A reappraisal of Tylenchina (Nemata). 8. The family 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, , , Aorolaimus, Aphasmatylenchus, Antarctylus,and Pararoty1enchus;and Rotylenchu- linae (syn. : Acontylinae) with , Acontylus, and Senegalonema. Peltamigratus and Nectopelta are proposed as new synonymsof Aorolaimus; Hoplolaimoides asa new synonym of ; 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 () 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 . 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,, and 1983 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 generally present but sometimes inconspicuous. the same subfamily. Male with secondarysexual dimorphism present, with Pararotylenchinae would be the most acceptable of anterior end less developed than in females, sometimes the proposed subfamilies. This monotypic taxon inclu- degenerated and non-functional. Caudal alae generally des a genus very different from therest of the Hoplolai- enveloping the tail end, rarely stopping short of it (Ro- midae. Some species of Pararotylenchus were originally rylenchulus). Gubernaculum with titillae. described in . In fact, Pararotylenchus Biology : Typically, ecto or semi endo-parasites of can be seen as an evolutionary dead-end where some higher plants. belonolaimid species engaged inthe way tothe hoplolaimids but were not quite successful in evolving the modifïed esophageal glands characteristics of the Synonymyzation of families and subfamilies family. However, the mainjustification for Pararotylen- Nemonchus, type genus of Nemonchinaeand Ne- chinae is the arrangement of the esophageal glands. As monchidae, has been shown by Thorne (1935) to be for Aphasmatylenchinae, Pararotylenchinae is rejected identical with Hoplo2ainzus because the differentiating to avoid an inflation in the number of subfamilies. characters of Nenzonchus galeatus are artefacts created by a bad fixation of Hoplolaimus coronutus. This makes the family Nemonchidaea junior synonym of Hoplolai- The subfamily Hoplolaiminae Filip'ev, 1934 inidaa. Rotylenchoidinae was created forthe taxa with a DIAGNOSIS single genital branch, differentiated fromthe other Hoplolaimidae.Adult female remains vermiform. hoplolaimids withtwo genital branches. It has been Lateral field with four lines or less. Phasmids near anus argued (Fortuner, 1984 and see below the discussion on or erratic, anteriorly migrated on body, rarely on tail, Helicotylenchus) that theregression of part of the female small or enlarged, rarely absent. Tai1 short, more curved genital system is not a Sound basis for generic differ- dorsally with or without a ventral projection, or regularly entiation, far less for the definition of a subfamily. rounded, rarely conoid. Caudalids and cephalids pre- Aphasmatylenchinae, elevated to familyrank by sent, labial framework and stylet strong. DG0 more or Fotedar and Handoo (1978),was differentiated from less,far from stylet. Esophageal glandseither of the same Hoplolaiminae by the absence of phasmids, the elonga- length and not overlapping the intestine, or variously ted amphid apertures, and the shape of the esophageal enlarged and overlapping the intestine. Genital branches r glands. The regression of an organ should not be used always outstretched, two of equal length or posterior for taxonomic purposes.The amphids of Aphasmatylen- branch smaller or reduced to a post-uterine sac (PUS). chus were studied with SEM by Sher and Bell (1975). Male with anterior end smaller than females but still They are of the same relative length as the amphids of functional. Caudal alae enveloping tail end. Helicotylenchushydrophilus and Antarctylushumus Migratoryecto or semi-endo parasites of higher (compare figs 6 A, 8 B, and 8 C in Sher & Bell, 1975). plants. Eggs not deposited in a gelatinous matrix. Sher and Bell (1975) concluded that Aphasmatylenchus Type genus : Hoplolaimus von Daday, 1905. ' nigen'ensis " has a face view similar to the basic pattern seen in the Hoplolaimidae ". Esophagealglands are variously arranged in different genera of Hoplolaimi- THEGENERA IN HOPLOLAIMINAE dae : abutti,ng lobes in Pararotylenchus; dorsal overlap Pararotylenchus Baldwin & Bell, 1981 ' in Rotylenchus,Scutellonema, and Hoplolainzus; equal ' ventral and dorsal overlap in Helicotylenchus; unequal Diagnosis : ventral and dorsal overlap in Antarcty1us;ventral overlap short in Aphasmatylenchus and long in Rotylenchulinae. Hoplolaiminae. Female :Body spiral to C-shaped. To accept the gland arrangementas a subfamily charac- Labial region annulated; oral opening slit-like; round ter would result inthe creation of nearly as many labial disc present; amphid apertures elongate at lateral subfamilies as there are valid genera in Hoplolaimidae. edge of disc; first lip annulusdivided intosix lip sectors; Rotylenchinaegroups Rotylenchus and Helicotylen- lateral sectors smaller than the others (SEM). Lateral ' chus with phasmids pore-like, differing from Hoplolai- field with four lines. Labial framework,stylet and stylet nzus and related genera with enlarged scutella. There knobs average sized for the family; knobsround to

Revue Nématol. 10 (2) :219-232 (1987) 22 1 R. Fortuner indented. DG0 about 4-7 pm from stylet base. Eso- = Onentylus Jairajpuri & Siddiqi, 1977 phageal glands not overlappingintestine, symmetri- = Calvatylus Jairajpuri & Siddiqi, 1977 cally arranged around esophageal lumen. Two genital ’ = Varotylus Siddiqi, 1986 (n. syn.) branchesoutstretched, equally developed. Epitygma double, present. Tail short, of variable shape, usually Diagnosis : more curved on dorsal side; phasmids small, pore-like, near levelof anus. Male : Caudal alae enveloping Hoplolaiminae. Female :Body spiral to C-shaped. tail. Gubernaculum reflexed distally and withsmall Labial region offset or continuous with body contours, titillae. Spicules trough-shaped. No secondary sexual anteriorly rounded or flattened, generally annulated, dimorphism. with or without longitudinal striae onbasal lip annulus. Lateral field with four lines, with or without scattered Type species : transverse striae. Labial framework,stylet, and stylet Pururotylenchus hoppen Baldwin & Bell, 1984 knobs average sized for thefamily; knobs with rounded = P. brevicaudutus (Hopper, 1959) Baldwin & to indented anterior surface, DG0 often close to stylet Bell, 1981. (6 pm) but with a tendancy to posteriorly directed migration (up to 16 pm). Esophageal glandsoverlap Other species : intestine dorsally and laterally; dorsal gland more devel- See list in Baldwin and Bell (1981). oped than subventral glands; intestine symmetrically arranged between the subventral glands. Two genital Discussion : branchesoutstretched, equally developed; posterior Pururotylenchus belongs to Hoplolaimidae because branch rarely degenerated. Epiptygma single or double of the short tail, phasmids near anus level, DG0 more present. Tail short, hemispherical, rarely with small than 4pm from the styletbase, high labial region, strong ventral projection; phasmids pore-like, small, near anus stylet and labial framework, and spiralhabitus. The level. Males : Caudal alae enveloping tail, not lobed. esophageal glands not overlappingthe intestine are Secondary sexual dimorphism not marked, sometimes similar to theglands in Tylenchorhynchusand some other anterior partof male body slightly smaller than female. . In the hypothesis that hoplolaimids originatedfrom belonolaimid-like ancestral forms, Type species : (de Man, 1876) Filip’ev, 1936. Pararotylenchus can be seen as a relict of the inter- = Hoplalaimus uniformis Thorne, 1979 mediary forms where some but not al1 of the characters = Scutellonema picea Gubina, ,1973 of Hoplolaimidae were present. If this hypothesis is true, the arrangementof the esophageal glands, that had Other species : been used (for example, by Thorne, 1949) to differen- R. abnormecaudatus Van den Berg & Heyns, 1974 tiate Tylenchorhynchus and the Tylenchinae from the R. acuspicaudatus Van den Berg & Heyns, 1974 Hoplolaimidae, was in fact the last characterto evolve. R. agnetis Szczygiel, 1968 Purarotylenchus is veryclose to Rotylenchus and, R. alpinus Eroshenko, 1976 except again for the arrangementof the glands, there is R. basiri (Khan & Khan, 1982) n. comb. littleto separate the two genera. It is probable that = Orientylus basiri Khan & Khan, 1982 Rotylenchus evolved directly from proto-Parurotylenchus = Varotylus basiri(Khan & Khan, 1982) Siddiqi, 1986 forms when the dorsal esophageal gland enlarged and R. bialaebursus Van den Berg & Heyns, 1974 overlapped the intestine. Other differences between the R. brevicaudatus Colbran, 1962 two genera, such as the presence of longitudinal striae R. breviglans Sher, 1965 on the lip annuli, appeared only later and are visible R. buxophilus Golden, 1956 today only in some species of Rotylenchus. Baldwin and = R. sheri Jairajpuri, 1964 Bell (198 1) noted that a morphological continuum exists R. calvus Sher, 1965 from Pararotylenchus, with glands of equal length, not = Calvatylus calvus (Sher, 1965) Jairajpuri & Siddiqi, 1977 overlapping the intestine, and esophageal lumen sym- R. capensis Van den Berg & Heyns, 1974 metrically arranged between the three glands, to Roty- R. capitatus Eroshenko, 1981 lenchus fallorobustus and R. breviglans Who also have R. catharinae Van den Berg & Heyns, 1974 glands of equal length not overlapping the intestine but R. caudaphasmidius Sher, 1965 where the esophageai lumen is shiifted ventraiiy between R. citri Rashid & Khan, 1974 the subventralglands, and finally to thetypical Rotylen- = Orientylus citri (Rashid & Khan, 1974) Jairajpuri & chus spp. with dorsal overlap. Siddiqi, 1979 = Varotylus citri (Rashid & Khan, 1974) Siddiqi, 1986 Roty2eenchu.s Filip’ev, 1936 R. cypriensis Antoniou, 1980 = Anguillulina (Rotylenchus) Filip’ev, 1936 R. dalhousiensis Sultan & Jairajpuri, 1979 = Gottholdsteineria Andrassy, 1958 R. desouzai (Kumar & Rao, 1976) Fortuner, 1984

222 Revue Nématol. 10 (2) :219-232 (1987) Reappraisal 01 Tylenchina. 8. Hoplolaimidae

= Rotylenchoides desouzai Kumar & Rao, 1976 = Orientylus peculiaris Khan & Khan, 1982 = Orientylus desouzai (Kumar & Rao, 1976) Orton = Varotylus peculiaris (Khan & Khan, 1982) Siddiqi, Williams, 1984 1986) R. devonensis Van den Berg, 1976 R. phaliurus Siddiqi & Pinochet, 1979 R. elegans (Khan & Khan, 1982) n. comb. R. pruni Rashid & Husain, 1972 = Orientylus elegans Khan & Khan, 1982 R. pumilus (Perry in Perry, Darling & Thorne, 1959) Sher, = Varotyluselegans (Khan & Khan,1982) Siddiqi, 1961 1986 R. quartus (Andrassy, 1958) Sher, 1961 R. eximius Siddiqi, 1964 = Gottholdsteineria quarta Andrassy, 1958 R. fabalus Baïdulova, 1984 R. ranapoi Darekar & Khan, 1982 R. fallorobustus Sher, 1965 = Varotylus ranapoi (Darekar & Khan, 1982) Siddiqi, R. jêroxcis Eroshenko, 1981 1986 R. geraerti (Jairajpuri & Siddiqi, 1979) Zancada& Lima, R. rugatocuticulatus Sher, 1965 1986 R. satsilinicus Sultan, 1985 = Orientylus geraerti Jairajpuri & Siddiqi, 1979 R. secondus Mulk & Jairajpuri, 1976 R. glabratus Kankina & Teben’kova, 1980 = Orientylus secondus (Mulk & Jairajpuri, 1976) Jai- R. goodeyi Loof & Oostenbrink, 1958 rajpuri & Siddiqi, 1979 R. gracilidens (Sauer, 1958) Sauer, 1958 = Varotylus secondus (Mulk & Jairajpuri, 1976) Sid- R. helicus Husain & Khan, 1967 diqi, 1986 = Varotylus helicus (Husain & Khan, 1967) Siddiqi, R. siddiqii (Mulk & Jairajpuri, 1986) 1986 = Orientylus siddiqii (Mulk & Jairajpuri, 1976) Jairaj- R. heredicus (Jairajpuri & Siddiqi, 1979) Ferraz, 1980 puri & Siddiqi, 1979 = Calvatylus heredicus Jairajpuri & Siddiqi, 1979 = Varotylus siddiqii(Mulk & Jairajpuri, 1976) Siddiqi, R. himprus (Sultan, 1980) n. comb. 1986 = Orientylus himprus Sultan, 1980 R. symmetricus (Sultan, 1980) n. comb. = Varotylus hinzprus (Sultan, 1980) Siddiqi, 1986 = Orientylus symmetricus Sultan, 1980 R. impur (Phillips, 1971) Germani, Baldwin, Bell & Wu, = Varotylus symn~etricus(Sultan, 1980) Siddiqi, 1986 1986 R. triannulatus Van den Berg & Heyns, 1974 = Scutellonenza inlpar Phillips, 1971 R. uniformis (Thorne, 1949) Loof & Oostenbrink, 1958 R. incisicaudatus (Phillips, 1971) Germani, Baldwin, Bell R. unisexus Sher, 1965 & Wu, 1986 R. usitatus Van den Berg & Heyns, 1974 = Scutellonema incisicaudatum Phillips, 1971 R. varus (Jairajpuri & Siddiqi, 1979) Zancada & Lima, R. incultus Sher,1965 . 1986 R. indorobustus Jairajpuri & Baqri, 1973 = Orientylus varus Jairajpuri & Siddiqi, 1979 = Scutellonenza petersi Mahajan, 1977 = Varotylus varus (Jairajpuri & Siddiqi, 1979) Siddiqi, R. insularis (Phillips, 1971) Germani, Baldwin, Bell & 1986 Wu, 1986 = Scutellonenza insulare Phillips, 1971 The names R. cognatus, R. julaharensis, R. yarikaen- R. ivanovae Kankina & Teben’kova, 1980 sis, O. populus, and O. prominens proposed in a thesis R. jagatpurensis Sultan, 1985 (Kapoor, 1982) are not available. Tylenchus kreisiFortu- R. karachiensis (Maqbool & Ghazala, 1984) n. comb. ner, 1985 (nom. nov. for T. robustusexiguus Kreis, = Orientylus karachiensis Maqbool & Ghazala, 1984 1926) and R. steineri (Stefanski, 1916) Siddiqi, 1986 R. laurentinus Scognamiglio & Talame’, 1973 (= Aphelenchus steineri Stefanski, 1916) were placed by R. leviflexus (Phillips, 1971) Germani, Baldwin, Bell & Siddiqi (1986) in species inquirendae under Rotylenchus. Wu, 1986 = Scutellonema leviflexus Phillips, 1971 R. lobatus Sultan, 1985 R. microstriatus Siddiqi & Corbett, 1983 Discussion : R. nzinutus (Sher, 1964) Germani, Baldwin, Bell & Wu, The name Rotylenchus was first proposed in 1934 1986 in two separate works by Filip’evbut was not described = Scutellonenza nlinutum Sher, 1964 at the time. The name became available only in July R. neorobustus Sultan & Jairajpuri, 1979 R. nexus Ferraz, 1980 1936 when Filip’ev proposed it again in a differentiating key (Sher, 1965). = Calvatylus nexus (Ferraz, 1980) Siddiqi, 1986 R. orientalis Siddiqi & Husain, 1964 The differentiation of Rotylenchus from related = Orientylus orientalis (Siddiqi & Husain, 1964) Jai- genera was difficult at first, partly because the statu of rajpuri & Siddiqi, 1977 the type species was not clear. Sher (1965) reviewed the = Helicotyleizchus orienta@ (Siddiqi & Husain, 1964) discussion about the identity of the type species and Geraert, 1976 accepted R. robustus (de Man, 1876) as the correct type R. peculiaris (Khan & Khan, 1982) n. comb. species. He also synonymized Gottholdsteineria Andras-

Revue Nématol. 10 (2) :219-232 (1987) 223 R. Fortmer __~ sy, 1958 with Rotylenchus and differentiated Rotylenchus from 9-13 pm (mean values) and 5-15 pm (individual from Helicotylenchus primarily by the prominent dori values) in different populations of H. pseudorobustus. A sa1 overlap of the esophageal glands. Sher’s con- similar specific variability in Rotylenchus would make clusions have been universally accepted. impossible in many cases the identification of the genus Recently someRotylenchus spp. were transferred into Orientylus. two new genera, Calvatylus and Onentylus. As already proposed by Zancada and Lima (1986) Calvatylus Jairajpuri & Siddiqi,1977 was differen- Orientylus Jairajpuri & Siddiqi, 1977 is here considered tiated from Rotylenchus by a labial region non offset as a junior synonym of Rotylenchus Filip’ev, 1936. and without annuli norlongitudinal striae, and by a male Some species in Onentylus were placed by Siddiqi , tail shorter than the body width. Ferraz (1980) showed (1986) in a new genus, Varotylus. The two genera are that labial characters may be variable within the same differentiated by the development of the female pos- population, and that short-tailed males are known for terior genital branch, non-functionalor absent in On’en- some species of Rotylenchus S. str. The synonymization tylus S. str., well developed in Varotylus. This charac- of Calvatylus with Rotylenchus proposed by Ferraz ter is not accepted at generic level. (See below the dis- (1980) is accepted here. cussion on the synonymization of Rotylenchoides with Orientylus Jairajpuri & Siddiqi,1977 differs from Helicotylenchus.) Rotylenchus by the more posterior position of the DG0 Varotylus is here proposed as a new junior synonym and by the smaller, narrower lip region. Also there are of Rotylenchus. no longitudinal striae on the basal lip annulus of the species in Onentylus. There are several species of Rotylenchus S. str. with no Scutellonema Andrassy, 1958 striae on thebasal annulus and/orwith narrowhead, for example R. caudaphasrnidius Sher, 1965 and R. brevi- Diagnosis : glans Sher, 1965. The position of the DG0 remains the Hoplolaiminae. Fernale :Body spiral to C-shaped only difference between Rotylenchus and Orientylus. or almost straight. Labial region narrow ‘truncate to The position of DG0 is traditionally given as ratio O. offset rounded; annulated, withor without longitudinal This ratio was shown by Fortuner (1984~)to be un- striae. First labial annulus divided intosix sectors, lateral suited for taxonomic purposes. The actual distance in sectors smaller than the others (SEM). Amphid aper- micrometers betweenstylet base and gland openingwas tures oval betweenlabial disc and lateral sectors. Lateral estimated from holotype measurements,calculated from field with four lines usually areolated near phasmids and figures, or found given in the text, of descriptions of anteriorly, sometimes transverse striae scattered over species of Rotylenchus. The distribution of specific whole field. Labial framework, stylet and stylet knobs values of DG0 in Rotylenchus s.1. ranges from 4 to 15 average sizedfor the family; knobs rounded toindented. pm and peaks at 6 pm. The distribution is heavy tailed DG0 4-8 pm from stylet base. Esophageal gland over- to the right. The proposa1 of Orientylus can be seen as lap dorsal and lateral. Two genital branches outstret- an attempt torestore normality in Rotylenchus by elim- ched, equally developed. Epiptygmapresent. Tai1 short, inating from this genus the species with DG0 equal or rounded; phasmids enlarged(scutella) situated opposite greater than 11 pm. (However, R. exirnius was kept in each other, near anus level. Male :Caudal alae envel- Rotylenchus in spite of its DG0 = 11-13 pm, probably oping tail tip, regular or rarely deeply lobed. No sec- because of its offset labial region with striated basal lip ondary sexual dimorphism. annulus.) There is no reason why the distribution of specific Type species : measurements should be normalin a genus. The distri- Scutellonernablaberum (Steiner, 1937)Andrassy, bution of specific DG0 values in Rotylenchus results 1958. from a trend for posteriorly directed migration of the gland opening. This trend is manifest only in about a Other species : third of the species, but these species are not different See list in Germani et al. (1986). (from The rest of their description) from the species Discussion : where the posterior migration has not yet started. The creation of a separate genus for those speciesis not Scutellonerna is closeto Rotylenchus, as seen by size, justified for systematic purposes. generalappearance, and mostlyarrangement of+e It may also be noted that the acceptationof Orientylus esophageal glands. There are some slight differences is also not justified for generic identification. The range between the two genera, for example, Scutellonerna has of specific values is about 4-12 pm in Rotylenchus S. str, lateral field generally areolated at phasmids and ante- 11-15 pm in Om’entylus. No study has been found in the riorly. However the primary difference between the two literature onintraspecific variability ofDG0 in Rotylen- genera is the size of the phasmids enlarged to scutella chus. In the related genus Helicotylenchus, DG0 varies in Scutellonema, pore-like in Rotylenchus.

224 Revue Nématol. 10 (2) :219-232 (1987) Reappraisal 01 Tylenchina. 8. Hoplolaiwidae

In this aspect Scutellonema is intermediate between A. longistylus (Doucet,1980) n.cornb. Rotylenchus withpore-like, adanalphasmids and Hoplo- = P. lonnistvlus Doucet,1980 laimus where the migration of the scutella anteriorly to = N lon&&lus (Douc& 1980) Siddiqi, 1986 anus is often accompanied by the evolution of other A. luci (Sher, 1964) n. comb. characters. = P. luci Sher, 1964 A. macbethi (Sher, 1964) n. comb. Aorolaimus Sher, 1963 = P. macbethi Sher, 1964 = Peltamigratus Sher, 1964 (n. syn.) A. nigeriensis (Sher, 1964) n. cornb. = = Nectopelta Siddiqi, 1986 (n. syn.) P. nigeriensis Sher, 1964 A. pachyurus (Loof, 1964) n. cornb. Diagnosis : = P. pachyurus Loof, 1964 A. perscitus (Doucet, 1980) n. comb. Hoplolaiminae. Female :Body spiral to C-shaped, = P. perscitus Doucet, 1980 medium sized. Lip region slightly offset or continuous = N. perscita (Doucet, 1980) Siddiqi, 1986 with body, with or without annuli and/or longitudinal A. sheri (Andrassy, 1968) n. comb. striae. Lateral field with four or less incisures. Labial = P. sheri Andrassy, 1968 framework and stylet medium sized; stylet knobs flat- A. striatus (Srnit, 1971) n. cornb. tenedto indented anteriorly. DG0 3-10 pmfrom = P. striatus Srnit, 1971 stylet base. Esophageal glands with threenuclei, overlap A. thornei (Ihobloch, 1969) n. cornb. intestine dorsally and laterally; intestine symmetrically = P. thonxi Knobloch, 1969 arranged between the subventral glands. Two genital A. torpidus Thorne & Malek, 1968 branches outstretched, equally developed. Tai1 short, A. triticeus (Doucet, 1984) n. comb. rounded. Phasmids enlarged to scutella erratically situ- = P. triticeus Doucet, 1984 ated on body, not opposite each other, anterior to anus = N. triticeus (Doucet, 1984) Siddiqi, 1986 level; sometimesone scutellum is anterior to vulva level. Male :Caudal alae enveloping tail, lobed or reg- Discussion : ular. Secondary sexual dimorphism visible in labial region and esophageal structures smaller in males. The original diagnosis of Peltanzigratus used three characters to differentiate this genus from Aorolaimus : Type species : (i) position of phasmids (bot11 between anus and vulva Aorolainlus helicus Sher, 1963. vs one anterior to vulva in Aorolaimus); (ii) lip annu- Other species : lation (lip regionwithout striation w annulipresent A. annulatus (Mulk & Siddiqi, 1982) n. cornb. in Aorolaimus); and (iii) shape of malecaudal alae = Peltamigratus annulatus Mulk & Siddiqi, 1982 (indented 'us regular in Aorolaimus). = Nectopelta annulata (Mulk & Siddiqi, 1982) Siddiqi, Smit (1971) noted that some species, placed in Pel- 1986 tamigratus because of the position of the phasmids, do A. baldus Thorne & Malek, 1968 not follow the generic diagnosis forthe other two A. brevicaudatus (Doucet, 1984) n. cornb. characters. P. christiei and P. holdemani have sometimes = P. brevicaudatus Doucet, 1984 a labial region finely striated; males of P. holdemani and = N. brevicaudata (Doucet, 1984) Siddiqi, 1986 P. machbethi havecaudal alae with only twosmall A. browni (Khan & Zakiuddin, 1968) n. cornb. indentations. Smit added a new species to the genus, P. = P. browni Khan & Zakiuddin, 1968 striatus, with lips markedly annulated and male caudal A: capsici Jiménez-Millan, Arias-Delgado & Fijo, 1964 alae not indented. A. christiei (Golden & Taylor, 1956) n. comb. Doucet (1980) recently described P. Zongistylus with = Rotylenchus christiei Golden & Taylor, 1956 lips annulated and P. perscitus also with lip annulated A. conicori (Doucet, 1984) n. cornb. and with caudal alae not indented. = P. conicori Doucet, 1984 Both Smit(1 97 1) and Doucet(1 980) noted that, of the = N. conicori (Doucet, 1984) Siddiqi, 1986 three diagnostic characters proposed by Sher (1964), A. holdemani (Sher, 1964) n. comb. only the first one position of the scutella - is valid. = P. holdemani Sher, 1984 - A. ibiboca (Monteiro & Choudhury, 1978) n. cornb. The small difference in position of scutella is certainly = P. ibiboca Monteiro & Choudhury, 1978 a goodcharacter for specific identification, but its value A. indicus (Khan, 1972) n. cornb. for the taxonomic differentiation of the genus is ques- = P. indicus Khan, 1972 tionable. The position of the scutella in therelated genus A. intennedius Suryawanshi, 1971 Hoplolainzus may also Vary. Some species in Hoplolai- A. leiotnerus (de Guiran, 1963) de Guiran & Sher, 1969 mus have both scutella posterior to vulva or both anterior = Hoplolaimus leiornerus de Guiran, 1963 to vulva. = A. israeli Sher, 1963 Because a single identifying character is not a Sound A. leipogramtnus Sher, 1963 basis for the definition of a genus, Peltamigratus Sher,

- Revue Nématol. IO (2) :219-232 (1987) 225 R. Fortuner

~

1964 is here proposed as a new synonym of Aorolaimus Other species : Sher, 1963. lis1 in 1981. Siddiqi (1986) proposed a new genus, Nectopelta, for See Luc, sixspecies in Peltamigratus (P. annulatus, [type], P. brevicaudatus, P. conicori, P. longistylus, P. perscitus, and Discussion : P. triticeus). Nectopeltawas differentiated from Peltami- Hoplolaimus was described originally from a single gratus by three characters : i) labial region distinctly specimen of the type species. This specimen was re- annulated; ii) marked by longitudinal grooves and; iii) described by Andrassy in 1954 then lost in 1956 (Sher, areolations of lateral field at phasmid level. 1961). A number of Peltamigratus spp. (S. str.) have labial The genus Nemonchus Cobb, 1913, was thought by region distinctly annulated, particularly so P. striatus Thorne (1935) to be the result of a fiiation artifact. and P. holdemani. Filip’ev and SchuurmansStekhoven (1941) accepted the In the absence of SEM face views, it is difficult to opinion of Thorne and listed Nemonchus galeatus, type prove the lack of longitudinal grooves in Peltamigratus and only species of,Nemonchus, as a synonym of Ho- S. str. Light microscopeface views of severalsuch species plolaimus coronatus. Sher (1 96 1) formally recognized in Sher (1964) do show a six-sectored structure. Evenif Nemonchus as a synonym of Hoplolaimus; transferred further studies prove the absence of grooves in Pelta- Nemonchus galeatus to Hoplolairnus galeatus; and re- migratus S. str., and their presence in Nectopelta, this stored the law of priority by listing H. coronatus Cobb, character may not be acceptable at generic level. Face 1923 as a synonym of H. galeatus (Cobb, 1913). views as seen with SEM can Vary within certain genera Two new genera were recently proposed for species such as Helicotylenchus (Fortuner, 1986) or Hemicyclio- previouslyin Hoplolaimus :Hoplolaimoides Shakil, 1973, phora (Loof, 1986). Use of such a csiterion needlessly and Basirolaimus Shamsi, 1979. would increase the number of genera. Hoplolaimoides was proposedduring theannual Differences in areolation have been reported in many meeting of the National Academy of Science, India, in genera; for example, Scutellonema (Germani et al., 1973. The proceedings of this meeting were printed and 1986). This character canbe used only for species the new name is available according to the provisions of identification. the International Code of Zoological Nomenclature. Nectopelta is heresynonymized with Aorolaimus Hoplolaimoides includes only one species, H. califor- (= Peltamigratus). nicus, distinct from the species in Hoplolaimus S. str. in having both scutella posterior to vulva, an arrangement Hoplolaimus von Daday, 1905 proposed as typical of Peltamigratus (Sher, 1964). Ex- = Newonchus Cobb, 1913 cept forthis posterior location of the scutella, H. califor- = Hoplolaimoides Shakil, 1973 nicus is a typical member of the genus Hoplolaimus and = Basirolaimus Shamsi, 1979 it does not seem justified to create a genus based on a Diagnosis : single character. If Hoplolaimoides were accepted, a second new genus should be proposed for H. puertori- Hoplolaiminae. Female : Body straight, large censis where the position of scutella is alsodifferent from (1-2 mm long). Lip region offset from body,wide, the rest of Hoplolaimus (in H. puertoricensis, both scu- anteriorly flattened, with clearly marked annuli, and tella are anterior to vulva). The position of the scutella with longitudinal striae. Lateral field with four lines or is agood character for identification but does not appear less, generally areolated at level of phasmids and ante- meaningul for the taxonomic differentiation of genera. riorly, sometimes with striae irregularly scattered over Hoplolaimoides is here accepted as a junior synonym of entire field, rarely not areolated. Labial framework and Hoplolaimus as already proposed without argumentsor stylet massive; stylet knobsanchor or tulip-shaped. justification by Siddiqi (1986). DG0 3-10 pmfrom stylet base. Esophagealglands Basirolaimus was established for thespecies in Hoplo- overlap intestine dorsally and laterally; sometimes gland Zaimus with six esophageal gland nuclei. Luc (1981) nuclei duplicated to a total of six nuclei; intestine argued that the duplication of the normal complement symmetrically arranged between the subventral glands. of three nuclei to five or six nuclei was only an intra- Two genital branches outstretched, equally developed. generic variation. Luc (1981) proposed Basirolaimus as Tai1 short, rounded,phasmids enlarged to scutella a junior synonym of Hoplolaimus. erratically situated on body, anteriorly to anuslevel, and Siddiqi (1986) rejected Luc’s (1981) conclusion on the sometimes anterior to vulvalevel, not oppositeeach grounds that the extra nuclei in Basirolaimus result of other. Male : Caudal alae enveloping tail, regular. duplication of the dorsal nucleus alone. This does not Secondary sexual dimorphism visible in labial region detract fromthe validityof Luc’sreasoning (mere and esophageal structures smaller in males. duplication of nuclei is not a valid generic character). Type species : The synonymization of Basirolamus with Hoplolaimus Hoplolaimus tylenchiformis von Daday, 1905. is reinstalled here.

226 Revue Nématol. 10 (2) :219-232 (1987) 1 Reappraisal 01 Tylenchina. 8. Hoplolaimidae

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Antarctylus Sher, 1973 near anus; cephalids andcaudalid present. Tail 1 to 2 1/2 body diameters long, typically more curved dor- Diagnosis : sally, with or without a terminal ventral process, some- Hoplolaiminae. Fernale :Body vermiform, spiral to timesrounded. Stylet and labial framework average- C-shaped. Labial region rounded, continuous, annula- sized. DG0 from 6 to 16 pm from stylet base. Median ted; anterior lipannulus notdivided into sectors (SEM). bulb rounded with average-sized valve. Glands overlap Lateral field with four lines. Phasmids small, near anus. intestine dorsally and ventrally, ail three glands of about Caudalidsnot described. Tailrather long (2-3 body the same length.Two genital branches, the posterior one diameters long), conoid, pointed. Stylet and cephalic sometimes degeneratedor reduced toa PUS. Epiptygma framework average sized. DG0 about 10pm from stylet present but folded inwards, into the vagina. Vulval flaps base in the only species known in this genus. Median present, inconspicuous. Male :Slight secondary sexual bulboval/rounded with average sized valve. Glands dimorphism seen in smaller anterior end. Caudal alae overlapping theintestine; the dorsal gland andone enveloping tail end. subventral gland overlap dorsally, the second subventral gland overlaps ventrally for a short distance. Both sub- Type species : ventralglands are shorter than the dorsal one. Eso- Helicotylenchus dihystera (Cobb, 1893) Sher, 1961. phago-intentinal junction a small triangular structure. = Tylenchus olaae Cobb, 1906* = T. spiralis Cassidy, 1930* Two equally developed genital branches. One or MO = Aphelenchus dubius pemensis Steiner, 1920 epitygma present, inconspicuous. Vulval flap not de- = H. nannus Steiner, 1945 scribed. Male :Slight secondary sexual dimorphism seen = H. crenatus Das, 1960 in smaller anteriorend. Tail with long hyaline end. = H. jlatus Roman, 1965 Caudal alae said to envelop tail end, but seen stopping = H. rotundicauda Sher, 1966 just shortof tail tip inoriginal figure. Gubernaculum not = H. punicae Swarup & Sethi, 1968 described; titillae not figured. = H. glissus Thorne & Malek, 1968 = H. dihysteroides Siddiqi, 1972 Type and only species : = H. teleductus Anderson, 1974 A. humus Sher, 1973. The original type species was H. nannus Steiner, 1945 Discussion : but Sher (1961) proposed H. nannus as a synonym of 19. The only species in thegenus presents al1 the dihystera. characters of Hoplolaiminae, except the female tail, Other species : longer and more pointed than usual. Some Rotylen- See list in Boag and Jairajpuri(1985) and additional chulinae also have long pointed tails. names in Fortuner (1984b). The arrangement of the esophageal glands is similar to thatof Helicotylenchus, but inAntarctylus the subven- Discussion : tral glands are shorter than the dorsal one; al1 three glands are of equal length inHelicotylenchus. Antarctylus The genus Rotylenchoides Whitehead, 1958 has been and mostHelicotylenchus spp. have an individed anterior synonymized with Helicotylenchus by Fortuner (1984 b) lip annulus, different from thesix-sectored lip annulus because these MO genera differ by a single character, of the other Hoplolaiminae(Sher & Bell, 1975). Because namely the regression of the posterior genital branchin of the long pointed tail, similar to tails of some taxa Rotylenchoides. A gradua1 regression of this organ is in Telotylenchinae (Triversus for example), this lone documented in the descriptionsof some species of Heli- species from Antarctica may be seen as a relict of cotylenchus S. 1. :(i)typical Helicotylenchus spp. have two the ancestral forms that evolved into Helicotylenchus. equally developed branches; fii) in some species (for example H. multicinctus)the posterior branch is smaller Helicotylenchus Steiner, 1945 than theanterior one, but is still functional; fiii)RotyZen- = Rotylenchoides Whitehead, 1958 choides intermedius Luc, 1960 and Helicotylenchus neo- = Zimrnermannia Shamsi, 1973 formis Siddiqi & Husain, 1964; have a posterior branch

Diagnosis : Hoplolaiminae. Female :Body vermiform, spiral to * Siddiqi (1986) proposesHelicotylenchus olaae and H. spiralis straight.Labial region continuous to slightly offset, (synonymsof H. dihystera) asnew combinations. In fact, rounded oranteriorly flattened, generally annulated but Tylenchus olaae and T. spiralis were brought into the genus never longitudinally striated; anterior lip annulus gen- Helicotylenchus by Sher (1961), when he synonymized both erally not divided into sectors, with elongate amphid nominal specieswith H. dihystera. This actionmade Sher spertures (SEM). Rarely faint or marked lip sectors are (1961) the authorof the combinationsof olaae and spiraliswith present. Lateral field with four lines. Phasmids small, Helicotylenchus, il and when they are needed.

Revue Nématol. 10 (2) :219-232 (1987) 221 R. Fortuner . reduced to arow of undifferentiated cells;a similar Other species : reduction is also observed in Rotylenchoides desouzai, A. straturatus Germani, 1970 however, because of the arrangement of its esophageal A. variabilis Germani & Luc, 1984 glands, this species is better placed in Rotylenchus, close to Rotylenchus orientalis; and (iv)the species that have Discussion : been proposed as typical representative of Rotylenchoi- Aphasmatylenchus is remarkable by the absence of des, namely R. brevis, R. variocaudatus and R. affinis phasmids andalso bythe conoid elongate tailof the male have a posterior branch reduced to a PUS. This syn- that resembles male tails in Belonolaimidae. The shape onymization was rejected by Siddiqi (1986). This author of the labial region,the well developed stylet and frame- did not realize that the gradua1 regression of the female work, the posterior DGO, the shape of the female tail, posterior genital branch in different species of Helico- are characteristics of Hoplolaimidae and Aphasmatylen- tylenchus S. 1. makes this character irrelevant for generic chus has been acceptedby its original descriptor and by differentiation. (See also discussion in Lucet al., 1987). later authors as a member of this farnily. Zimmermannia was proposed as a sub-genus of Heli- WithinHoplolaiminae, Aphasmatylenchus stands cotylenchus by Shamsi (1973) during a congress. Minu- apart by its vend glandular overlap. The exact arrange- tes of the congress were published accordingto therules ment of the glands has not been described, but seerns of the International Code of Zoological Nomenclature. to differ from both the symmetrical dorsal overlap of The new sub-genus, with H. (Zimmermannia)e ythri- Rotylenchus and the asymmetrical dorsal and ventral nae as type and only species, was characterized by the overlap of Helicotylenchus. In fact theglands in Aphas- male tail, with a terminal process longer than thecaudal matylenchus are somewhat similar to those in Rotylen- alae. The terminal ventral process in some species of chulinae but are shorter than in this subfamily. HeZicotylenchus and other genera is the remnant of the Aphasmatylenchus is know only from West Africa. dorsal asymmetrical regressionof the tail (Fortuner, Because of the absence of any known intermediate 1986~).This remnant can be longer or shorter depen- genus between Aphasmatylenchus andthe ancestral dingon the species andon individual variation (its proto-Hoplolaimidae from which it evolved, it is irn- length varies from 1 to 7 pm in a population of H. possible to propose any hypothesison the origin of the eythrinae from Riverside, California). When longer, it genus. However there is no need to disguise Our ignor- is quite natural that itmay extend past the caudal alae. ance by creating a separate taxon for this genus and the It seems inadequate to proposea separate genus on such subfamily Aphasmatylenchinae proposedby Sher (1965) a character. is not accepted here. Zimmemzannia is here considered as a synonym of Helicotylenchus as already proposed withoutjustification or arguments by Siddiqi (1986), and also by Fortuner The subfamily Rotylenchulinae Husain & Khan, 1967 (1986). DIAGNOSIS Aphasmatylenchus Sher, 1965 Hoplolaimidae. Body of mature females swollen or kidney shaped.Lip region not as high as in Hoplolaimi- Diagnosis : nae. Esophageal glands enlarged into a long lobe over- Hoplolaiminae. Female :Body vermiform, circle to lapping the intestine mostly laterally. Rotylenchulinae C-shaped. Labial regionslightly offset from body, annu- are sessile semi-endoparasites and lay their eggs in a lated, but without longitudinal striae. First labial annu- gelatinous matrix, whereas Hoplolaiminae areectopara- lusdivided into six equal sectors, elongateamphid sites or migratory semi-endoparasitesand lay their eggs apertures (SEM). Lateralfield with four lines. Phasmids free in the soil. absent. Cephalids and caudalid not described. Tail 1.5 to 2 body diameters long, more curved dorsally with DESCRIPTIONOF ROTYLENCHULINAE rounded end. Stylet and labial framework well devel- Lip region high, but lower than typical Hoplolaimi- oped. DG0 about 8 pm from stylet base. Median bulb nae, never striated longitudinally. Lateral field with four rounded.Glands overlap the intestine ventrally and lines. Phasmids always pore-like, near anus or on tail. laterally. Two genital branches equally developed. Epi- Tail short, rounded, or longer, conoid. Labial framework ptygma and vulval flap not described. Mule : Slight and sryh me&um-sized. DGÛ far to very far from secondary sexual dimorphism seen in the smaller an- stylet. Genital branches outstretched or reflexed, pos- terior end. Tail conoid, elongate, with a hyaline end. terior branch sometimes reduced to a PUS. Caudal alae enveloping tail end. Gubernaculum with titillae. Male sexual dimorphism well marked with anterior end smaller than female, sometimes esophagus degener- Type species : ate, nonfunctional. Caudal alae enveloping tail, some- Aphasmatylenchus nigeriensis Sher, 1965. times not reaching tail end.

228 Reappraisal 01 Tylenchina. 8. Hoplolainzidae

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Rotylenchulus and the related genera are quite dif- Cephalids and caudalidnot described. Tai1 short, hemi- ferent from the genera in Hoplolaiminae (see below). spherical. Labial framework and stylet well developed. Originally proposed as a " ... with charac- Stylet knobsslopping backwards inthe only known ters of Rotylenchus... " (Lindford & Oliveira,1940), species. DG0 13 pm from stylet base. Median bulb Rotylenchulus was considered in Pratylenchinaeby strong, rounded. Anterior genital branch outstretched. Thorne (1949) and Tom Goodey (195 l), and in Na- Posterior branchreduced to a PUS. Epiptygma and cobbinae by Chitwood (1950) and J. B. Goodey (1963). vulval flapnot described. Male : Vermiform.Labial It was made the typeof Rotylenchulinae by Husainand region knob-like, high, rounded, distinctly offset. Stylet, Khan, 1967. This subfamily was placed in Hoplolaimi- labial framework and esophagus reducedin size. Caudal dae by the original authorsand by Siddiqi (1971). alae enveloping tail. Juveniles :Tail rounded witha long Golden (1971) placedit inNacobbidae, Heteroderoidea. terminal projection. Fotedarand Handoo (1978) evaded the problemby Biology : elevating Rotylenchulinae to family rank. The resemblance between Rotylenchulus and Nacob- The females are semi-endoparasitic, their anterior bus is probably .due to a convergence. Enlargement of portion is embedded in the root up to the level of the the mature female body and sedentary semi-endopara- excretory pore. Males andlarvae have not been observed sitism are phenomena that occurred in several lines of feeding. The eggs are deposited singly in a gelatinous plant parasitic nematodes. A tendencytowards body matrix. enlargement isalready attested in some anguinids. Type and only species : Sedentary saccate females are known in Hoplolaimidae Acontylus vipriensis Meagher, 1968. (Rotylenchulus), Pratylenchidae (), Criconema- toidea (Tylenchulus), and Heteroderidae. The enlarge- Discussion : ment of the female body is not a reliable character for This genus is represented by a single species found the definition of generic relationships and differen- in two localities in Australia. It is here accepted as a tiation of families. Rotylenchulus should not be placed member of Hoplolaimidae because of the shape of the near Nacobbus. labial region, the strongstylet and labial framework, the Rotylenchulus has some characteristics of Hoplolai- posterior migration of the DGO, and the short rounded midae (see below) but it has evolved so far from the tail. typical hoplolaimids thatis was tempting toaccept it in It is close to the genera in Rotylenchulinae by several a family of its own. However, two genera link Rotylen- characters : i) themature females are swollen and chulus to the rest of the hoplolaimids : Senegalonema, a embedded in theroots; ii) the esophageal glands overlap genus recently described fromAfrica, and Acontylus, an the intestine dorsally and laterally over a long distance; Australian genuswhose taxonomic statuswas heretofore iii) the eggs are laid within a gelatinous matrix; and iv) unsettled. Acontylus has been in the past linked to Ho- the maleshave an esophagealregion reduced and plotylus, a pratylenchid, or placed in a subfamily of its probably nonfunctional. own, Acontylinae. It is proposed to accept it asa The arrangement of the esophageal glands in both Rotylenchulinae. sexes, and the degenerescence of the anterior extremity The inclusion of Senegalonema and Acontylus in of the male are similar to thoseobserved in several Rotylenchulinae makes it possible to accept this sub- genera in Pratylenchidae, namely Radopholus and Ho- family in Hoplolaimidae. This action makes unnecess- plotylus. These genera lack the hoplolaimid characteris- ary the elevation of Rotylenchulinae to family rank and tics that are evident in Acontylus. the creation of Acontylinae. The regression of the posterior genital branch on Verutus Esser, 1981 was placed by Siddiqi (1986), in Acontylus has also been observed in some speciesof Rotylenchulidae. It is considered here to be closer to Helicotylenchus and Rotylenchus. In these two genera the Heteroderidae (as seenamong other characters by arrangement of the esophageal glandsis different to that morphology of male sexual organs). It will be treated in Acontylus, there is no markedsecondary sexual with this later family. dimorphism, mature females are not swollen and do not lay eggs in a gelatinous matrix. THEGENERA IN ROTYLENCHULINAE Senegalonema Germani, Luc €k Baldwin, 1984 Acontylus Meagher, 1968 Description : Diagnosis : Rotylenchulinae. Immature fernale : Body vermi- Rotylenchulinae. Female :Body straight to slightly form,C-shaped. Labial region high, not offset, not ventrally arcuate, swollen in the vulval area. Labial annulated; anterior lip annulus divided into six sectors, region high, slightly offset, trapezoid in lateral outline. four submediansectors triangular, adjacent two by two, Lateral field with four lines. Phasmids at anus level. dorsally and ventrally. Lateral field withfour lines.

Revue Nématol. IO (2) :219-232 (1987) 229 R. Fortuner

Phasmids pore-like near anus level. Tail conoid, 2 to 2.5 Type and only species : body anal diameter long, more than half its length Senegalonema sorghi Germani, Luc & Baldwin, nonprotoplasmic.Labial framework and stylet me- 1984. dium-sized. DG0 about 5-7 pm from stylet base in the only known species. Medianbulb oval,with strong valve. Glandular overlap mostlylateral, sometimes ven- Discussion : tral. Two genital branches outstretched. Maturefernales : Body posteriorly swollen, variously shaped to kidney- The relationships of this genus with Rotylenchulus shaped, annulated. Phasmidsenlarged. DG0 at 5-8 pm and other genera have been discussed in detail by the from stylet. Gland overlap expandedlaterally. Tail short, original authors (Germani, Luc & Baldwin, 1984). Sene- conical. Vulvapostequatorial. Two genital branches very galonema differs from Rotylenchulus by shape of lip long, convoluted. Male :Slight secondary sexual dimor- region, position of DGO, outstretched genital branches phism seen in smaller anterior end. Esophagus func- inimmature females, nondegenerate anterior end of tional. Caudal alae enveloping tail end.Gubernacu- males, and caudal alae and titillae of males. The genus lum with titillae. shares some characteristics with some generain Hetero- Malesand immature females free in soil. Mature deridae, which points towardsa common origin of this females semi-endoparasitic, sessile in roots. family and Hoplolaimidae.

Table 1 Tabular key to the genera in Hoplolaimidae (Females)

Body Glands DG0 (Pm) Tail Phasrnids Genital branches

Parurotylenchus vermiform no overlap 4-7 short, pore-like 2 more curved D near anus Rotylenchus vermiform D. overlap 6-16 short, more curved D pore-like 2 or 1 to rounded near anus Scutellonema vermiform D. overlap 4-8 short, scutella 2 rounded near anus Aorolairnus vermiform D. overlap 3-10 short, scutella 2 rounded on body HopIolaimus vermiform D. overlap 3-10 short, scutella 2 rounded on body Antarctylus vermiform D + V, unequal 10 long (2-3 0) pore-like 2 overlap conoid, near anus pointed Helicotylenchus vermiform D + V, equd 6-16 short, pore-like 2 or 1 overlap more curved D near anus to round Aphasrnatylenchus vermiform short lateral 8 short, no phasmids 2 overlap more curved D Acontylus swollen at long lateral 13 short, pore-like 1 vulva overlap rounded near anus Senegalonerna kidney- long lateral 5-7 short, large 2, convoluted shaped overlap conoid Senegalonerna vermiform long lateral 5-7 long (2-2.5 W) pore-like 2, straight (immature female) overlap conoid near anus Rotylenchulus kidney- long lateral 13-33 short, pore-like 2, convoluted shaped overlap conoid on tail Rotylenchulus vermiform long lateral 13-33 long (2-3 0) pore-like 2, reflexed (immature female) overlap conoid on tail

230 Nérnatol. Revue 10 (2) :219-232 (1987) Reappraisal 01 Tylenchina. 8. Hoplolaimidae

Rotylenchulus Linford & Oliveira, 1940 REFERENCESI) = Spyrotylenchus (= Spirotylenchus) Lordello & Cesnik, 1958. ANDRASSY, 1. (1985). A dozennew species from = Leiperotylenchus Das, 1960 Hungary. Opusc. 2001.Bpest, 19-20 : 3-39. BALDWIN,J. G. & BELL,A. H. (1981). Pararotylenchus n. gen. Diagnosis : (Pararotylenchinae n. subfam., Hoplolaimidae) withsix new Rotylenchulinae. Immature female : Body vermi- speciesand two new combinations. J. Nematol., 13 : form, spiral to C-shaped. Labial region variable, from 11 1-128. low rounded to high flattened, not offset, annulated or BOAG, B.& JAIRAJPURI, M. S. (1985). Helicotylenchus scoticus not annulated; anterior lipannulus dividedinto six n. sp. and a conspectus of the genus Helicotylenchus Stei- sectors of about the same size. Lateral field with four ner,1945 ( : Nematoda). Syst. Parasitol., 7 : lines. Phasmids at mid-rail. Tail 2 to 3 body anal di- 47-58. ameters long, conoid with rounded end. Labial frame- CHITWOOD, B. G. (1950). General structure of nematodes.In : work and stylet strong. DG0 13 to 33 pm from stylet Chitwood, B. G. & Chitwood, M. B. (Eds) An Introduction base. Median bulb oval with strong valve. Glandular to Nenzatology, Section1: Anatonly, Baltimore, Monumental overlap very long, mostly lateral. Two genital branches Printing Co : 7-27. opposed, with double flexure. Mature female : Body obese, kidney-shaped. Cuticle thick. Tail conical pointed DOUCET,M. E. (1980). Description de deux nouveaux Pelta- with or without hyaline extremity. Ovaries very long, migratus et d'une population d' (Ne- : &- convoluted. Male :Vermiform. Anterior end reduced. rnatoda Tylenchida)de la Province de Cordoba, gentine. Nematologica, 26 : 34-46. Caudal alae difficult to see, not quite reaching tail end. EROSHENKO,A. S. (1984).[Plant nematodes of coniferous Type species : plants in the Primorsk Territory.]In Parazity zhiuotnykh i Rotylenchulus renifornlis Linford & Oliveira, 1940. rastenii. Vladivostok, USSR, Akad. Nauk SSSR : 87-97. FERRAZ,S. (1980). Description Rotylenchus nexus n. sp. Other species : of (Nematoda : Hoplolaiminae) from Brazil, with some ob- Rotylenchulus anamictus Dasgupta, Raski & Sher, 1968 servations on the nematode genus Calvatylus. Syst. Parasi- R. borealis Loof & Oostenbrink, 1962 td., 2 : 21-24. R. clavicaudatus Dasgupta, Raski & Sher, 1968 R. Ieptus Dasgupta, Raski & Sher, 1968 FILIP'EV,1. N. (1934). The classificationof the free-living R. macrodoratus Dasgupta, Raski & Sher, 1968 nematodes and their relation to the parasitic nematodes. R. nzacrosoma Dasgupta, Raski & Sher, 1968 Smithson. misc. Collect., 89, 63 p. R. parvus (Williams, 1960) Sher, 1961 FILIP'EV,1. N. & SCHUURMANS STEKHOVEN,J. H. (1941). A = Helicotylenchus parvus Williams, 1960 Manual of Agricultural Helnzinthology. Brill, Leiden, 878 p. R. sacchari Van der Berg & Spaull, 1981 R. variabilis Dasgupta, Raski & Sher, 1968 FORTUNER,R. (1984~). Morphometrical variability Helicoty-in lenchus Steiner, 1945. 5 : On the validity of ratios. Revue NématoL, .7 : 137-146. Generic identification FORTUNER,R. (1984b). Morphometrical variabilityHelicoty- in lenchus Steiner, 1945. 6 : Value of the characters used for TO help with the practical differentiation of the genera specific identification. Revue Nématol., 7 : 244-263. discussed above, a tabular key for generic identification in Hoplolaimidae is presented in Table 1. FORTUNER,R. (1984~). List and status of thegenera and families,of plant-parasitic nematodes. Helnzinth,Abstr., Ser. B, 53 : 87-133. Note FORTUNER,R. (1986). Variabilité et identifcation des espèces chezlesnématodes d_u genre Helicotylenchus.Thèse Andrassy (1985) has proposed a new genus in Ho- Doct. État, Univ. Claude Bernard-Lyon 1, xv 223 p. plolaimidae forHoplorhynchus riparius.Luc (1986) gave + good evidence that this species belongs to the genus FOTEDAR,D. N. & HANDOO,Z. A. (1978). A revised scheme Pratylenchoides Winslow (Pratylenchidae), and he pro- of classification to order Tylenchida. Thorne, 1949. (Nema- posed Hoplorhynchus as a junior synonym of the genus toda.) J. Sci., Univ. Kashmir, 3 (1975) : 55-82. Pratylenchoides. GERMANI,G., Luc, M. & BALDWIN,J. G. (1984). A new Eroshenko (1984) proposed two new genera in a new Rotylenchulinae : Senegalonenla sorghi n. gen., n. sp. (Ne- family in Hoplolaimoidea (Interrotylenchidae with- In- matoda : Tylenchida). Revue Nématol., 7 : 49-56. terrotylenchinae : Interrotylenchus, and Scutellonemoi- GEWANI, G., BALDWIN,J. G.,BELL, A. H. & Wu, X. Y. dinae : Scutellonenloides), for taxa previously in Rotylen- (1986). Revision ofthe genus Scutellonenza Andrassy, 1958 chus. This article has notbeen seen and cannot be (Nematoda : Tylenchida). RevueNématol., 8(1985) : commented on here. 289-320.

Revue Nématol. 10 (2) :219-232 (1987) 23 1 R. Fortuner

GOLDEN, A. M. (197 1). Classification of the genera and higherSHAMSI, M. A. (1973). Taxonomic notes on the genusHelicoty- categories of the order Tylenchida (Nematoda). In : Zuc- lenchus Steiner, 1945 (Nematoda : Rotylenchinae) with a kerman, B. N., Mai, W. F. & Rohde, R. A. (Eds). Plant proposa1 forZimmemannia n. sub-genus. Proc. natn. Acad. parasitic nematodes, Vol. 1, morphology, anatomy, , Sci., India, Biol. Sci., 43 : 80. and ecology. New York, Academic Press : 191-232. SHER,S. A. (1961). Revision of the Hoplolaiminae (Nema- GOODEY,T. (1951). Soil and freshwater nematodes - a mono- toda). 1. Classification of nominalgenera and nominal graph. London, Methuen & Co., Ltd., 390 p. species. Nematologica, 6 : 155-169. GOODEY,T. (1963). Soil and freshwater nematodes. 2nd ed., SHER,S. A. (1964). Revision of the Hoplolaiminae (Nema- rev. J. B. Goodey, London, Methuen & Co., Ltd., 544 p. toda). IV. Peltamigratus n.gen. Nematologica, 9(1963) : , HUSAIN,S. 1. & KHAN,A. M. (1967). A new subfamily, a new 455-467. subgenus, and eight new species of nematodes from India SHER,S. A. (1965). Revision of the Hoplolaiminae (Nema- belonging to superfamily Tylenchoidea.Proc. helminth. Soc. toda). V. Rotylenchus Filipjev,1936. Nematologica, 11 : Wash., 34 : 175-186. 173-198. s’JACOB, J. J. (1960). Hoplotylus femina n. g., n. sp. (Pratylen- SHER,S. A. & BELL,A. H. (1975). Scanning electron micro- chinae : Tylenchida) associated with ornamental trees. Ne- graphs of the anterior region of some species of Tylen- matologica, 4 (1959) : 320-323. choidea (Tylenchida : Nematoda). J. Nevnatol., 7 : 69-83. IZAPOOR,M. (1982). Taxonomic studies on nematodes,of some SIDDIQI,M. R. (1971).Structure of theesophagus in the medicinal and aromatic plantsofNorth India. Thesis, Panjab classification of the superfamily Tylenchoidea (Nematoda). Univ., Chandigarh, India, x 418 p. + Indian J. Nematol., 1 : 25-43. LINFORD,M. B. & OLIVEIRA,J. M. (1940). Rotylenchulus re- niformis nov. gen., n. sp., a nematode parasiteof roots. Proc. SIDDIQI,M. R. (1986). Tylenchidaparasites of plants and helminth. Soc. Wash., 7 : 35-42. insects. Slough, UK, Commonwealth Agricultural Bureaux, IX + 645 p. LOOF,P. A. A. (1986). Scanning electron microscope studies onthe genus Hemicycliophora deMan, 1921 sensu lato SMIT,J. J. (1971). Deux nouvelles espèces africaines d’Ho- (Nematoda : Criconematoidea). Revue Nématol., 8 (1985) : plolaiminae (Nematoda:Tylenchoidea). Peltamigratus stria- 113-123. tus n. sp.et Scutellonerna afFicanumn. sp.Nematologica, 17 : 113-126. Luc, M. (1981). Basirolaimus Shamsi, 1979, a junior synonym of Hoplolaimus von Daday, 1905 (Nematoda : Tylenchida) TAYLOR,A. L. (1 936). The genera and speciesof the Cricone- Nematol. medit., 9 : 197-199. matinae, a subfamily of the Anguillulinidae (Nematoda). Trans. Am. microsc. Soc., 55 : 391-421. Luc, M. (1986). Hophorhynchus Andrassy,1985, a junior synonym of Pratylenchoides Winslow,1958 (Nemata : THORNE,G. (1935). Notes on free-living and plant parasitic Pratylenchidae). Revue Nématol., 9 : 198. nematodes, II. l‘roc. helminth. Soc. Wash., 2 : 96-98. Luc, M., MAGGENTI, A. R., FORTUNER,R., RASIZI, D. J., & THORNE,G. (1949). On the classification of the Tylenchida, GERAERT,E. (1987). A reappraisal of Tylenchina (Nemata). new order (Nemata: Phasmidia). Pm. helminth. Soc. Wash., 1.For a new approach of the taxonomy of Tylenchina. 16 : 37-73. Revue Nématol., 10 : 127-134. WIESER, W. (1953). Die Beziehung zwischen Mundhohlenges- MAGGENTI,A. R., Luc, M., RASKI, D. J., FORTUNER,R., & talt,Ernahrungsweise und Vorkommen bei freilebenden GERAERT,E. (1987). A reappraisalof Tylenchina (Nemata). marinen Nematoden. Ark, Zool. (Ser. 2), 4 : 439-484. 2. The suborder Tylenchina.Revue Néwzatol., 10 : 135-142. ZANCADA,M. C. & LIMAM. B. (1986). Numerical taxonomy PARAMONOV,A. A. (1967). [A critical review of the suborder of the genera Rotylenchus Filipjev,1936 and Orientylus Tylenchina (Filip’ev, 1934) Nematoda: .] Akad. Jairajpuri & Siddiqi, 1977 (Nematoda : Tylenchida). Ne- Nuuk, SSSR Tmdy gehint.Lab., 18 : 78-101. matologica, 31 (1985) : 44-61.

Accepté pour publication le 29 aotît 1986.

232 Revue Nématol. 10 (2) :219-232 (1987)