A reappraisal of Tylenchina (Nemata). 3. The family Orley, 1880 (l)

Étienne GERAERTand DEWEYJ. RASKI Rijksuniversiteit Gent, Instituut voor Dierkunde, K. L. Ledegamkstraat 35, 9000 Gent, Belgiuwz and Division of Nenzatology, University of California, Davis, CA 95616, USA.

SUMMARY The family Tylenchidae contains those tylenchs with the relatively shortest, not overlapping oesophagus and shortest, delicate stylet. Female reproductive system predominantly monodelphic, prodelphic, rarely amphidelphic. Tai1 long, conoid to filiform. Caudal alae adanal, small.It is the only family where amphidial apertures can be on the of lateral the head side and where the anterior conical part of the stylet can be much shorter than the posterior cylindrical part. As subfamilies we recognize : Tylenchinae, Ecphyadophorinae, Tylodorinae; Atylenchinae and Boleodorinae. Four families are synonymized with the Tylenchidae and ten subfamilies are synonymized with the five subfamilies mentioned.

RBSUMB

Réévaluation des Tylenchina (Nemata). 3. La fande des Tylenchidae Orley, 1880

La famille des Tylenchidae comprend les Tylenchides présentant : un œsophage très court et ne recouvrant pas distalement l'intestin, un stylet fin et très court, un système reproducteur femelle très généralement monodelphique, prodelphique, plus rarement amphidelphique; queue longue, conoïde9 filiforme; ailes caudales adanales, peu développées. C'est la seule familleoù les ouvertures amphidiennes peuvent être situées sur le côté de la région labiale et où le cône du stylet peut être nettement plus court que la hampe. Les sous-familles reconnues valides par les auteurs sont : Tylenchinae, Ecphyadophorinae, Tylodorinae, Atylenchinae et Boleodorinae. Quatre familles sont synonymisées avec les Tylenchidae et dix sous-familles avec les cinq sous- familles citées.

An historical review of the Tylenchidaewas published as in other families which must bereviewed as possible in1983 (Raski & Maggenti, 1983) which traced the members of Tylenchidae. As examples,Antarctenchinae changing concepts of that taxon from the time Orley was proposed by Spaull (1972) as a subfamily in the (1880) first proposed it. There were few subfamiliesand Dolichodoridae and latertransfered to Boleodoridae genera in Tylenchidae until recent years. In fact Thor- (Brzeski & Sauer, 1982); Campbellenchinae in the ne's (1949) proposal for had three subfam- Tylodoridae (Wouts,1977); Pleurotylenchinae first ilies in the Tylenchidae only one of which, Tylenchinae, placed inthe Atylenchidae (Andrassy,1976) later remains in this family. There were, of course,some more transfered to Tylodoridae (Wouts, 1977); Psilenchinae genera and subfamilies in the Neotylenchoidea that are originally placed in Tylenchidae (Paramonov, 1968)later now being transfered to the Tylenchidae, but even transfered to Boleodoridae (Brzeski & Sauer,1983); takingthose into account there were few suchhigh similarly Boleodorinae first proposedin Neotylenchidae categories in those days. (Khan, 1964) transfered to Tylenchidae (Geraert, 1971) Since the 1940's there has beena tremendous increase and finally to Boleodoridae (Brzeski & Sauer,1983); in proposals for new genera, subfamilies and families Ecphyadophorinae and Atylenchinae each had separate taxa. These have been proposed in Tylenchidae as well family status as well.

(1) This article is part of a studyon the classificationof Tylenchina by the present authors and R. Fortuner (CDFA, Sacramento), M. Luc (ORSTOM, Paris) and A. R. Maggenti (University of California, Davis).

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Family Tylenchidae Orley, 1980 absent. Phasmid-like structures present on somespecies, usually advulval and dorsad near lateral field; small, = Atylenchidae Skarbilovich, 1959 rounded, coveredby cuticle, not an aperture; rarely = Ecphyadophoridae Skarbilovich, 1959 pore-like, on tail (Anturctenchus, Atetylenchus). Procor- = Boleodoridae Khan, i964 pus elongate; median bulb slender-spindle-shaped, ap- = Tylodoridae Paramonov, 1967 parentlynon-muscular in many;rounded, muscular well-developed in others; valvular apparatus variously TYPESUBFAMILY : developed. Isthmus long, slender; oesophageal glands, Tylenchinae Orley, 1880 symmetrically swollen,usually pyriform rarely with short irregularly-lobed overlap of intestine (except = Dactylotylenchinae Wu, 1969 Epi- churinema andsome Cephalenchus). Small cardia = Duosulciinae Siddlqi, 1979 syn. n. usually projecting into intestine. Excretory poreusually near hemizonid and nerve ring at isthmus, rarely more OTHERSUBFAMILIES : posteriad; occasionally exceptionalin prominentcuticu- Atylenchinae Skarbilovich, 1959 larization. Spermatheca small or large, rounded, ovate = Aglenchinae Siddiqui & Khan, 1983 syn. n. or elongated; axial or offset; sperms small (less than = Antarctenchinae Spaull, 1972 syn. n. 2 Pm), rounded.Female reproductive system single, = Pleurotylenchinae Andrassy, 1976 anteriorly directed; rarely two, amphidelphic (Anturcten- Ecphyadophorinae Skarbilovich, 1959 chus, Atetylenchus, Psilenchus).Uterus quadricolumellar. = Ecphyadophoroidinae Siddiqi, 1986 syn. n. Post-uterine branch usually present, shorter than one = Epicharinematinae Maqbool & Shahina, 1985 vulval body diameter(VBD); sometimes lacking. Spicu- syn. n. les slender, ventrally arcuate, cephalated; gubernaculum Boleodorinae Khan, 1964 simple, trough-shaped.Caudal alae usually adanal; = Basiriinae Decker, 1972 never extending to terminus; rarely asymmetrical (one = Leipotylenchinae Sher, 1974 syn. n. side, usually the right one, shorter than other); occa- = Psilenchinae Paramonov, 1967 sionally reduced (Polenchus)or totally lacking (Miculen- Tylodorinae Paramonov, 1967 chus). Tail elongate-conoid, mostly narrowing to long slender, filiform, even hair-like outline, similar in both = CampbellenchinaeWouts, 1978 ' sexes. Sexual dimorphismon labial plate and amphidial = Eutylenchinae Siddiqi, 1986 syn. n. apertures sometimes present.

Description of the family Tylenchidae Diagnosis Small (from 0.33 mm) to medium (up to 1.3 mm) . Usually slender, elongate, small spe- sized, a few much larger; most always slender, vermi- cies. Sexes similar. Stylet usually small, delicate. Oeso- form in shape. Body marked with manytransverse striae phagus with slender procorpus, median bulb more or (some species appear smooth when seenby light micro- less developed; long, slender isthmus,oesophageal scopy but some smae have been found in everycase glands symmetrically arranged, pyriform,rarely with where study by SEM has been possible). Lateral field slight'overlapping of intestine. One female genital with O, 2, 3, 4, 6 or multiple longitudinal lines (some- branch, anteriorly developed; rarely two branches; col- times so delicate or shallow to be seen only by SEM). umneduterus with four rows. Tail long, conoid to Lip regionusually elevated, rounded; transversestriae filiform. Caudal alae adanal, small, occasionallylacking. usually extend up ont0 labial region, some species with smooth labial region. Labial frameworkdelicate, weakly Relationships developped (moderately developed in Anturctenchus). Stylet mostly small (3 pm-20 pm),delicate, with Most of the characteristics of the species of Tylenchi- distinct basal knobs (rarely without knobs,a few species dae conform with thedefinition of primitivity as covered with long to very long stylet). Conical part of stylet in Luc et al. (1987) : small stylet; delicate labial frame- shorter, as long as or longer than posterior part. Amphid work; amphid openings fromsmall to elongate, sinuous apemes quite variable; mostly elongated slits extend- or straight extending posteriaci longitudinally; median ing posteriad laterally, or arc-shaped to rounded pits on bulb spindle shaped, small, delicateor rounded, muscu- labial plate; rarely small oblique slits on labial plate. lar; oesophageal glands symmetrically arranged, pyri- Four cephalic sensillae or remnants thereof present on form; tail elongate,long. Little is known of the life some species. Oral disc or plate sometimes distinctly history of most species but inso far as known theyhave elevated, rounded; rarely six-lobed (Busiria) withevi- no specialized cycles, resting or resistant stages. Yet it dence of six (inner?) labial papillae. Deirids present or is one of the mostdiverse of the Tylenchina; apparently

144 Revue Nématol. 10 (2) :143-161 (1987) - Reappraisal of Tylenchina. 3. Tylenchidae actively evolving in some characters suchas the amphid tral (Triversus Sher, 1974 and Amplimerlinius icarus, moving from large elongate post labial varying to arc- " belonolaimids ", have, however, a somewhat similar shaped or oval pits limited to the labial plate or even en-face view). small elliptical apertures near the oral opening(Miculen- Genera with this labial plate are : Aglenchus, Coslen- chus, Ecphyadophora) with many links and relationships chus, Antarctenchus,Atylenchus (with largecephalic with other groups (i.e. Tylodorus to the dolichodorids; setae). Filenchus to the anguinids, etc.). II. A second groupdiffers because the amphidial It is most closely related to the Anguinidaewhich were apertures are not confined to the oral plate but continue considered part of the Tylenchidae until recent years on thelateral side aslongitudinal slits; in some theseslits (Siddiqi, 1971) and differ by the smallelliptical amphids are only on the lateral side (Irantylenchus). The region in Anguinidae, which also have elongated, axial sperma- immediately surrounding the mouth opening can be theca, spermslarge with prominent cytoplasm, long slightly raisedbut thereis no oraldisc. The end-on view post-uterine branch (more than two body diameters at can be quadrangular or dorso-ventrally flattened. vulva level). Genera belongingto this group : Irantylenchus, Tylen- chus, Malenchus (the entire headis dorso-ventrally flat- tened), Filenchus (several species now included in Filen- chus have the amphidial apertures within the oralplate). Comments on morphology of Tylenchidae III. A third group has slit-like amphidial apertures confined to theoral plate but slitsthe are dorso-ventrally LIPAREA (Fig. 1) directed. Genus Miculenchus and some species of Filenchus Al1 studies on the sensillae in the Tylenchida head belong to this group. have proved that within the Tylenchida the same basic pattern(De Coninck, 1942)as in al1 is IV. A fourth group has an offset oral disc, the head present : six internal labial, six external labial, four is dorso-ventrally flattened; the amphidial aperture is cephalic and two amphids. In most Tylenchida, how- very long and mostly sinuous, it starts immediately or ever, several of these sensillae have no opening to the close to theoral disc and continues longitudinally on the outside but the nerve endings are found close to or narrow lateral side of the head. withinthe head cuticle (De Grisse, 1977). Wecan Genera belonging to this group : Lelenchus, Ecphyu- assume that the prototylench had the four circlesof dophoroides, Epicharinema. sensillae al1 showing at the outside and with the am- V. A fifthgroup has also an offsetoral disc, the phidial apertures as the most external ones : a situation amphidial slits start immediatelyat theoral disc, they are presentin most free-living nematodes. None of the laterally directed but areonly found on the front endof tylenchs shows that pattern, in al1 tylenchs that have the head, the amphidial apertures are surrounded by a been studied the six external labial sensillae end within plate that bears the four cephalic papillae(if visible), that the head cuticle or close to it. The tylenchs closest to the plate is constricted dorso-ventrally to a varying degree, originalpattern are found within theTylenchidae : the constriction is extreme in Cephalenchus so that four Macrotrophurus, Psilenchus, Atetyylenchus, and separate lobes are formed, inEutylenchus there is a cleft relatives with large slit-like apertures on thelateral side and in Tylodorus there is only a shallow constriction. of the head. Generabelonging to this group : Cephalenchus, The remainingTylenchidae show adifferent ar- Tylodorus, Eutylenchus (with largecephalic setae), rangement and by the SEM pictures some grouping Campbellenchus.Dolichodorus species of which SEM can be proposed (Fig. 1). pictures are available show a similar head. Gracilancea 1. A first groupshows a laterally elongated, undivided differs slightly fromthe foregoing ones because the front plate that carries al1 the sensillae; the amphidial amphidial apertures are rounded. aperturesare entirely within theplate and the four VI. A sixthgroup, quite distinct, is that of the cephalic papillae (if visible) are more towards the out- Boleodorinae. Labialplate undivided, more or less side. In a few species the six internal labial papillae quadrangular;four prominent cephalic papillae (apparently without pores) can be observed in a sym- dome-shaped, outside of anterior surface, usually sur- metrical arrangement around the central mouth openingrounded by adepression. Amphidial apertures start (= a very small dorsoventral slit). Mid-dorsaland between or outside the four cephalic papillae and are mid-ventral the plate is more or less constricted. simple oblique slits or have an invertedV-shape. There Such an oral plate (here identical to labial plate) is is a perioral depression bearing six papillae. [Genera only found in theTylenchidae, in no other tylenchdoes included : Boleodurus, Neopsilenchus, Basiria, Psilenchus the frontplate contain the amphidial aperturesif theand and Basirienchus; also included : Atetylenchus and front plateis elongated, then the elongationis dorsoven- Neothada (amphidial apertures smaller)].

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Aglenchussp.Aglenchus sp. d Coslenchuspolonicus Antarctenchlts hooperi d

/

Tylenchus Malenchus Filenchus Filenchus F. facultativus davainei williamsi normanjonesi vulgaris

II

Miculenchus Filenchus salvus chilensis Letenchus Epicharinema leptosoma keralense Ecphyadophoroides indicus

Eutylen chus Cephalenchus C. hexalineatusCampbellenchus Tylodorus fuequensis brevicaudafus filicauda acuminatus V

Boleodorus Basiria NeopsilenchusBasirienchus Neothada thylactus paragracilis costatus d magnidens geraerti \

Dolichodorus u pulvinus Macrotrophurus Gracilancea EcphyadophoraEcphyadophoraarbusticola graciloides quadralata caelata

Fig. 1. Comparison of en-face views of several Tylenchidae; drawings are made from published SEM-photographs, someof them are schematised so scales have not been added. Al1 head arranged the same way, where known, the ventral side is downwards, amphidial apertures are dashed.

146 Revice Nématol. IO (2) : 143-161 (1987) Reappraisal of Tylenchina. 3. Tylenchidae

VII. Standingapart are Macrotrophurus witha C. The V-relationship ring-like disc and large T-shaped amphidial apertures InGeraert (1979) it was concludedthat in some (Sher & Bell,1975; De Grisse, 1977) and Ecphyado- long-tailed species the vulva position is better correlated phora with very small pore-like amphidial apertures. with the distance from head to anus (V') than with the On the anterior end of several males belonging to body length 0. various genera (Atetylenchus,, Filenchus, The study of V' in several genera with many species Aglenchus, Coslenchus) two papillae on the ventral side revealed that this V' is rather constant within a genus are extremely welldeveloped. These are probably two of and different among the genera. the four cephalic ones. Tylenchus has a V' of 76.8 -1- 3.1; T. butteus Thorne & Malek, 1968 and T. sandneri Wasilewska, 1965 have much higher values of 82 and 85 %;they were MORPHOMETRICRELATIONS V' considered by Andrassy (1979) as Tylenchus species but because of their short conical stylet part they are trans- A. The oesophagus length-body length relationship ferred to Filenchus. Geraertand Crappé (1981) pointed outthat the Malenchus has a V'of 79.9 f 1.7; M. piauhyensis Tylenchidae can be characterized by a proper oesoph- Monteiro, 1974 considered by Brzeski (1982) as a syn- agus/body relationship (not ratio b), significantly dif- onym of Ottolenchus facultativus (Szczygiel, 1964) ferent from the other families of the Tylenchida; it was Brzeski, 1982, wasthe only one in Andrassy's (1981) list also shownthat Antarctenchus,Gracilancea, Atetylen- with an exceptionally high VI-value of 83.5 %. chus, Macrotrophurus, Sauertylenchus and Tylodorus are Aglenchus and Coslenchus have a V of 80.6 5 1.4. within the prolongation of the Tylenchidae line (from Cephalenchus hasamuch more posterior V'of the SEMview Sauertylenchus is considered to belong to 83.6 f 2.0; in Geraert (1968) a V' of approximately 84 was calculated. the " belonolaimids "). A closer examination of the genera usually considered So it is obvious that the afore mentioned genera can as Tylenchidae revealed that most genera have indeed be characterized by their V' value; unfortunately not a very similar relationship with one exception : Malen- every genus has a typical relation : the species with a chus. Malenchus has a slightly longer oesophagus than bodylength above 0.5 mm havegenerally a more species of other genera with a similar body length; for anterior V' (78.2 -t 2.1) while the shorter species have species with a body lengthbelow 0.5 mm theb-value can a more posterior V' (82.5 f 2.8). even be expressed as a ratio, for Malenchus this ratio is 4.82 5 0.47 andfor Aglenchus and Coslenchus it is 5.60 -t 0.32. FEMALEREPRODUCTIVE SYSTEM (Fig. 2)

B. The stylet In al1 females where the arrangement of the several cells of the female reproductive system could be studied Geraert and Crappé (1981) found that the Tylenchi- we found the " crustaformeria " cells of the uterus in dae and Anguinidae form a distinct group within the four rows, the so-called quadricolumella of Wu (1958, Tylenchidawhen the relationship body length/stylet 1967). The very thin Ecphyadophorinae could have a length was considered : they have the shortest stylets. different arrangement but because of this thinness we Study of the relationship between thetwo stylet parts could not make a definite statement. (for which the symbolm is used) revealed that al1 species Observations by Geraert (1981) mentioned that in of a certain, well established genus have a similar re- Filenchus the uterus is tricolumellar, but these obser- lationship and that a definite grouping of genera can vations have not been confirmed. be done : The arrangement of uterus and spermathecacells can - genera with the conical part much smaller than the be used to group some genera : cylindncal part (conical part about 1/3 of total stylet 1) in Eutylenchus, Cephalenchus, Camnpbellenchusand, length : m = 30-40 "O) : the genera of the Boleodorinae, Tylodorus we find along post-vulval and pre-vulval Lelenchus, Ecphyadophora, Ecphyadophoroides, Malen- uterine sac, a distinct transition zone of several cells,the chus, Irantylenchus, Filenchus, Miculenchus. crustaformeria-part of the uterus with aboutfive or six - genera with the conical partabout equalto the cells in each of the fourrows, another transition zone of cylindncal part i.e. m = 40-55 YO : closer to 40 % in several cells, a non-offset elongated spermatheca, ovi- Atylenchus, Aglenchus, Coslenchus, and closer to 50 % in duct, ovary. Tylenchus, Gracilancea, Cephalenchus, Eutylenchus, Epi- A similar arrangement is found in the didelphic Psi- channema. lenchus. - genera with conical part larger than cylindncalpart 2) in most other genera we find a generally smaller i.e. m more than 55 O/O : Macrotropl2urus, Tylodorus. structure with three or four cells in each of the four

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10 prn 1

E

Fig. 2. Genital system of some Tylenchida. A-C : Comparison of the female system of Eutylenchus fueguensis (A), Tylodorus acuminatus (B) and Cephalenchus brevicaudatus (C); D-E : Atylenchus decalineutus .- D : male with large cloaca1 protuberances; E :female with longitudinal vulval flap, large pre-vulval uterine sac, empty, offset spermatheca.In the drawings of the female genital system the columned cells of the uterus have been stippled.

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~~~ ~ uterus rows and with a spermatheca usually offset, DIAGNOSIS sometimes in line. Vulva and vagina also give useful characteristics : Tylenchidae. Head not or slightly offset, amphidial - a thick vaginal Wall, a vulva with epiptygmataand openings variously shaped (with twomajor types, a longitudinal and a transverse slit), usually no distinct lateral flaps dikes ” when smaller; see Andrassy, 1981) is found in Aglenchus, Coslenchus and Malenchus; oral disc. Stylet small to very small, usually with knobs. - a longitudinal flap covering the vulva is found in Female genital tract short with only three or four cells Pleurotylenchus and in Atylenchus; in each of the fourrows ofthe crustaformeria partof the - a vulva outside the mid-ventral line is found in uterus, spermatheca offset or not. Tai1 elongated, top many Cephalenchus species. variously formed. Usually the vulva is a simple transverse slit, and the vagina a thin walled structure. TYPE GENUS : Tylenchus Bastian, 1865. = Aerotylenchus Fotedar & Handoo, 1979 Subfamilies and genera in Tylenchidae (Areotylenchusin Fortuner, 1984) Type species : Many of the about 30 genera can be easily grouped T. davainei Bastian, 1865 into a few subfamilies; as usual others are moredifficult Other species : to place. This is particularly true for thetype subfamily T. arcuatus Siddiqi, 1963 Tylenchinae : the genera left in this subfamily do not T. capitatus Andrassy, 1979 have any particular character(s) that unite them except T. elegans de Man, 1876 that they do not have any particular character(s) at all. T. hayati Khan, 1985 The type-genus of the Ecphyadophorinae is Ecphya- T. kashmirensis Mahajan, 1973 dophora, a genuswith a quiteunique end-on view, T. kirjanovae Andrassy, 1954 Ecphyadophoroides has a different head; Lelenchus has T. maius Andrassy, 1979 been considered to belong in this subfamily because its T. neoandrassyi nom. nov. head isvery similar tothat of Ecphyadophoroides. = T. andrassyi Fotedar & Kaul, 1985 Lelenchus shares,however, many characteristics with nec T. andrassyi Szczygiel, 1969 Filenchus from the Tylenchinae. T. neodavainei Wu, 1969 The Tylodorinae, Atylenchinae and Boleodorinae are T. rex Andrassy, 1979 distinct groupings based on many characteristics. T. ritae Siddiqi, 1963 Genera difficult to include are : Gracilancea (perhaps T. rohtangus Khan, 1985 belonging tothe Atylenchinae), Macrotrophurus and T. safroni (Fotedar & Handoo, 1979) Siddiqi, 1986 Luella. T. sherianus Andrassy, 1981 T. stylolineatus Wu, 1969 T. subdavainei Mukhina, 1981 The subfamily Tylenchinae Orley, 1880 T. tortus Andrassy, 1979 = Dactylotylenchinae Wu, 1969 = Duosulciinae Siddiqi, 1979 OTHER GENERA This subfamily contains the Tylenchidae that are Filenchus Andrassy, 1954 most similar to the type genus Tylenchus. The genus = Dactylotylenchus Wu, 1968 Filenchus is not always easy to distinguish from Tylen- = Discotylenchus Siddiqi, 1980 chus and several species considered to be Tylenchus by = Duosulcius Siddiqi, 1979 Andrassy(1979) belong to Filenchus (cf. Raski & = Lalnbertia Brzeski, 1977 Geraert, 1987); moreover not al1 species considered to = Ottolenchus Husain & Khan, 1965 be Filenchus are well known. In that recent review we = Zanenchus Siddiqi, 1979 synonymize several generawith Filenchus, amongst Type species : them Ottolenchus. Ottolenchus and Malenchus have not F. vulgaris (Brzeski, 1963) Lownsbery & Lowns- been easy to differentiate and several transfers have been bery, 1983 made between them. Other generathat can beincluded Species list in Raski and Geraert (1987) are Miculenchus, Irantylenchus,Polenchus and Allotylen- chus. Al1 these genera show, however,characteristics that Miculenchus Andrassy, 1951 separate them from the rest, moreover Our knowledge Type species : about some of these genera is rather restricted. Miculenchus salvus Andrassy, 1959 So, we will not add a detailed diagnosis (mentioning Other species : the exceptions) but only a short one. M. elegans Raski & Geraert, 1985

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Irantylenchus IZheiri, 1972 slender bodies with long, almost filamentous tails are Type species : indicative of closer relations with the latter. It is also I. clavidorus Kheiri, 1972 preferable to place alinking group with the more derived Other species : to emphasize the direction of change. I. vicinus (Szczygiel, 1970) Brzeski & Sauer, 1983 Ecphyadophora is inconsistent with the elongate head Both species are probably synonyms. and long, slit-like amphids of the other three genera by Malenchus Andrassy, 1968 virtue of its small, oval amphids located close to the anterior endof its symmetrically-shaped cephalic region = Neomalenchus Siddiqi, 1979 annulated almost to oral plate or aperture. However, Type species : considering al1 the other similarities linking these four M. machadoi (Andrassy, 1963) Andrassy, 1968 genera, it seems best to consider Ecphyadophoroides, Species list in Geraert and Raski (1986b) Epicharinema and Lelenchus as most closely related to Allotylenchus Andrassy, 1984 Ecphyadophora in this subfamily. Type and only species : A subfamily Ecphyadophorinae and family Ecphya- A. excretorius Andrassy, 1984 dophoridae were proposed by Skarbilovich (1959). Allen Polenchus Andrassy, 1980 and Sher(1 967) accepted the subfamily taxonbut placed Type and only species : it in the Neotylenchidae. Jairajpuri and Siddiqi (1969) P. politus Andrassy, 1980 restored the family Ecphyadophoridaewithin a new This genus is probably synonymoüs with Tylenchus. superfamilyNeotylenchoidea which was acceptedby Andrassy (1976). Raski et al. (1980) added the genus Cucullitylenchus Huang & Raski, 1986 Epicharinema to the subfamily Ecphyadophorinae but Type and only species : transferred it to theTylenchidae. To these is addod the C. arnazonensis Huang & Raski, 1986 genus Lelenchus as discussed earlier. Maqbool and Shahina(1985) proposed anew generic taxon Karachinema as a monotypic genus based on a Thesubfamily Ecphyadophorinae Skarbilovich, new species, K. elongatum. That species was described 1959 with a prominent stylet, well-developed median bulb = Epicharinematinae Maqbool & Shahina, 1985 with valvular apparatus, a long slender isthmus and a = Ecphyadophoroidinae Siddiqi, 1986 glandular region symmetrically arranged in a bulb-like structure set off from intestine, not overlapping. They This subfamily was reviewed by Raski et al. (1980) considered this species related to Epicharinema by virtue and again by Raski, Koshy and Sosamma(1982). It was of its valvate metacorpus and tylenchoid oesophagus. ofmajor significance to this taxonthat Raski and The report of Raski and Geraert (1986) indicates the Geraert (1986~)established with certainty thetrue vague nature of the oesophagus of Ecphyadophora and nature of the oesophagealmorphology. Until now Ecphyadophoroides is attributed to fEative artifact and speciesof Ecphyadophora and Ecphyadophoroides have indeed both genera have a tylenchoid oesophagus with been characterized with an irregularly tubular oesoph- variously valvated metacorpus. K. elongatum is judged agus without a median bulb; glands in a lobe overlap- most closely related to Ecphyadophora teres Raski, ping the intestine and oesophago-intestinal junction in- Koshy & Sosamma, 1982 and differs by its shorter stylet distinct orobscure. Despite this variancefrom typical (8 pm vs 9-12 pm for E. teres) and smaller size Tylenchidae, the Ecphyadophorinae appearedrelated to (L = 0.68 - 0.71 mm vs 0.83 - 1.00 mm for E. teres). members of that family and were transferred to it (Raski, Karuchinerna is proposed as a new synonym of Ecphya- Koshy & Sosamma, 1982). The œsophageal structure dophora. K. elongatum belongs to Ecphyadophora and gives further indication of closerelationship of Lelenchus transferred to that genus as Ecphyadophora elongatum spp. to Ecphyadophora, Ecphyadophoroides and Epicha- (Maqbool & Shahina, 1985) comb. n. (= Karuchinema rinema. The long, extremely slender body shape (high elongatum Maqbool & Shahina, 1985). " a " values), configuration of cephalic region as high, Epicharinematinae is judged a synonym of Ecphya- smooth, dorsoventrally narrowed with long amphidial dophorinae as syn. n. apertures, somewhat sinuousin shape, and exceptionally long tail - al1 link Lelenchus with Ecphyadophoroides DIAGNOSIS and Epicharinema and in partwith Ecphyadophora. The character mostly relating Lelenchus to Tylenchinae is the Tylenchidae.Moderate-sized; very slender Y a " typical slender, elongate, low caudal alae quite distinct valuesrange from 37-176) cephalic regionwith fine from the flap-like alae of the Ecphyadophorinae. Con- annuli present up to labial plate which bears ' small, sidering Lelenchus as a linkbetween Tylenchinae and ovate amphidial apertures (Ecphyadophora, Mitranema) Ecphyadophorinae we judge the morphology of the or cephalic region high, ' dorso-ventrally narrowed, cephalic region including amphidial structure and the smooth,amphidial apertures long dits (Ecphyadopho-

150 Revue Nématol. 10 (2) : 143-161 (1987) Reappraisal of Tylenchina. 3. Tylenchidae roides, Epickarinema, Lelenchus). Males with flap-like The subfamily Tylodorinae Paramonov, 1967 caudal alae (except Lelenchus); tail narrows markedly = Campbellenchinae Wouts, 1978 after cloaca1 opening; tail long, filiform. = Eutylenchinae Siddiqi, 1986 syn. n.

TYPE GENUS : Ecphyadophora de Man, 1921 The study of the female reproductive systemand the = Karaclzinema Maqbool & Shahina, 1985 anterior head end with SEM showed that the genera Type species : Eutylenchus Cobb, 1913 (Fig. 3), Tylodorus Meagher, E. tenuissima de Man, 1921 1963 (Fig.4) and Cephalenchus Goodey, 1962(syn. Other species : Inzphalenchus) show many charcateristics in common. E. caelata Raski & Geraert, 1986 Canzpbellenchus Wouts, 1977 can also be included. E. elongata (Maqbool & Shahina, 1985) comb. n. Eutylenchus and Atylenchus Cobb, 1913 were con- = I

Revue Neinatol. .IO (2) : 143-161 (1987) 151 E. Geraert & D. S. Raki

Fig. 3. Eutylenchusfueguensis. Female : A : vulva (Bar = 3 Pm); B : body vulva to anus (Bar = 10 p).

part thatis elongated. Stylet always with roundedor oval TYPE GENUS : Tylodorus Meagher, 1963 knobs. The median bulb is welldeveloped andanteriorly Type and only species : situated (except for Tylodorus), oesophagealglands elongated, symmetrically arranged, onlyexceptionally T. acuminatus Meagher, 1963 overlapping. The first intestinal cells are not always well distinguished to form an oesophago-intestinal valve (a central valve is apparently absent). OTHER GENERA Female reproductive system : vulva a transverse slit Campbellenchus Wouts, 1977 with slight modifications in each genus; vagina simple; Type species : only the anterior side of the tract is fully developed, C. poae Wouts, 1977 posteriorly a long post-vulval uterine sac is found; the Other species : uterus is subdivided in a few cellsforming thetransition C. jïlicauda Wouts, 1977 zone withthe uterinesac, acrustaformeria part withfive This genus is discussed in a separate paper (Geraert or six cells in each of the four rows, another transition & Raski, 1986~). zone of several cells, and is followed by a non-offset elongated spermatheca, oviductand ovary (in Campbel- Cephalenchus Goodey, 1962 lenchus the ovary shows a double curvature). = Imphalenchus Dhanachand & Jairajpuri, 1980 Males are known in al1 the genera and almost al1 the Type species : species, there is no sexual dimorphism; spicules are only C. hexalineatus (Geraert, 1962) Geraert& Goodey, slightly curved, caudal alae are always well developed. 1964 Tails elongated with terminus acureto finely rounded. This genus is discussed in a separate paper (Raski and Phasmids on tail in Tylodorus, ad- or postvulval in the Geraert, 1986b). remaining genera. Eutylenchus Cobb, 1913 BIAGNOSIS Type species : E. setifends (Cobb, 1893) Cobb, 1913 Tylodorinae can be characterized by their head end Other species : with an oral disc, slit-like amphid apertures longitudi- E. aficanus Sher, Corbett & Colbran, 1966 nally oriented in a lobed or slightly divided labial disc or = E. orientalis Husain & Khan, 1968 ring; stylet long to very long, the female reproductive E. excretorius Ebsary & Eveleigh, 1981 systemwith more than twentycrustaformeria cells E. heguensis Valenzuela & Raski, 1985 in the uterus and anelongated spermatheca in line. E. vitiensis Orton Williams, 1979

152 Revue Neinatol. 10 (2) :143-161 (1987) Reappraisal of Tylenchina. 3. Tylenchidae

Fig. 4. Tylodorus acuwzinatus.Female :A : head, lateral view; C : lateral field and excretory pore;E : vulva. Male : B :head, lateral view; D : lateral field ending at caudal ala; F : cloacal opening. (Bar on A, B = 2 Pm; C, D = 10 Pm; E, F = 5 Pm). (Arrows on A and B indicate cephalic sensillae).

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Macrotrophurus Loof, 1958 except in Atylenchus (setae) and in themale of Aglenchus Type and only species : sp. where only two of them are well developed. Lateral M. arbusticola Loof, 1958 field with two, three, four or six lines; cuticle provided This genus andspecies are tentatively included in this with longitudinal ridges in Atylenchus, Pleurotylenchus subfamily. and Coslenchus species. Cuticle distinctly annulated with wide annuli. Stylet medium sized, moderately developed with The subfamily Atylenchinae Skarbilovich, 1959 rounded knobs : from 7 pm in Aglenchzu to 22 pm in Antarctenchus; the anterior part is slightly smaller than = Antarctenchinae Spaull, 1972 syn. n. to about equal to the posterior part. = Pleurotylenchinae Andrassy, 1976 Median bulbwell developed, in themiddle or slightly = Aglenchinae Siddiqui & Khan, 1983 syn. n. anterior to the middle of the oesophagus, the posterior Skarbilovich (1959) was the fmt to propose a subfam- bulb short rounded or elongate. ily and family for Atylenchus Cobb, 1913 and Eutylen- Female reproductive systern : vulva with inner lips chus Cobb, 1913. Sher, Corbettand Colbran (1966) (epiptygmata) and lateral flaps covering the transverse revised and accepted the family taxon but rejected the vulvalslit slightly or partly (Aglenchus, Coslenchus, subfamily. Golden (1971) raised Atylenchidae to super- Antarctenchus) or vulva covered by flap-like anterior lip family rank Atylenchoidea for these two genera in the directed backwards (Atylenchus, Pleurotylenchus). Vagi- single subfamily. nal Wall usually distinctly thickened at thevulval side in Tylenchus sachsi Hirschmann,1952 was described Aglenchus, Coslenchus and Antarctenchus. Antarctenchus from specimens collected in Germany. Szczygiel(l969) is didelphic, the remaining generahave onlythe anterior redescribed T. sachsifrom specimens collected in Poland genital tract developed : in Pleurotylenchus and Atylen- and proposed a separate genus, Pleurotylenchus for that chus the post-vulval uterine sac is well developed, in species. Andrassy (1976)proposed a subfamilyPleuroty- Aglenchus and Coslenchus this sac is rudimentary or lenchinae for this monotypic genus and transfered the absent. Uterus short(cells probably always in fourrows taxon to Atylenchidae. Raski et al. (1980) accepted the with three or four cells in a row), spermatheca usually relationship of Atylenchus, Eutylenchus and Pleurotylen- offset. chus but placed them al1 in one subfamilyAtylenchinae Malespresent in al1 genera, but most Coslenchzu in the family Tylenchidae. species appear to be parthenogenetic. Bursawell devel- Eutylenchus differs in many characteristics from the oped in Antarctenchus, Aglenchus and Coslenchzu, very other two genera and shares many characteristics with poorly developed in Atylenchus and Pleurotylenchus. In the Tylodorinae and is dealt with under that heading. Atylenchus, Antarctenchus and Coslenchus polonicus the Atylenchus decalineatusCobb, 1913, the single species cloacal opening is provided withlarge hypoptygmata; in of this genus, shows some interesting characteristics in Aglenchus and remaining Coslenchus species a cloacal female andmale reproductive systemthat have not been tube is present. reported before : the vulva is backwardly covered by a Tai1 long, attenuated, tail tip setose, acute or finely longitudinal flap and the cloacal opening in the male is rounded. Phasmidsad-vulval or post-vulval in Aglenchus bordered by large papilla-likestructures (Figs 5,2). The and Coslenchus, on the tail in Antarctenchus. same longitudinal flap is described for Pleurotylenchus, DIAGNOSIS thesame cloacal ornamentsare foundin Coslenchus polonicus and Antarctenchus hoopen Spaull, 1972.As Tylenchidae.Undivided laterally elongated front explained before, Atylenchus, Aglenchus, Coslenchusand plate of the head with large, roundish amphidial aper- Antarctenchus have also a very similar en-faceview; tures; stylet withabout equal parts, vulvacovered unfortunately it has not been possible to obtain SEM by lateral or longitudinal flaps; males oÎtenwith pictures of P. sachsi. large hypoptygmata.

DESCRIPTION TYPE GENUS : Atylenchus Cobb, 1913 Body smallto moderate size (from 0.33 mm in Coslen- Type and only species : chus to 1.29 mm in Antarctenchus). Head slightly or not Atylenchus decalineatus Cobb, 1913 offset, interna1 sclerotization Îainr. End on view as seen OTHERGENERA by SEM consists of one large, laterally elongated plate more or less constricted dorso-ventrally containing the Agletzchus Andrassy, 1954 large, more or less rounded amphidial apertures (with Type species : lateral offshoot in themale of Aglenchus sp.); around the A. agncola (de Man, 1921) Andrassy, 1954. dorso-ventral mouth slit six papillae are sometimes The genus is discussed in a separate paper (Geraert distinct; the fourcephalic papiIIae are usually indistinct & Raski, 1987).

154 Revue Nérnatol. 10 (2) : 143-161 (1987) Reappraisal of Tyenchina. 3. Tylenchidae

Fig. 5. Atylenchus decalineutus. Female : A : En-face; B : head end, lateral view; C : excretoxy pore (arrow indicates deirid (?)); D : lateral field at deirid (3); E : vulva; F : anus. (Bar on A, B, C = 2 prn; D, E = 5 pn; F = 10 Pm).

Amtarctenchus Spaull, 1972 This genus is discussed in a separate paper (Geraert Type and only species : & Raski, 1987). A. hoopen' Spaull, 1972 Pleurotyletzchus Szczygiel, 1969 Coslenchus Siddiqi, 1978 Type and only species : = Cosaglenchus Siddiqui & JChan, 1983 P. sachsi (Hirschmann, 1952) Szczygiel, 1969 = Paktylenchus Maqbool, 1983 This species shows no differences with Atylenchus Type species : decalineatus except for theabsence of the long cephalic C. costatus (de Man, 1921) Siddiqi, 1978. setae.

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Gracilancea Siddiqi, 1976 The study of a paratype slide of L. amiri revealed that Type and only species : i) the stylet cone measured4 Pm; i.e. about 1/3 (and not G. graciloides (Micoletzky, 1925) Siddiqi, 1976. 1/2) of the stylet length (12.5 Pm); ii) the opening of the In theoriginal description Gracilancea was considered dorsal oesophageal gland is at about 5 pm posterior to to belong to the Tylodorinae,Tylodoridae (Siddiqi, the stylet knobs (and not less than 2 Pm); iii)the lateral 1976); in Brzeski and Sauer’s (1984) redescription the field is indistinct : the original description gives four similarities with Cephalenchus (on one hand) andAglen- lines, the accompanying Fig. 1 E five lines; Siddiqi chus (on the other hand) are stressed. The perioral disc (1986) transfered the species to Merlinius suggesting being well set off makes it difficult to include Graci- there are six lines. In Our study of A. abulbosus the lateral lancea inthe Atylenchinae butthe structure of the field is shown to be slightlyvariable and weaklyex- amphidial apertures within the labial plate, of stylet, pressed (cf. Fig. 6 E, F). vulva and vagina shows some relationship with Tylo- Although the end-on view is not available we prefer dorinae and is tentatively included here. to consider L. amiri as a representative of Atetylenchus, differing from thedescribed species bythe small, flange- like stylet knobs (absent in the other two species). The genus Neobasiria Javed, 1982 was synonymized The subfamily Boleodorinae Khan, 1964 with Basiria by Hashim (1985) and with Boleodorus by Siddiqi (1986); we follow Hashim (1985) as we consider = Psilenchinae Paramonov, 1967 a weakly developed median bulb (vs a normally devel- = Basiriinae Decker, 1972 oped bulb) not a genus character. Geraert and Raski = Leipotylenchinae Sher, 1973 syn. n. (1986a) proposed a new genus Basirienchus and under- lined the similarity of Neothada with theremaining The family Boleodoridae has beentreated in Brzeski Boleodorinae. and Sauer (1984), including the Boleodorinae with Bo- The genus Duotylenchus Saha & Khan, 1982 probably ZeodorusThorne, 1941, Basiria Siddiqi, 1959 and Neopsi- belongs to this group (Siddiqi, 1986); its single species ZenchusThorne &Malek, 1968 and thePsilenchinae with could be a Basiria species. Psilenchus deMan, 1921. and Antarctenchus Spaull, 1972. Antarctenchus isnow treated under a different DIAGNOSIS(cf. Brzeski & Sauer, 1983) subfamily (Atylenchinae); Psilenchus differs from the Tylenchidae. Small to medium size nematodes. Oral remaining gene’a by a paired female genital system a opening surrounded by six papillae, a second circle of character that we consider has not sufficient merit to four papillae on theedges of anterior surface. Amphidial sustain a separate subfamily and so we synonymize the apertures in form of open V or oblique, on lateral side Psilenchinae with the Boleodorinae. ofhead, usually partially coveredby a flap. Stylet A genus that we have also to consider is Atetylenchus delicate, anterior partabout half the posterior part, Khan, 1973 (= LeipotyZenchus Sher, 1974). This genus knobs small, often flange-like, stylet sometimes without was based on a single species, originally described as knobs. Medianbulb with or withoutthickenings of Tetylenchus abulbosus Thorne, 1949. lumen walls. Oesophageal glands symmetrically arran- Sher (1974) differentiated it from Psilenchus by the ged, offset from intestine. Female reproductive system absence of slit-like amphid apertures on the lip region paired or unpaired; usually with few cells in uterusand and thelow flat lip region. He could also haveadded the spermatheca except for Psilenchus; spermatheca offset difference in the position of the median bulb : in A. or in line. Bursa small, adanal. Tai1 elongated, often abulbosus the median bulb is slightly anterior to the rounded at the end, seldom spicate. middle while in the usual Psilenchus species it is dis- tinctly behind the middle. TYPEGENUS : Boleodorus Thorne, 1941 Restudy of the material used by Sher (1974) revealed Type species : that large slit-like amphid apertures are presenttogether B. thylactus Thorne, 1941 with six papillae around theoral opening (Fig. 6). In the Other species : male two of the four cephalic papillae are very distinct. B. abnomus Khan & Basir, 1964 As for the median bulb position the recently described B. acuruus Jairajpuri, 1982 Psilenchus graminus Bajaj, Kaushit & Bhatti, 1982 has B. acutus Thorne & Malek, 1968 also an anterior position; Siddiqi (1986), after studying B. brevistylus Khera, 1970 a paratype, transfeked it to Atetylenchus. B. citri Edward & Rai, 1978 Leipotylenchzu amiri Maqbool & Shahina, 1984 was B. clavicaudatus Thorne, 1941 described unaware of the synonymy of the genus with B. cynodonti Fotedar & Mahajan, 1974 Atetylenchus and unaware of the description of P. gra- B. filiformis Husain & Khan, 1977 minus :from thedescriptions both species can hardly be B. flexuosus Eroshenko, 1982 distinguished. B. hyderi Husain & Khan, 1965

156 Revue Nérnatol. 10 (2) : 143-161 (1987) Reappraisal of Tylenchina. 3. Tylenchidae - - . __ ."

Fig. 6. Atetylenchus abulbosus. Female : A : En-face (by S. A. Sher & A. Bell. Utah specimen); C. En-face (Oregon specimen); E. vulva (Oregon specimen). Male:B :En-face (by S. A. Sher & A. Bell. Utah specimen);D :En-face (Oregon specimen);F :lateral field (Oregon specimen). (Bar on A, B = 4 Fm; C, D = 2 pm; E, F = 5 Fm).

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B. impur Khan & Basir, 1964 Genus incertae sedis :Luella Massey, 1974 B. innuptus (Andrassy, 1961) Siddiqi, 1963 B. mirus Khan, 1964 This taxon was proposed byMassey (1974) as a B. neosimilis Geraert, 1971 monotypic new genus for a new species, Luella lucu- B. pakistanensis Siddiqi, 1963 lenta, found inassociation with Ips pini in red pine. The B. rajïqi Husain & Khan, 1965 description was based on a single female and male and B. similis Khan & Basir, 1963 the taon judged to be a member of Nothotylenchinae. B. spirulis Egunjobi, 1968 The revision of Neotylenchidae (Fortuner & Raski, B. 'volutus Lima & Siddiqi, 1963 1987) has found mostmembers of Nothotylenchinae to B. zaini Maqbool, 1982 belong to theAnguinidae. Luella does not entirely fit the definition of the Anguinidae because of the very short OTHER GENERA post uterine sac(about 46 O/O of VBD as measured from Massey's drawing - Fig. III, C), and the small sperms Basiria Siddiqi, 1959 without surrounding cytoplasm. = Clavilenchus Jairajpuri, 1966 Other characteristicsgiven by Massey include : i)cuti- = Basiroides Thorne & Malek, 1968 cle finely striated,without lateral incisures; id stylet = Neobasiria Javed, 1982 without knobs; iii) oesophageal glands slightly overlap- Type species ping intestine; iv) ovarysingle, reflexed; v) terminus B. graminophila Siddiqi, 1954 filiform; vi) spiculesand gubernaculum tylenchoid; Other species : cf. Hashim (1985) vii) caudal alae adanal. The median bulb is illustrated Basirienchus Geraert & Raski, 1986 as spindle-shaped but no suggestion of musculature or Type species valvular thickenings. B. uncinatus Geraert & Raski, 1986 None of the above are. diagnostic for a particular Other species : cf. Geraert & Raski (1986~) genus, most can be found in various unrelated genera. Nothing is known about the anterior morphology as to Duotylenchus Saha & Khan, 1982 amphids, labial plate, the fourcephalic papillae, oral disc Type and only species or the six labial papillae. D. bilineatus Saha & Khan, 1982 The description byMassey limited by the above Probably junior synonym of Basiria. uncertainties and lack of knowledge lead to the conclu- Neopsilenchus Thorne & Malek, 1968 sion Luella is most probably related to Tylenchidae but Type species : cannot beassigned with confidence to any of the subfa- N. magnidens (Thorne, 1949) Thorne & Malek, milies as presently defined. Therefore it is proposed 1968 Luella be designated genus incertae sedis in the family Other species : cf. Siddiqi (1986) Tylenchidae. Neothada Khan, 1973 Type species N. tatra (Thorne & Malek, 1968) Khan, 1973 Key to the genera (sometimes to species) Other species of Tylenchidae N. cancelZata (Thorne, 1941) Khan, 1973 N. geraerti (Andrassy, 1982), Siddiqi, 1986 1 - Femalesdidelphic ...... 2 - Femalesmonodelphic ...... 5 Psilenchus de Man, 1921 2 Tail short,subcylindrical, rounded; stylet very Type species - long (90-1 10 Pm) ...... Macrotrophurusarbusticola P. hilarulus de Man, 1921 Loof, 1958 Other species : cf. Kheiri (1970) - Tail elongated, attenuated; stylet small (less than Atetylenchus Khan, 1973 20 Pm) ...... 3 = Leipotylenchus Sher, 1971 3 - Cephalic framework moderately sclerotized; vulva Type species providedwith lateral vulval membranes; male A. abulbosus (Thorne, 1949) Khan, 1973 cloaca provided with large hypoptygmata ...... Other species ...... Antarctenchus hooperi A. amin' (Maqbool & Shahina, 1984) n. comb. Spaull, 1972 A. graminus (Bajaj, Kaushik & Bhatti, 1982) Sid- - Cephalic framework not sclerotized; vulva without diqi, 1986 lateralvulval membranes; male ythout hypo- This genus is tentatively included under the Boleo- ptygmata ...... 4 dorinae Lecause it shares many characteristics with Psi- 4 - Head high with distinct amphidial slit on the lenchus; the en-face view is, however, more Filenchus- lateral side; median bulb usually behind middleof like. oesophagus ...... Psilenchus

158 Rèvue Nétnatol. 10 (2) : 143-161 (1987) Reappraisal of Tylenchina. 3. Tylenchidae ...

-Head low with indistinct amphidial slit; median 18 - Stylet without knobs, anterior part with wide lu- bulb anterior to middle of oesophagus .... Atetylenchus men ...... Neopsilenchus 5 - Stylet = 76-104 pm ...... Tylodorus acuminatus - Stylet with or without knobs, anterior part conical Meagher, 1964 withvery fine lumen ...... Basiria - Stylet = 38-52 ...... Epicharinema keralense 19 - Headwith disc-like structure ...... 20 Raski et al., 1980 - Headwith smooth contour ...... 22 - Stylet = 22-34pm ...... 6 20 - Head with small disc at the front end ...... 21 - Stylet less than 22km ...... 7 - Head with large dome shaped structure ...... 6 - Cuticle with longitudinal ridges...... Carnpbellenchus ...... Cuczlllitylenchus amazonensis - Cuticlewithout ridges ...... Gracilancea Huang & Raski, 1986 7 - Headprovided with setae ...... 8 21 - Very slender (a = 62-67); caudal alae posteriorly - Headwithout setae ...... 9 concave ...... Mitranema mitrum 8 - Vulva covered by longitudinal flap, male without Siddiqi, 1986 caudalalae; cloaca provided with large hypo- - Not so,slender; caudal alae rounded ...... Filenchus ptygmata ...... Atylenchusdecalineatus (some Filenchus species, before considered. as a Cobb, 1913 separate genus Discotylenchus) - Vulva with lateral vulval flaps; male with caudal 22 - Veryslender (a = 60-180);bursa lobed ...... 23 alae;cloaca raised ...... Eutylenchus - Relative body width variable, from thick to slender; 9 - Cuticleprovided with longitudinal ridges ...... 10 caudalalae, if present, rounded (five genera) ...... 24 - Cuticle without longitudinal ridges ...... 14 23 - Head quadrangular; amphidial pores on front; 10 - Conusabout 1/3 ofstylet length ...... Il bodyconstricted at vulva ...... Ecphyadophora - Conusslightly less to about 1/2 ofstylet length .. 13 - Head flattened; long sinuous amphidial aperture; bodynot constricted at vulva ...... Ecphyadophoroides 11 - The longitudinal ridges mask the transverse body annulation; no distinct lateral field (the midlateral 24 - Cuticledeeply incised ...... 25 ridge appears slightly larger than the others) ...... - Cuticlenot deeply incised ...... 26 ...... Basirienchuselenae 25 - Head quadrangular in front; body annulations Geraert & Raski, 1986 zigzagin surface view, thisdistinct annulation - The longitudinal ridges dividethe transverse annu- continues on the head; lateral field a small ridge; lationinto blocks, lateral field with 4 lines ...... 12 malewithout caudal alae ...... Miculenchus 12 - Head with 2-3 annuli; stylet without knobs ...... - Head flattened; lateral field a small ridge with 4, ...... Neothada 6 or many lines (the many lines visible only using - Head with 6-7 annuli; stylet with delicate knobs SEM); maleswith caudal alae ...... Malenchus ...... Basirienchuscostatus 26 - Very slender species; annulation not so distinct; Geraert & Raski, 1986 head very flattened with long sinuous amphidial aperture ...... Lelenchus 13 - Vulvacovered by longitudinal flap; stylet = 17-19pm ...... Pleurotylenchussachsi - Relatively thicker species; annulation usually dis- (Hirschmann, 1952) tinct; head usually quadrangular; amphidial aper- - Vulvawith lateral vulval flaps; stylet less than ture whenlong, not sinuous ...... 27 15pm ...... Coslenchus 27 - Head high with longitudinal amphidial apertureon lateralside; clavate stylet knobs with ventrally 14 Conusabout 1/3 ofstylet length 15 - ...... situatedopening of oesophageal lumen; dorsal - Conus slightly less to about 1/2 of stylet length .. 28 oesophageal gland opening one half to one stylet 15 - Headhigh with distinct amphidial slit on the lengthposterior to knobs ...... Irantylenchus lateralside (monodelphic Boleodorinae) ...... 16 - Headquadrangular; annulation distinct; stylet - Head variously shaped, amphidial slit longitudi- withrounded knobs; lateral field with 2, 3 or nallyorientated or only present.on the front ...... 19 4 lines; dorsal gland opening close to knobs ... Filenchus 16 -Body ventrallycurved, sometimes in aspiral; 28 - Vulvaprovided with lateral flaps ...... 29 female reproductive system with offset sperma- - Vulvawithout flaps ...... 30 theca filled with refractive speim and ovary with 29 - Lateral field 2 lines; vagina thin; post-vulval sac oocytes in multiplerows ...... Boleodorus short ...... Allotylenchus -Body more or less straight; sperm not refractive - Lateral field with 3 lines (central ridge can also and oocytesnot inmultiple rows ...... 17 appearas 2 lines close to each other); vagina 17 - Tail hooked or bent near terminus which is pointed thickened;post-vulval sac short ...... Aglenclzus ...... Basirienchusuncinatus - Lateral field with 4 or usually 6 lines; vagina not - Geraert & Raski,1986 thickened; post-vulval sac well developed ...... - Tailmore or less straight ...... 18 ...... Cephalenchus .

Revue Nérnatol. 10 (2) :143-141 (1987) 159 E. Geraert di D. .y. Raski - -

30 - Lateral field and body annulation inconspicuous; GERAERT,E. (1981). The femalereproductive system in caudalalae very small ...... Polenchus systernatics. Annls Soc.r.zoo1. Belg., 110 : 73-86. - Lateral field and body annulation distinct; caudal GERAERT, &E CRAPPE,D. (1981). Morphometric comparison alaedistinct ...... Tylenchus of theTylenchida species (Nematoda). Meded.Fac. LandbWet. Rijksuniv. Gent, 46 : 713-720. GERAERT, E.& RASKI, D. J. (1986a). Three new species of REFERENCES Basirienchus from southern Chile compared withCampbel- ALLEN, M. W. & SHER,S. A. (1967). Taxonornic problems lenchus,Neothada and Basiria.Nematologica, 31 (1985) : concerning the phytoparasitic nematodes.A. Rev. Phytopha- 266-288. thol., 5 : 247-264. GERAERT,E. & RASKI, D.J. (1986b).Unusual Malenchus ANDRASSY,1. (1976). Evolzction as a basis for the systematization species (Nemata : Tylenchidae). Nematologica, 31 : 27-55. of nematodes. London-San Francisco-Melbourne, Pitman GERAERT,E. & RASKI, D. J. 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Accepté pour publicationle 22 août 1986.

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