1936), Thorne (1949) higherrank taxa. The taxonomic position of some rearranged the hoplolaimids to include the genera Ho- genera (Rotylenchulus, Acontylus)is still unsettled. plolaimus, Rotylenchus,and Helicotylenchus that are still Wieser (1953) is generally credited for the upgradeof (1) This article ispart of a studyon the classification of Tylenchina by the present author E.and Geraert (Rijksuniversiteit, Gent), M. LUC(ORSTOM, Paris), and A. R. Maggenti and D. J. Raski (University of California, Davis). * Associate in the Division of Nematology, University of California, Davis, USA. R evue Nématol. Revue (1987) 10 (2) :219-232 219 R. Fortuner ~ ~~ ~ ~ ~~~~~ Hoplolaiminae to family level. In fact, the author uses located anterior to anus, rarely on thetail. Tail two body thename " Hoplolaimidae " in only onesentence diameters long or less. Caudal alae of male leptoderan. (p. 452) out of a long article on the morphology of the Gubernaculum withtitillae. Deirids absent. Y " oral cavity andits relationships with the ecologyof The relationship between Hoplolaimidae and related marine nematodes. Discussing the cuticular differen- families in Tylenchina have been discussed by Maggenti tiations of the " Monoposthiinen ", Weiser continues, et al. (1987). " Typisch ist diese Funktion der Kutikularanhange bei der terrestrischen Familie Hoplolaimidae ausgepragt, wo SUBFAMILIESAND GENERA IN HOPLOLAIMIDAE die Hautstacheln als Antagonisten des durchdas Type subfamily : Vorschnellendes Speeres erzeugten " Riickstosses Hoplolaiminae Filip'ev, 1934. wirken (S. Stauffer, 1924) (Transl. : " Typically, this = Nemonchinae Skarbilovich, 1959 function of the cuticular appendagesis well marked in = Rotylenchoidinae Whitehead, 1958 the terrestrial family Hoplolaimidae, in which the skin = Aphasmatylenchinae Sher, 1965 (n. syn.) spikes act as antagonists of the " reverse push caused = Rotylenchinae Golden, 1971 (n. syn.) by the thurst of the spear (Stauffer, 1924). ") = PararotylenchinaeBaldwin & Bell, 1981 It is evident that Wieser was not proposingto upgrade (n. syn.) the subfamily, but that he was using the name " Ho- Hoplolaimus von Daday, 1905 plolaimidae " as a convenient term to refer to a group Rotylenchus Filip'ev, 1936 of taxa (see Art. 11, e of the International Code of Helicotylenchus Steiner, 1945 Zoological Nomenclature). It can be noted that Wieser Scutellonema Andrassy, 1958 was unfamiliar withthe family and was still including in Aorolaimus Sher, 1963 it the criconematids in spite of the works of Taylor Aphasmatylenchus Sher, 1965 (1936) and Thorne (1949). It remains that, because of Antarctylus Sher, 1973 the coordinate status in the family group (Art. 36), the Pararotylenchus Baldwin & Bell, 1981 author of Hoplolaimidae is Filip'ev, 1936. Other subfamily : The nominal subfamily Hoplolaiminaewas included Rotylenchulinae Husain & Khan, 1967 by Goodey (1963) with several other subfamilies (Roty- = Acontylinae Fotedar & Handoo, 1978 lenchoidinae, Belonolaiminae, Dolichodorinae, Praty- Rotylenchulus Lindford & Oliveira, 1940 lenchinae, and Nacobbinae) in a higher rank taon : Acontylus Meagher, 1968 Hoplolaimidae. Similarly, but ata higherlevel, Paramo- Senegalonema Germani, Luc & Baldwin, nov (1967) grouped Hoplolaimidae,Pratylenchidae, and 1983 Heteroderidae under the superfamily Hoplolaimoidea. The presentstudy does not include these higher COMMENTS rankings but presents a reasoned classification of the genera in Hoplolaimidaeaccording to the guidelines Description of Hoplolaimidae fromLuc etal. (1987) and to the classificationof Tylenchoidea. Female vermiform to kidney-shaped; Tylenchina proposed by Maggenti et al. (1987). when vermiform, habitus often spiral. Lip region high, Minimal lists of species are given for some genera. typically higher than 1/2 the diameter of the basal lip Additional information on species nomenclature can be annulus; anterior end with roundedor trapezoidal found in Fortuner (1986b). outline in lateral view, annulated, sometimeswith longi- tudinal striae on basal lip annulus,rarely striae on other lip annuli. Lateral field typically with four lines, some- The family Hoplolaimidae Filip'ev, 1934 times regressed(some Hoplolaimus spp.). Phasmids = Nemonchidae Skarbilovich, 1959 typically nearanus level, rarely on tail, sometimes = Aphasmatylenchidae Sher, 1965 migrated far anteriorly (Hoplolaimus), generally small = Rotylenchulidae Husain & Khan, 1967 pore-like structures, sometimes enlarged into scutella, rarely absent (Aphasmatylenchus). Tail typically short, DIAGNOSIS less than two tail diameters long, rarely longer; generally morecurved dorsally, sometimes regularly rounded, Tylenchoidea. Lip region higherthan 1/2 thediameter rarely conical. Caudalids and cephalids generz!!y pres- of the basal lip annulus, with rounded or trapezoidal ent; deirids absent. outline inlateral view. Stylet strong, 2-1/2-3 times longer Labialframework strong, withhigh arches. Stylet than thediameter of the basal lip annulus. CEsophageal strong, its length typically equals to 2-1/2 to 3 times the glands generally overlapping the anterior intestine. diameter of the basal lip annulus. Stylet knobs strong, Females with two genital branches, posterior branch rounded to indented, sometimes anchor-shaped. DG0 sometimesreduced to a post-uterine sac. Columned at least 4 Pm, sometimes more than 20 Pm, from the uterus with three rows of four cells. Phasmids generally stylet base. Median bulb strong, rounded. Esophageal 220 Revue Nématol. 10 (2) :219-232 (1987) Reappraisal 01 Tylenchina. 8. Hoplolaimidae glands anrangement variable but mostly overlappingthe exists a number of differences between Rotylenchus and intestine. Esophago-intestinal junction a small triangu- Helicotylenchus (i.e., position of DGO,esophageal lar structure. glands arrangement,striations of the basal lip annulus), Two genital branches opposed, outstretchedor rarely that speaks against a close relationship between the two 1 reflexed (Rotylenchulus); posterior branch may bede- genera. On the other hand, Rotylenchus, except for the generated or reduced to a PUS. Columneduterus size of the phasmids, is remarkably similar to Scutello- with three rowsof four cells. Epiptygma and vulval nema. Rotylenchus and Scutellonema should be kept in flaps
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