A reappraisal of Tylenchina (Nemata). 6. The family Belonolaimidae Whitehead, 1960 (l) Renaud FORTUNER* and Michel LUC** California Department of Food & Agriculture, Analysis & Identification, 1220 N Street, Sacramento, CA 95814, USA, and Muséum national d’Histoire naturelle, Laboratoire des Vers, 61, rue de Buffopz, 75005 Paris, Prame. SUMMARY The family Belonolaimidae is redefinedto include two subfamilies, Belonolaiminae withBelonolainlus, Carphodorus, Morulaimus, Geocenamus, and Sauertylenchus, andTelotylenchinae with Tylenchorhynchus (= Telotylenchus),Trophurus, Trichotylenchus, Nagelus, Paratrophurus, Merlinius, Triversus, and Anzplinzerlinius.The following synonymizations are proposed : Ibipora = Mo- rulaimus, Hexadoms = Geocenamus, Uliginotylenchus = Trichotylenchus, Meiodorus and Mulveyotus = Triversus, Histotylenchus and Telotylenchoides = Paratrophurus. The synonymy of Tylenchorhynchusincludes nine generic names.Doliclzorhynchus is a junior homonym. Tetylenchus is kept in genera dubia vel incertae sedis. A tabular key is proposed for identification of the genera in the family. RESUME Réévaluation des Tylenchina (Nemata). 5. La famille des Belonolainzidae Whitehea4 1960 La famille des Belonolaimidae est redéfinie avec deux sous-familles: Belonolaiminae, comprenant Belonolaimus, Carphodorus, Morulaimus,Geocenanzus et Sauertylenchus etTelotylenchinae, comprenant Tylenchorhynchus (= Telotylenchus), Trophuws, Trichotylenchus,Nagelus, Paratrophurus, Medinius, Triversus et Amnplinlerlinius. Lessynonymies suivantes sont proposées : Ibipora = Mondaimus, Hexadorus = Geocenamus,Uliginotylenchus = Trichotylenchus,Meiodorus et Mulveyotus = Triversus, Histotylenchus et Telotylenchoides = Paratrophurus. La synonymie deTylenchorhynchuscomprend neuf noms génériques.Dolichor- hynchus est un homonyme plus récent. Tetylenchtls est classé comme genus dubium. Une clef tabulaire est proposée pour aider à l’identification des genres de la famille. The family Belonolaimidae is here redefined to in- A few months later, Siddiqi (1960) independently pro- clude two groups of taxa that untilnow were considered posed Telotylenchinae for a new genus, Telotylenchus, to be distinct : the belonolaimids (Belonolaiminae, and for Pseudhalenchus, both with overlapping glands. Telotylenchinae) with overlapping oesophageal glands, The structure of the glandular oesophagus was empha- and the tylenchorhynchids (Tylenchorhynchinae, Mer- sized again by the same author (Siddiqi, 1971~)when liniinae, Trophurinae) withglands abutting and he placed Tylenchorhynchinae and related subfamilies bulb-shaped. with abutting glands in Dolichodoridae, while Telotylen- Belonolaimus was placed by Chitwood (1950) in the chinae was placed in Belonolaimidae. family Dolichodoridae. Thorne (1949) did not include It is Our opinion that there is no structural difference this genus inhis revision of Tylenchida, but Loof (1958), betweenforms with abutting glands(the so-called using the system of Thorne and its emphasis on the “ bulb ”), and forms with glands overlapping the an- aspect of the glandular partof the oesophagus, separated terior part of the intestine. These forms differ only in Belonolaimus from Dolichodorus byclassifying, them the lengthof the glands, and in theposition of the oeso- into two different subfamilies (Hoplolaiminaeand phageal lumen relative to the glands (Seinhorst, 1971). Tylenchinae, respectively). The twokinds of arrangementsmay Co-exist in the When Whitehead (1960) proposed a new subfamily, same family(for example, Pararotylenchus is a Hoplolai- Belonolaiminae, heincluded init only generawith midae with abutting glands), in the same genus (Pmyletz- glandularoverlap (Belonolaimus and Trichotylenchus). choides magnicauda has abutting glands, while P. ritteri (1) This article is part of a study on the classification of Tylenchina by the present authors and D. J. Raski, A. R. Maggenti (University of California, Davis) and E. Geraert (Rijksuniversiteit, Gent). * Associate in the Division of Nematology, University of California, Davis, USA. ** Nematologist fiom ORSTOM. Revue Nénzatol. 10 (2) : 183-202 (1987) 183 t R. Fortuner & M. Luc has a longoverlapping lobe) andeven inthe same biology of the various genera in the family are best species, for example Ditylenchus myceliophagus as dis- treated at subfamily level. Telotylenchinae are surface cussed by Fortuner (1982). grazers feeding on the epidermal cells of plant roots. Once admitted that morphology, size, and location of Belonolaiminae have a long stylet that enables them to oesophageal glands mayVary within anytaxa, including reach deeper into the root while the rest of their body families, it becomes evident that belonolaimids, tylen- remains outside of the plant. chorhynchids, and related groups belong to the same family. The subfamily Telotylenchinae Telotylenchinae Siddiqi, 1960 The family Belonolaimidae = Tylenchorhynchinae Eliava, 1964 (n. syn.) Belonolaimidae Whitehead, 1960 = Trophurinae Paramonov, 1967 (n. syn.) = = Telotylenchidae Siddiqi, 1960 Tetylenchinae Siddiqi, 1970 = Tylenchorhynchidae Eliava, 1974 = Merliniinae Siddiqi, 1971 = Meiodorinae Siddiqi, 1971 (n. syn.) DIAGNOSIS DIAGNOSIS Tylenchoidea. Medium to large sized nematodes, with tail cylindroid to conoid, more than twice as long Belonolaimidae. Cephalicframework with weak to medium sclerotization. Stylet 15 to 40 pm long, with as wide but never elongate filiform (typically c' = 2-5). Phasmids always on posterior half of tail,never en- cone about as long as shaft. Corpus not enlarged and larged into scutella. Deirids presentor absent. Face view metacorporal valveof mediumdevelopment. Labial as seen with SEM either ancestral (first lip annulus regioncontinuous or with slight indentation, never six-sectored) or with lateral sectors regressed and face bulbous. SEM face viewwith six lip sectors or with viewevolving towards either agrossly quadrangular lateral lip sectors regressed. Labial disc lemon shapedor shape ora four leaf clover shape. Females typically with variously fused with lip sectors. Disc and lip sectors two genital branches (except Trophurus). Columned sometimesfused together. Sensillae openings often uterus with three rows of cells. Males with peloderan visible on the sub-median lip sectors. caudal alae, rarelylobed or stopping just short of the tail tip. Spicules with or without pronounced velum. TYPE GENUS Belonolaimidae are migratory ectoparasites of plant Tylenchorhynchus Cobb, 1913 roots. A few species are endoparasitic. = Bitylenchus Filip'ev, 1934 = Telotylenchus Siddiqi, 1960 (n. syn.) TYPESUBFAMILY = Quinisulcius Siddiqi, 1971 (n. syn.) = "Dolichorhynchus "(primary homonym; n: syn.) Belonolaiminae Whitehead, 1960 = Trilineellus Lewis & Golden, 1981 (n. syn.) = Divittus Jairajpuri, 1984 OTHERSUBFAMILY = Morasinema Javed, 1984 Telotylenchinae Siddiqi, 1960 = Tessellus Jairajpuri & Hunt, 1984 = Neodolichorhynchus Jairajpuri & Hunt, 1984 (n. COMMENTS SYn.1 The family Belonolaimidae occupiesan intennediate OTHER GENERA position between Tylenchidae and Hoplolaimidae. It is Trophurus Loof, 1956 differentiated from Tylenchidae and Dolichodoridae by = Clavaurotylenchus Caveness, 1958 its columned uterus with three rows of cells (four rows Trichotylenchus Whitehead, 1960 in the othertwo families). It differs from Hoplolaimidae = Uliginotylenchzu Siddiqi, 1971 (n. syn.) by having a longer tail, and by having phasmids always Nagelus Thorne & Malek, 1968 on tail. Paratrophurus Arias, 1970 Telotylenchinae is accepted only as a subfamily in = Histotylenchus Siddiqi, 1971 (n. syn.) Beionoiaimidaebecause of themany similarities be- = Telotylenchoides Siddiqi, 1971 tween belonolaimidsand tylenchorhynchids. Faceviews Merlinius Siddiqi, 1970 areoften similar amongst various members of both = Scutylenchus Jairajpuri, 1971 groups (for example, Momlaimus is similar to Merlinius Triversus Sher, 1974 for this character); the cylindroid tail of belonolaimids = Meiodonls Siddiqi, 1976 (n. syn.) is similar to that of Paratrophurus and Amplimerlinius. = Mulveyotus Anderson & Ebsary, 1982 The few differences that exist in morphology and Amplimerlinius Siddiqi, 1976 184 Revue Nématol. 10 (2) : 183-202 (1987) Reappraisal of Tylenchina. 6. Belonolaimidae GENUSDUBIUM The genera in Telotylenchinae Tetylenchus Filip’ev, 1936 Triversus Sher, 1974 DIAGNOSIS COMMENTSON REJECTED SUBFAMILIES Telotylenchinae. Body about 1 mm long. Lip region Tetylenchinae must berejected after the placement of low, flattened. SEM face view with labial disc and first its type genus in genus dubium (see below). labial annulus fused together; rounded amphid aper- Tylenchorhynchinae is rejected because of the syn- tures often conspicuous. Stylet 11-25 pm long, robust ‘ onymization of Tylenchorhynchus and Telotylenchus. Ty- or with needle-like cone. Labial frameworkthin, weakly lenchorhynchus is the senior synonymat generic level, but sclerotized, with wide basal ring. Metacorporal bulb Telotylenchinae is the senior subfamily and it should be fusiform. Lateral field with four or three lines. Female accepted in accordance withArticle 40 (a) of the Inter- tail conoid, pointed, medium to long(c’ = 3-8). Deirids national Code of Zoological Nomenclature. absent. Male caudal alae sometimes trilobed. Merlinius, the typegenus of Merliniinae, is here accepted as a valid taxon. Merliniinae was differentiated TYPESPECIES by Siddiqi (1971a, 1979) as having six lines in lateral Triversus annulatus (Merny, 1964) Sher, 1974 field, spicules cylindroid with distal end rounded and = Tetylenchus