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1987

A Reappraisal of Tylenchina (Nemata). 4. The family Nicoll, 1935 (1926)

Renaud Fortuner California Department of Food and Agriculture

Armand R. Maggenti University of California - Davis

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Fortuner, Renaud and Maggenti, Armand R., "A Reappraisal of Tylenchina (Nemata). 4. The family Anguinidae Nicoll, 1935 (1926)" (1987). Faculty Publications from the Harold W. Manter Laboratory of Parasitology. 108. https://digitalcommons.unl.edu/parasitologyfacpubs/108

This Article is brought to you for free and open access by the Parasitology, Harold W. Manter Laboratory of at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Faculty Publications from the Harold W. Manter Laboratory of Parasitology by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. A reappraisal of Tylenchina (Nemata). 4. The family Anguinidae Nicoll, 1935 (1926) (l)

Renaud FORTUNER*and Armand R. MAGGENTI California Department of Food and Agriculture, Analysis and Identification (Nematology), 1220 N Street, Sacramento, Ca 95814; and University of Califonlia, Division of Nematology, Davis, Ca 95616, USA

SUMMARK The familyAnguinidae isredefined. The familiesand subfamilies Nothotylenchidaehnae, Sychnotylenchidae/inae, Halenchidaelinae, Pseudhalenchinae, Ditylenchidaehnae, Cynipanguininae, Neoditylenchinae, Nothanguininae and Thadinae are rejected. The genera , Halenchus, , Pseudhalenchus, Sychnotylenchus, Thada, Subanguina, Cynipanguina, and Pterotylenchus are recognized as valid in Anguinidae.Chitinotylenchus is a genus inquirenduln or incertae sedisin Anguinidae. The following new synonyms are proposed:Nothotylenchus, Boleodoroidesand Om’na synonyms of Ditylenchus; Nothanguina synonym of Anguina; Afrinaand Mesoanguina synonyms of Subanguina; and Neoditylenchus synonym of Sychnotylenchus. Hypotheses on the evolution within Anguinidae are presented. A tabular key is includedto help identify the valid genera.

RESUME

Réévaluation des Tylenchina (Nemata). 4. La famille des Anguinidae Nicoll, 1935 (1926). La famille des Anguinidae est redéfinie. Les familles et sous-familles Nothotylenchidae/inae, Sychnotylenchidae/inae, Halen- chidae/inae, Pseudhalenchinae, Ditylenchidaelinae, Cynipanguininae, Neoditylenchinae, Nothanguininae et Thadinae sont rejetées. Les genresAnguina, Halenchus, Ditylenchus, Pseudhalenchus, Sychnotylenchus, Thada, Subanguina, Cynipanguinaet Pterotylenchus sont acceptés comme genres validesa l’intérieur d’Anguinidae. Chitinotylenchus est un genus inquirendunz ou incertae sedis, dans lesAnguinidae. Les synonymes suivants sont proposes : Nothotylenchus,Boleodoroides et Om’na synonymesde Ditylenchus, Nothanguina synonyme d’dnguina, Afn‘na et Mesoanguina synonymes deSubanguina et Neoditylenchus synonyme deSychnotylen- chus. Des hypotheses sur l’évolution probable des formes a l’intérieur des Anguinidae sont proposées, ainsi qu’une clef tabulaire pour aider à l’identification des genres valides de cette famille.

The name Anguinidae was first proposed by Nicoll proposed by Paramonov (1962) at subfamily level with (1935)**, in replacement of Anguillulinidae Baylis & the genera Anguina, Paranguina,and Nothanguina. The Daubney, 1926. same author proposed thefamily Sychnotylenchidaefor When Baylis and Daubney(1926), then Nicoll(l935), some insect-associate (Paramonov, 1967). proposed respectively Anguillulinidae and Anguinidae, Wu (1967a, b) studiedthe relationships between they included in these families genera that are now in ,Ditylenchus, and Anguina. She treated al1 other families in (, , three genera as members of Tylenchinae, but she laid Tylenchulus, etc.), or in other orders ( in the groundwork that eventually permitted theseparation Aphelenchida; Tylopharynx in ). The of Tylenchus from the other two genera. modern conceptof Anguinidae was first recognized, and Siddiqi (1971) placed Ditylenchus in Anguinidae. Golden (1971) separated the genera in Anguinidae into the subfamilies Ditylenchinae and Anguininae in Tylen- ** According to the International Code of Zoological No- chidae. This arrangement is not generally accepted menclature, Article40, a, the correct authority for this family (Brzeski, 1981). Siddiqi (1980) treated Anguinidae as a is : Anguinidae Nicoll, 1935 (1926), see Fortuner (1984). superfamily. Brzeski (1981) attemptedto clarify the

(1) This article is part of a study on the classification of Tylenchina by the present authors and E. Geraert (Rijksuniversiteit, Gent), M. Luc (ORSTOM, Paris), and D.J. Raski (Universityof California, Davis). * Associate in the Division of Nenzatology, university of California, Davis, CA 95614, USA.

Revue Nématpl. Revue (1987)10 (2) :163-176 163 R. Fortuner & A. R. Maggenti status of the genera in Anguinidae. Siddiqi (1986) placed Subanguina Paramonov, 1967 Anguinoidea in Hexatylina. Maggenti et al. (1987) Cynipanguina Maggenti, Hart & Paxman, 1974 proposed to accept the family Anguinidae under Tylen- Pterotylenchus Siddiqi & Lenne, 1983 choidea in the suborder Tylenchina. Genus incertae sedis or inquirendum in Anguinidae : The validity and status of the genera and families of Chitinotylenchus Micoletzky, 1922 anguinids are here discussed accordingto the principles of Luc et al. (1987) and the general classification of COMMENTS Tylenchina of Maggenti et al. (1987). Minimal list of species are given for some genera. Description of Anguinidae Additional information onspecies nomenclature can be Vermiformnematodes; mature femalessometimes found in Fortuner (1987). enlarged, but never globose or kidney-shaped. Lip re- gion low, anteriorly flattened, not or slightly offset, not annulated orwith faint annuli. First lip annulus not The family Anguinidae Nicoll, 1935 (1926) divided into sectors; amphid apertures small, elliptical, directed towards the oral opening.Lateral field with = Anguillulinidae Baylis & Daubney, 1926 either four or six and more lines. Deirids and phasmids = Nothotylenchidae Thorne, 1941 generally absent. Tai1 long, slender, often withlast third = Sychnotylenchidae Paramonov, 1967 ventrally bent; sometimes shorter and rounded. = Ditylenchidae Golden, 1971 Labialframework delicate. Stylet thinand short, = Halenchidae Jairajpuri & Siddiqi, 1969 (n. syn.) about as long as basallip annulus diameter; stylet knobs Synonym subfamilies : present, small, rounded. Stylet muscles parallel to stylet Anguillulininae Baylis & Daubney, 1926 axis. Dorsal esophageal gland opens just below stylet Nothotylenchinae Thorne, 1941 (n. syn.) base. Procorpus thin or wide, separated or not from the Sychnotylenchinae Paramonov, 1967 (n. syn.) median bulb by a constriction. Median bulb (metacor- Halenchinae Jairajpuri & Siddiqi, 1969 (n. syn.) pus) fusiformto rounded, rarely absent, with or without Pseudhalenchinae Siddiqi, 1971 (n. syn.) ’ valve. Isthmus thin to wide. Esophageal glands short, Ditylenchinae Golden, 1971 pyriform,not overlapping the intestine, or longer, Neoditylenchinae Kakuliya & Devdariani, 1975 (n. stopping short of the intestine, or overlapping it for a syn.) short or a long distance. Esophago-intestinal junction Cynipanguininae Fotedar & Handoo, 1978 (cardia) with hyaline cells large, of same diameter as Nothanguininae Fotedar& Handoo, 1978 (n. syn.) intestine. Thadinae Siddiqi, 1986 (n. syn.) One anterior genital branch; posterior branch reduced to a post-uterine sac (PUS) or absent. Ovary straight or DIAGNOSIS with flexures, sometimes very long and reaching to the with low, flattened anterior end; with esophageal region; oocytes in either oneor two rows, or small, delicatestylet and labialframework; female geni- in many rows. Oviduct with two rows of fivecells. ta1 systemwith long sixteen-celled tubularsperma- Spermatheca long, with sixteen cells, tubular, always in theca, in line with genital tract; sperm cells with large line withgenital tract, shorter in Pseudhalenchus. Col- amount of cytoplasm (except Pseudhalenchus). umned uterus variable, with four rows of four or more Anguinidae differs from al1 families in Tylenchoidea cells, or with manysmall cells irregularly arranged. except by the aspect of its anterior extre- Sphincter present at the junction uterus-vagina. Vagina mity with low flattened end, and small, delicate stylet oblique or perpendicular to body axis; no vulval flaps and labial framework. It differs from Tylenchidaeby the (except in one genus) or epiptygma. characteristics of the genital system (Tylenchidae has Males similar to females, no secondary sexual dimor- spermatheca short, twelve-celled, rounded and some- phismexcept in species with enlarged adult females times off-set, andsperm cells with little cytoplasm) where males remain thin. Sperm cells with large amount and by the oesophageal glands, often overlapping the of cytoplasm (except Pseudhalenchus). Caudal alae small beginning of the intestine. adanal (leptoderan), ending atless than 1/3 of taillength, or longer, enveloping up to 213 of tail length, or com- pletely enveloping tail (peloderan). LISTOF GENERA IN ANGUINIDAE Biology. Mycetophagous, insect associates, or para- Anguina Scopoli, 1777 sites of above ground parts of higher plants. Halenchus Cobb, 1933 Ditylenchus Filip’ev, 1936 Discussion on synonymy of Anguinidae Sychnotylenchus Rühm, 1956 In the present article, the genera in Anguinidae are Pseydhalenchus Tarjan, 1958 not placedinto separate subfamilies. The subfamily Thada Thorne, 194 1 name Anguininae will not be used.

164 Revue Nérnatol. 10 (2) : 163-176 (1987)

, Reappraisal of Tylenehina. 4. Anguinidae -

Nothotylenchus, the type genusof the family Nothoty- Halenchus fucicola are present in Ditylenchus (0.drepa- lenchidae, is proposed below as a junior synonymof nocercus). Halenchidae was characterized by Siddiqi Ditylenchus, a genus in Anguinidae. For the same rea- (1986)by sclerotization of excretorycanal, " prehen- sons as those given for thesynonymization at the generic sile '' tail, and marine habitat. Sclerotization can vary. level, Nothotylenchidae is hereconsidered a junior H. dumzonicus has excretory duct not quite so heavily synonym of Anguinidae, as already proposed by Brzeski cuticularized as in thetype species (H.fucicola). Hooked (1981). tails exist, for example, in Ditylenchus drepanocercus. The family Ditylenchidae was differentiated from Differences in habitat cannot be accepted for family Anguinidae by Golden (1971) because of smaller size characterization (this would lead to an excessive multi- (but size is not a reliable generic or family character), plication of families). Because of their variability and slimmer body (there is a great variability in this feature), because they are irrelevant at family level, the criteria ovary without flexure (in fact long ovaries with flexures used by Siddiqi cannot be used and the family Halen- are observed in D. dipsaci, D. angustus, and D. destruc- chidae is here rejected. tor), and oocytes in oneltwo rows'(but the genusSuban- The subfamilyPseudhalenchinae was proposed by guina combines this characteristic with otherfeatures of Siddiqi (1971) after the transfer of Pseudhalenchus in the genital system closer to those in Anguina). Plant- Tylenchinae because this genus is markedly different parasite species of Ditylenchus causestunting and from al1 other generain thelatter subfamily. Pseudhalen- swellings to plants, but nogalls. Their cryptobiotic stage chus is hereconsidered in Anguinidae.Within this is the 4th juvenile stage. Anguinu spp. cause galls or family, the only remarkable character of the genus are gall-like distortions to plants, and their cryptobiotic the smallsperm cellswith little cytoplasm. A single stage is the 2nd juvenilestage. These differences in character is not considered to be diagnostic at family biology do not support the validity of Ditylenchinae level and Pseudhalenchinae is here rejected. becausenot al1 species of the genera in Anguinidae The subfamilyCynipanguininae was proposed by follow to either one of the two above-defined schemes. Fotedar and Handoo (1978) because of the stem-like Some Ditylenchus spp. for example are mycetophagous. process at the posterior end of the esophageal glands. The cryptobiotic stage is the. adult stage in Cynipan- The other morphological characters of Cynipanguina guina. The biologyof Sychnotylenchus ispeculiar as are close to Anguina to such an extent that Brzeski discussed below. Using biology as a criterion for sub- (1981) synonymized the genera Cynipanguina and An- families within Anguinidae wouldlead to accept almost guina. These genera are here considered valid separate as many such taxa as there exist valid genera in the taxa, but the morphological differences between them family. We prefer to follow Hooper (1978),Siddiqi are too slight to warrant the creation of a separate (1980), and Ryss and Krall' (1981) and maintain Dity- subfamily for Cynipanguina. lenchus and Anguina in the same family. Nothanguina is proposed belowas asynonym of The biologyof Sychnotylenchus the type genus of Anguina. This makesthe subfamily Nothanguininae Sychnotylenchidae is different from that of the other Fotedar & Handoo, 1978 a synonym of Anguinidae. anguinids in that the species in this genus are always Thadinae was proposed by Siddiqi (1986) to accom- found associated with bark beetles. However, they are modate Thada and Neothada in Tylenchidae. Thada is not insect parasites but feed on the fungi that grow in here placed in Anguinidae very close to Ditylenchus as the frass of the beetles. Their general morphology is shown below. This closenessmakes the creation of similar to that of the Anguinidae except forthe shorter Thadinaeunnecessaq and this subfamilyis here re- male and female tails. This is not enough to justify the jected. placement of these forms in a family or a subfamily of their own. The family Sychnotylenchidae has already been treated as a junior synonym of Anguinidae by The genera in Anguinidae Siddiqi (1971), Golden (1971), and Andrassy (1976). Pseudhalenchus Tarjan, 1958 The subfamily Sychnotylenchinaeis here considered as a synonym of Anguinidae. Because Neoditylenchus is DIAGNOSIS synonymized below with Sychnotylenchus, the subfamily Neoditylenchinae isalso considered a synonymof Anguinidae. Amphids as longitudinal slits. Median Anguinidae. bulb with valve; isthmus not separated from glandular Halenchinae was proposed in Nothotylenchidae by bulb by aconstriction; glandular bulb long, overlapping Jairajpuri and Siddiqi (1969), and elevated to family rank intestine. Ovary short; oocytes in oneltwo rows; sperma- by Siddiqi (1986). It was differentiated from Nothoty- theca short; columned uterus with four rows of four lenchinae by esophageal glands overlapping intestine, cells; post-uterine sac (PUS) present. Testes without and tail tip strongly hooked. Overlapping versus abut- flexures; caudal alae leptoderan, short, adanal. Sperm tingglands is not accepted as afamily or generic cells small, with little cytoplasm. Mature females not criterion (Fortuner, 1982); hooked tails similar to that of swollen. Feeding habits unknown.

Revue Nématol. 10 (2) :163-1 76 (1987) 165 R. Fortuner & A. R. Maggenti

TYPE SPECIES = Diptenchus Khan, Chawla, & Seshadri, 1969 P. minutus Tarjan, 1958 = Safianema Siddiqi, 1980 = Om’na Brzeski, 1981 (n. syn.) OTHER SPECIES P. hylobii Massey, 1967 DIAGNOSIS Anguinidae. Medianbulb with or without valve; DISCUSSION isthmus not separated from glandular bulba constric- by Pseudhalenchus was originally proposed by Tarjan tion; glandular bulbshort or long,when long may (1958) for two new species,P. minutus and P. anchilispo- overlap the intestine for a short or long distance. Ovary somus, and placed in Tylenchinae by its author. It was short or long, sometimesreaching esophageal region transferred to anew subfamily,Telotylenchinae by andior flexed; oocytes in oneltwo rows; columned uterus Siddiqi (1960). This action was rejected by Loof (1963). with four rowsof four cells; post-uterine sac (PUS) Siddiqi (1971) later created a monotypicsubfamily, present or absent. Testes usually without flexures; Pseudhalenchinae in Anguinidae, for Pseudhalenchus. caudal alae leptoderan, short adanal or long, but never At the same time, Golden (1971) placed this genus in reaching tail end. Mature female notor slightly swollen. Ditylenchinae. Siddiqi (1980) transferred P. anchilispo- Mycetophagous or parasites of higher plants, found in somus and several other species to a new genus, Safia- soi1 or above ground. nema in Anguininae, andplaced what was left of Pseud- halenchus and thefamily Pseudhalenchinae in Tylenchi- dae. Ryss and Krall’ (1981) transferred this subfamily to TYPE SPECIES . Fortuner (1982) accepted the transfer D. dipsaci (Kiihn, 1857) Filip’ev, 1936 of Pseudhalenchus to Tylenchidae, but rejected Pseudha- lenchinae. OTHER SPECIES The unsettledclassification of Pseudhalenchus isa direct consequence of its systematic position, interme- Ditylenchus acris (Thorne, 1941) n. comb. = Nothotylenchus acris Thorne, 1941 diate between Tylenchidae and Anguinidae. Fortuner Ditylenchus acuminatus nom. novum (1982) placed it in Tylenchidae becauseof the structure = Pseudhalenchus acutus Khan & Nanjappa, 1972 of the sperm cells, smalland with small amount of = Ditylenchus acutus (Khan & Nanjappa, 1972) Fortu- cytoplasm, a character of Tylenchidae. However, Raski ner, 1982 nec Nothotylenchus acutzts Khan, 1965 (pers. comm.) pointed out that its longglandular overlap Ditylenchus acutus (Khan, 1965) n. comb. is quite unknown in al1 Tylenchidae (with the exception = Nothotylenchus acutus Khan, 1965 of Epicharinema witha very slight overlap),whereas Ditylenchzcs adasi (Sykes, 1980) n. comb. many species in Anguinidaehave a longglandular = Nothotylenchus adasi Sykes, 1980 overlap. Face views of a population of P. minutus from Ditylenchus affinis(Thorne, 1941) n. comb. Brazilwere observed with SEM byRaski. Amphids = Nothotylenchus affinis Thorne, 1941 appear as small longitudinal slits (i.e., pointing towards Ditylenchus alliphilus nom. novum the oral opening) and are generally similar to face views = Nothotylenchus allii Khan & Siddiqi, 1968 nec Dity- of Anguina and Ditylenchus (as shown in Raski & lenchusallii (Beijerinck,1883) Filip’ev & Schuur- Maggenti, 1983). Face views in Tylenchidae are quite mans-Stekhoven, 1941 different. Only Filenchus has small dits, but they are Ditylenclus anchilisposomzts (Tarjan, 1958) Fortuner, 1982 dorso-ventrally directed (Raski & Geraert, 1987). (Butler, 1913) Filip’ev, 1936 Pseudhalenchus can be seen as a form evolving to- Ditylenchus antricolus (Andrassy, 1961) n. comb. wards thetypical anguinids. It already had some charac- = Nothotylenchus antricolus Andrassy 1961 teristics of this family (face view, glandular overlap), but Ditylenchus attenuatus (Mulvey, 1969) n. comb. it still shows ancestral characters found in Tylenchidae = Nothotylenchus attenzcatzcs Mulvey, 1969 (sperm cells). Because classification should be basedon Ditylenchus atypicus (Khera & Chaturvedi, 1977) n. comb. = Boleodorus atypicus Khera & Chaturvedi, 1977 derived, ratherthan ancestral characters, Pseudhalen- = Nothotylencusatypicus (Khera & Chaturvedi,1977) chus is now reinstalled in Anguinidae. The subfamily Siddiqi, 1986 Pseudhalenchinae has been rejected (see above). Ditylenchzls ausafi Husain & Khan, 1967 Ditylenchus basiri (Khan, 1965) n. comb. Ditylenchus Filip’ev, 1936 = Nothotylenchus basiri Khan, 1965 Ditylenchus bhatnagari (Tikyani & Khera, 1969) n. comb. = Anguillulina (Ditylenchus) Filip’ev, 1936 = Nothotylenchus bhatnagari Tikyani & Khera, 1969 = Nothotylenchus Thorne, 1941 (n. syn.) Ditylenchus brassicae Husain & Khan, 1976 = Boleodoroides Mathur, Khan & Prasad, 1966 (n. Ditylenchus buckleyi (Das, 1960) n. comb. syn.1 = Nothotylenchus buckleyi Das, 1960

166 Revue Nématol. 10 (2) :163-176 (1987) Reappraisal of Tylenchina. 4. Anguinidae

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Ditylenchus callidus (Izatullaeva, 1967) n. cornb. = Nothotylenchusmajor Thorne & Malek,1968 nec = Nothotylenchus callidus Izatullaeva, 1967 Ditylenchus major (Fuchs, 1915) Filip’ev, 1936 Ditylenchus caudatus Thorne & Malek, 1968 Ditylenchus medians (Thorne & Malek, 1968) n. comb. Ditylenchus citri (Varaprasad, Khan & Lal, 1981) n. cornb. = Nothotylenchus nzedians Thorne & Malek, 1968 = Paurodontus citri Varaprasad, Khan & Lal, 1981 Ditylenchus nzedicaginis Wasilewska, 1965 = Nothotylenchus citri (Varaprasad, Khan & Lal, 1981) Ditylenchus melongena Bhatnagar & Kadyan, 1969 Siddiqi, 1986 Ditylenchus microdens Thorne & Malek, 1968 Ditylenchus clarus Thorne & Malek, 1968 Ditylenchus minutus Husain & Khan, 1967 Ditylenchus compactus (Massey, 1974) n. cornb. Ditylenchus mirus Siddiqi, 1963 = Nothotylenchus compactus Massey, 1974 Ditylenchus,?nyceliophagusGoodey, !958 Ditylenchus convallariae Sturhan & Friedman, 1965 Ditylenchus nanus Siddiqi, 1963 Ditylenchus cylindricollis (Thorne, 1941) n. cornb. Ditylenchus nortoni (Elrniligy, 1971) Bello & Geraert, 1972 = Nothotylenchus cylindricollis Thorne, 1941 = Basiroides nortoni Elmiligy, 1971 Ditylenchus cylindricus (Khan & Siddiqi, 1968) n. cornb. = Basiri,a nortoni (Elrniligy, 1971) Fotedar & Mahajan, = Nothotylenchus cylindricus Khan & Siddiqi 1968 1973 Ditylenchus cyperi Husain & Khan, 1967 Ditylenchus obesus Thorne & Malek, 1968 Ditylenchus damnatus (Massey, 1966) Fortuner, 1982 Ditylenchus oyzae (Mathur, Khan& Prasad, 1966) n. cornb. Ditylenchus danubialis (Andrassy, 1960) n. comb. = Boleodoroides oyzae Mathur, Khan & Prasad, 1966 = Nothotylenchus danzdbialis Andrassy, 1960 = Paurodontus oyzae (Mathur et al., 1966) Surnenkova, Ditylenchus deiridus Thorne & Malek, 1968 1975 Thorne, 1945 = Nothotylenchus ozyzae (Mathur et al., 1966) Siddiqi, Ditylenchus dipsacoideus (Andrassy, 1952) Andrassy, 1956 1986 Ditylenchus drepanocercus (Goodey, 1953) Ditylenchus paramonovi (Gagarin, 1974) n. cornb. Ditylenchus dyadis Anderson & Mulvey, 1980 = Nothotylenchus paranzonovi Gagarin, 1974 Ditylenchus elongatus (Husain & Khan, 1974) n. cornb. Ditylenchus parasindis (Massey, 1974) n. cornb. = Nothotylenchus elongatus Husain & Khan, 1974 = Nothotylenchus parasindis Massey, 1974 Ditylenchus emus Khan, Chawla & Prasad, 1969 Ditylenchus petilus (Massey, 1974) n. cornb. Ditylenchus equalis Heyns, 1964 = Nothotylenchus petilus Massey, 1974 Ditylenchus exiguus (Andrassy, 1958) n. cornb. Ditylenchus plzyllobius (Thorne, 1934) Filip’ev, 1936 = Nothotylenchus exiguus Andrassy, 1958 = Anguillulina phyllobia Thorne, 1934 falcariae Pogosyan, 1967 = Om’na phlyllobia (Thorne, 1934) Brzeski, 1981 Ditylenchus ferepolitor (Kazachenko, 1980) n. comb. Ditylenchus saxenai nom. nov. = Nothotylenchus ferepolitor Kazachenko, 1980 = Nothotylenchus indicussaxena, Chhabra& Joshi, 1973 Ditylenclzus filinzus Anderson, 1983 nec Ditylenchus indicus (Sethi & Swarup, 1967) For- Ditylenchus fotedari (Mahajan, 1977) n. cornb. tuner, 1982 = Nothotylenchus fotedari Mahajan, 1977 Ditylenchus sibiricus Gerrnan, 1969 Ditylenchusgaleopsidis Teploukhova,1968 . Ditylenchus silvestris (Kazachenko, 1980) n. cornb. Ditylenchus goldeni (Maqbool, 1982) n. cornb. = Nothotylenchus silvestris Kazachenko, 1980 = Nothotylenchus goldeni Maqbool, 1982 Ditylenchus sintilis (Thorne & Malek, 1968) n. cornb. Ditylenchus hexaglyphus (Khan & Siddiqi, 1968) n. cornb. = Nothotylenchus similis Thorne & Malek, 1968 = Nothotylenchus hexaglyphus Khan & Siddiqi, 1968 Ditylenchus singhi (Das & Shivaswarny, 1980) n. cornb. Ditylenchus indicus (Sethi & Swarup, 1967) Fortuner, 1982 = Notlzotylenchus singhi Das & Shivaswamy, 1980 Ditylenchus innuptus (Andrassy, 1961) n. cornb. Ditylenchus solani Husain & Khan, 1976 = Nothotylenchus innuptus Andrassy, 1961 Ditylenchus sonchophila (Kir’yanova, 1958) Ditylenchusinobseruabilis (Kir’yanova,1938) Kir’yanova, Ditylenchussrinagarensis (Fotedar & Mahajan,1974) n. 195 1 comb. Ditylenchus istatae Sarnibaeva, 1966 = Nothotylenchussrinagarensis Fotedar & Mahajan, Ditylenchus khani Fortuner, 1982 1974 Ditylenchus kheirii nom. novurn. = Nothotylenchus geraerti Kheiri, 1971 nec Ditylenchus Ditylenchus taleolus (Kir’yanova, 1938) Kir’yanova, 1961 geraerti (Pararnonov, 1970) Bello & Geraert in Bello, Ditylenchustausaghyzatus (Kir’yanova,1938) Kir’yanova, 197 1 196 1 Ditylenchus loksai (Andrassy, 1959) n. cornb. Ditylenchus taylori (Husain & Khan, 1974) n. cornb. = Nothotylenchus loksai Andrassy, 1959 = Nothotylenchus taylori Husain & Khan, 1974 Ditylenchus longistylus(Khera & Chaturvedi, 1977) n. cornb. Ditylenchus tenuidens Gritsenko, 1971 = Boleodorus longistylus Khera & Chaturvedi, 1977 Ditylenchus thonzei (Andrassy, 1958) n. cornb. = Notlzotylenchuslongistylus (Khera & Chaturvedi, = Nothotylenchus thonzei Andrassy, 1958 1977) Siddiqi, 1986 Ditylenchus triformis Hirschrnann & Sasser, 1955 Ditylenchus lutonensis (Siddiqi, 1980) Fortuner, 1982 Ditylenclzustruncatus (Eliashvili & Vacheishvili,1980) n. Ditylenchus maleki nom. nov. comb.

Revue Nématol. 10 (2) :163-176 (1987) 167 R. Fortuner di A. R. Maggenti -

= Nothotylenchustruncatus Eliashvili & Vacheishvili, bulb valve, elongation of the esophageal glands thatmay 1980 overlap the intestine, regression of the post-uterine sac. Ditylenchus tuberosus (IZheiri, 1971) n. comb. Such species are easier to differentiate from the tylen- = Nothotylenchus tuberosus IZheiri, 1971 chids, and sometimes have been removed from Ditylen- Ditylenchus tudus (Yokoo, 1968) n. comb. chus to other genera. = Neotylenchus turfilsYokoo, 1968 Safianema and Diptenchus were proposed as junior = Nothotylenchus turfus (Yokoo, 1968) Siddiqi, 1986 synonyms of Ditylenchus by Fortuner (1982). Safianema Ditylenchus uniformis (Truskova & Eroshenko,1977) n. was differentiated from Ditylenchus primarily because of comb. the long esophageal gland overlap over the intestine. = Nothotylenchus unifonnis Truskova & Eroshenko, 1977 This character is variable within the same species and Ditylenchus utschini (Gagarin, 1974) n. comb. even varies during thelife of the same specimen (Fortu- = Nothotylenchus utschini Gagarin, 1974 ner,1982). Diptenchus was differentiated primarily Ditylenchus valveus Thorne & Malek, 1968 because of the absence of a PUS. These two synony- Ditylenchus virtudesae Tobar-Jimenez, 1964 mizations were rejected bySiddiqi (1986) without argu- Ditylenchus varaprasadi nom. nov. ment or justification. = Paurodontus solani Varaprasad, Khan & Lal, 1981 Thorne (1941) described Nothotylenchus in Neotylen- = Nothotylenchussolani (Varaprasad et al., 1981)Sid- chidae becauseof the valveless median esophageal bulb. diqi, 1986 nec D. solani Husain & IZhan, 1976 He noted in thediagnosis that thespecies in Nothotylen- chus mayeasily bemistaken for Ditylenchusdipsaci The name Nothotylenchus strictusproposed in a thesis (Kiihn) Filipjev or D. intemedius (de Man) Filipjev, by Kapoor (1982), is not available. Neotylenchus nitidus especially since they are so frequently associated with Massey, 1969, wastranferred by Siddiqi (1986) to Notho- alfalfa crown where these two species often are found.” tylenchus underthe erroneous authority “ Massey, The description of Nothotylenchus given by Thorne 1974 ”. This decision is not accepted herein view of the (1941) fits Ditylenchus, except forthe absence of valves. columned uterus illustrated by Massey as closer to that This description was left unemendeduntil Brzeski in Subanguina. Boleodorustypicus Husain & Khan, (1981) added a few details (isthmus not separated by a 1967 was also transferred to Nothotylenchus by Siddiqi constriction; columned uterus with four rows of four (1986).Oocytes were said to be arranged in multiple cells; etc.). These features also are present in Ditylen- rows in this species. Both species will have to be inves- chus. The biology of speciesin Nothotylenchus is not well tigated further before their taxonomic position can be known. Most are suspected to be mycetophagous. Some decided. For the moment they are considered incertae are found in the frass of bark beetles. At least one sedis. species, N. acris, has been associated with a disease of strawberry inJapan (Nishizawa & Iyatomi,1955). DISCUSSION Geraert (1976) wrote that Nothotylenchus was probably synonymous with DityZenchus. Sumenkova (1974) and DityZenchus is the genus in Anguinidae thatis closest Brzeski (1981) were also of the opinion that these two to the Tylenchidae, except Pseudhalenchus. Many Dity- may be congeneric. lenchus spp. have an esophageal regionsimilar to that in Tylenchidaewith fusiform median bulband short The only morphological difference remainingbe- pyriform glands. The female genital system is the least tween Ditylenchus and Nothotylenchus is the absence of derived among Anguinidae with oocytes in one or two median bulb valves in the lattergenus. Some species of rows and columned uterus as four rows of four cells. DityZenchus have large, well distinct, valves (D. dipsaci, Many males have short leptoderan caudal alae, as in D. destndctor, etc.). Manyother have smaller valves. Tylenchidae. It may be difficult to differentiate some Africanpopulations of D. myceliophagus haveweak species in Ditylenchus (with short pyriformglands, short valves that often become invisible in fïed specimens ovary, and males with short caudalalae) from members (Fortuner, 1982). There is no well-marked difference for of Tylenchidae. Wu (1967) showedthat the spermatheca this character between the two genera, but we obsenre always long, tubular, and in-line with the genital tract a continuum of specific forms where valves, at first in Ditylenchus was a good character to differentiate this strong and well-marked, regress and eventually disap- genus from tylenchids with spermatheca small, round, pear completely. and oftenoffset from thegenital tract. The size ofsperm Because there is no marked difference in the biology cells visiblein thespermatheca (always large with a large and themorphology of the two genera; becauseNothoty- amount of cytoplasm in Ditylenchus) is another good Zenchus and Ditylenchus differ bya single character; differentiating character from tylenchids ‘(with small because different species in DityZenchus present success- sperm cells with reduced cytoplasm). ive stages in the regression, from typical clearly valvate Other members of thegenus show somederived DityZenchus to typical valveless Nothotylenchus; and characters : reduction and disappearanceof the median because a regressed character is not a Sound basis for

168 Revue Nématol. 10 (2) :163-176 (1987) Reappraisal of Tylenchina. 4. Anguinidae -~ systematic differentiation, Nothotylenchus Thorne, 1941 mada Thorne, 1941 is here proposed as a synonym of Ditylenchus Filip’ev, 1936. Boleodoroides was originally described in Boleodori- DIAGNOSIS nae, close to Boleodorus andother genera now in Anguinidae. Similar to Ditylenchus. Wide (2.5-3 pm) Tylenchidae, and to Nothotylenchus now a synonym of well marked bodyannuli. Stylet knobs as small swellings Ditylenchus in Anguinidae. at the end of the shaft. DG0 at more than 4 pm from Khera (1970) reduced Boleodoroides as a subgenus stylet base. Median bulbwithout valve. Deirids present. under Boleodorus. Geraert (1971) studied paratypes of Spicules cephalated. Boleodoroides oyzae (type of species) and found many differences from the original description. He concluded that Boleodoroides does not resemble Boleodorus at all, TYPEAND ONLY SPECIES but is much closer to Paurodolztus. Jairajpuri (1982) agreed that B. oyzae and B. brevistylus do not belong T. striata Thorne, 1941 to the Boleodorus group, and he preferred to consider Boleodoroides as a genus incertae sedis. COMMENTS There is no esophagealstem in B. oyzae, so this species differs from Paurodontus. Some characters re- Thada was originally differentiated by Thorne (1941) semble Ditylenchus andthe anguinids (head shape, because of its unusually thick, deeply striated cuticle, female gonad, and particularly .the spermatheca). The cap-like valvular apparatus at the esophago-intestinal esophagusresembles that of Nothotylenchus. Siddiqi junction, and ovate cephalation of the spicules. Geraert (1974) found that thecuticle of type material (1986)gave new illustrations of B. o yzae (type of species) and proposed to synonymize Boleodoroides to of the two species described by Thorne (1941) in thenew Nothotylenchus. His action is accepted here, but because genus nada (T. striata, type species, and T. cancellata) Nothotylenchus is here treated as a junior synonym of was no thicker than in other Tylenchida (0.5 to 1.5 pm Ditylenchus,Boleodoroides is proposed as a newsyn- thick compared to about0.7, up to2 pm,in Tylenchida). onym of this latter genus. Geraert (1974) commented that Thada is a “ disturb- ing mixture ” of characteristics of tylenchids and ,of Om’na was differentiated from the rest of the An- Ditylenchus. The anterior end and esophageal region is guinidae ‘< by having overlapping oesophageal lobe ” typical of the tylenchids. The genital system is closer to (Brzeski, 1981). An esophageal overlap exists in some Ditylenchus. Thada has wide, well-marked body annuli, species of Ditylenchus (Fortuner, 1982). The type and sometimes with longitudinal striae (T. cancellata), and a only species in the genus, O. phyllobia is very close to conicaltail with a rounded end (Geraert, 1974). The Ditylenchus, and in factwas long knownas D. phyllobius taxonomic position of. Thada becomes clearer if the (Thorne, 1934) Filip’ev, 1936. It has many characteris- species that have been placed in this genus are con- tics of the genus, including stout female body, general sidered separately. shape of anterior and posterior ends,female genital The genital system of T. cancellata, as illustrated by tract, male caudal alae and tail. The type species is an Geraert (1974) with round spermathecafilled with small above-ground plant parasite and produces leaf galls on spermsresembles genital systems in tylenchids. The the weed Solanum elaeagnifolium. It differs from typical species T. camellata, with slit-like amphids and deep members of Ditylenchus by .the complete absence of a longitudinal and transversal striae as in Coslenchus for valve and median esophageal bulb.It has been discussed example, is a typical member of Tylenchinae. T. can- above that regression of valves was not a good justifi- cellata and T. tatra Thorne & Malek, 1968 have been cation for the differentiation of the genus Nothotylen- transferred to a different genus,Neothada Khan, 1973. chus which was synonymized to Ditylenchus. The ab- Neothada belongs to Tylenchidae(Raski & Geraert, sence of the median bulb itself is a bettercharacter, but 1987). some species in Nothotylenchus now in Ditylenchus also T. striata is now the only species left in Thada. The have no median bulb at al1 (N. antricollus, N. cylindri- genital system of T. striata, as illustrated by Geraert collis, and N. cylindricus). (1974) is identical to the genital system in Ditylenchus, Absence of median bulb occurs in many unrelated with elongate spermathecafilled with large sperms. The species. Regression of this organ in insect associates and amphid apertures inT. striata are certainly smaller than parasites and in plantparasites needs to be investigated in T. cancellata and it is even questionable that theyare and its taxonomic significance evaluated. In the mean- slit-like ”. Face view, preferably studied with SEM,are time, because of its general appearanceand b~ology,D. needed to define the exact shape of this feature. T. phyllobia is transferred back to Ditylenchus phyllobius striata has wide annuli (2.5-3 Pm), but no longitudinal and thè genus Om‘na is proposed as a junior synonym striae. The esophago-intestinal junctionand the tail of Ditylenchus. shape are not very different from the similar structures

Revue Nématol. 10 (2) :163-176 (1987) 169 R. Fortuner & A. R. Maggenti

in Ditylenchus. The spicules are said to be cephalated, OTHERSPECIES but so are the spicules in some Ditylenchus spp. (D. dipsaci, D. destmctor, etc.). H. dumnonicus Coles, 1958 Because of thestructure of its genital system, T. H. mediterraneus (Micoletzky, 1922) Cobb, 1933. striata, type species of the genus nada, belongs to the Anguinidae. The differences between T. striata and the COMMENTS species in Ditylenchus are small : possibly different Halenchus, originally described in Tylenchinae, was amphid apertures, smaller stylet knobs, DG0 farther transferred to Nothotylenchinae by Thorne (1949) be- from the stylet,wider body annuli, slightly different cause of the absence of median bulb valve. Jairajpuri spicule shape. Additional collections of T. stnata need, and Siddiqi (1969) placed it ina separate subfamily, Ha- to be studied to decideif T. stnata, and thegenus nada, lenchinae, in Nothotylenchidae, because of the gland are different from Ditylenchus. Inthe meantime, it overlap and of the hooked tail tip. Sumenkova (1974) seems best to accept 7'hada as a valid genuswithin argued that esophagus structure should form the basis Anguinidae. for classification of Neotylenchoidea. She transferred Halenchus to characterized by isthmus reduced, esophageal glands separated from esophagus Pterotylenchus Siddiqi & Lenne, 1984 proper, and intestine joining directly with corpus. Sid- diqi (1980) considered Halenchus as related to Notho- DIAGNOSIS tylenchinae, in Anguinoidea. The esophagus of Halenchus is different from that of Anguinidae.Median bulb not differentiated, valve neotylenchids (Hexatylus). It has no well-defined me- lacking. Deirids present. Vulva partly covered by large dian bulb (similar in this respect to some Ditylenchus cuticular flaps. Ovary straight with oocytes mostly in spp.), a well defined isthmusencircled by nerve ring,and single file; columned uterus withfour rows of 8-9 cells. intestine seemingly joining esophagus in the glandular Males unknown. area. Glandular overlap is very long, but no different from that of some Ditylenchus spp. TYPE AND ONLY SPECIES Halenchus fits well in the family Anguinidae as al- ready recognized bySiddiqi (1980) :female gonad, male P. cecidogenus Siddiqi & Lenne, 1984. large sperms, small caudal alae, tail of both sexes. It is The type species forms galls on thestems of a tropical close to Ditylenchus and mada by oocytes in one or two Pasture legume, Desmodium ovalifolium. rows, and esophagus with non constrictions. SEM face views are needed to better differentiate these three COMMENTS genera. Pterotylenchus is close to Ditylenchus by its general The marine habitat of Halenchus is unique in An- appearance, its esophagus, similar to some species for- guinidae. In Tylenchina, only some Hirschmanniella merly in Nothotylenchus and Om'na, and its ovary. It spp. () are known marine . H. differs from Ditylenchus mostly in thecolumned uterus mexicana and H. zoostericola were once placed in Halen- similar to Subanguina, and the presence of cuticular chus. This genusdiffers from Hirschmanniella by shape flaps. of labial area, stylet more slender, median bulb notwell defined, monodelphicfemale gonad, large sperms in tubular elongate spermatheca, and tail shape. Halenchus Cobb, 1933 Subanguina Paramonov, 1967 DIAGNOSIS = Heteroanguina Chizhov, 1980 Anguinidae. Labial regionslightly offset and narrower = Afrina Brzeski, 1981 (n. syn.) than body. Corpus without distinct median bulb, val- = Mesoanguina Chizhov & Subbotin, 1985 (n. syn.) veless. Esophagealglands a long overlapping lobe. Excretory duct wide. Oocytes in one/two rows. Caudal DIAGNOSIS alae short, adanal. Tai1 of both sexes with hooked tip. Marine nematodes; formgalls on seaweeds (Ascophyl- Ànguinidae. Median buïb with vaives; isthmus may lzun, Fucus). beseparated fromglandular bulb by a constriction, esophageal glands mayor may not overlap the beginning of the intestine. Oocytes in oneltwo rows; columned TYPE SPECIES uterus with four long rows of cells (about eight/twelve H. fucicola(de Man & Barton in de Man,1892) Cobb, cells per row). Testes usually without flexures; bursa 1933. medium sized to long but not reaching tail end.

170 Revue Nénaatol. 10 (2) :163-1 76 (1987) Reappraisal Tylenchina. 4. Anguinidae - of

Mature females swollen or not swollen. Form galls on genus was characterized by : z) being parasitic on dicoty- root and/or above ground parts of higher plants. ledons only (vs monocotyledons only or both mono- and dicotyledons); iz) having third stage larvae as the infective stage (vs second or fourth stage); iiz) forming TYPE SPECIES unstained galls with well-marked cavity (vs partially or S. radicicola (Greef, 1872) Paramonov, 1968 completely stained galls without well-markedcavity) and; iv) having two morphologicallydistinct generations in the galls (us one generation only). OTHERSPECIES Differences in hostlist should not beused as generic A list of speciesin Subanguina has recently been given criteria. If they were,other genera wouldhave to besplit. by Brzeski (1981). To this list must be added : In Ditylenchus, for example, some species feed onfungi, Subanguina hyparrheniae (Corbett, 1966) comb. others can feed on both fungi and higher plants (D. = Anguina hyparrheniae Corbett, 1966 destructor), while D. dipsaci can survive on only two = Afrina hyparrheniae (Corbett, 1966)Brzeski, fungus species (Maggenti, 1981). 1981 The authors of Mesoanguina use differences in in- Subanguina tumefaciens(Cobb, 1932) comb. fective larval stages to justify the proposa1 of this new = Tylenchus tumefaciens Cobb, 1932 genus. In thesame article (Chizhov & Subbotin, 1985), = Afrina tumefaciens (Cobb, 1932) Brzeski, 1981 they reject Cynipanguina which is the only anguinid genus with adultsas the infective stage (Maggenti, Hart & Paxman, 1974). Cynipanguina is accepted here be- COMMENTS cause of morphological differenciations. The synonymy of Heteroanguina and Subanguina has Gall stains are not necessarily caused by the parasite, been proposedby Brzeski (1981).The morphology of S. but moreoften by the reaction of the plant to the granzinophila given as type species of Heteroanguina is presence of the parasite. Some other factors may also different fromAnguina as indicated by Chizhov (1980), induce gall stains. Coynebacterium, for example, turn but is no different from Subanguina as redefined by galls into a bright yellow. Brzeski (1981). Two generations of Anguillulinamillefolii (= Me- The genus Afrtna was proposed by Brzeski (1981) for soanguina millefolii, type species)were described by A. hyparrheniae (Corbett), type species and A. tumefa- Goodey (1938). They differ in body proportions and in ciens (Cobb). It was said to be intermediate between somemorphological details. The firstgeneration is Anguina and Subanguina because it presents some larger,with spiral habitusand reflexedovary. The characters of either genera (esophagus,ovary, and flexed smaller second generation hasstraighter body and ovary testis as in Anguina, columned uterus as in Suban- mostly outstretched. Chizhov andSubbotin (1985) guina). proposed Mesoanguina with first generationadults smal- Examination of paratypes of A. hyparrheniae revealed ler, straigher, and with ovary straight or reflexed. They that the female genital system is identical to that in also differentiate thearrangement of oocytes inthe Subanguina. In theovary, after a zone of multiplication, ovary, in a single row (first generation) vs two or three the oocytes are arranged in two rows, and even in a single rows (second generation). From the study of Goodey file at the end of the ovary of some specimens, not in (1938), the maturation areas of both generations look multiple rows as in typical Anguina. Characteristics of remarkably similar with oocytes in one row. Mature esophagus and testis may be variable in Anguina and oocytes do seemto crowd each other at the proximal end Subanguina. Some Anguina have no constriction be- of the ovary in the first generation females, but the tween isthmus and glandular bulb (A. agrostis, A. grami- aspect is quite different from that in Anguina sensu nis, A. microlaenae, etc.). The testis of S. radicicola are stricto. This alternance of generations is said byHooper flexed and no different from the testis in Anguina. and Southey (1978) to be characteristic of A. millefolii. Because the structure of the female genital system is It is not clear how Chizhov and Subbotin extended its the best differentiating character between Subanguina description to thirteen otherspecies. and Anguina and because the structure of this system While the introductionof biological considerations in in A. hyparrheniae, type species of Afrina is similar to diagnoses of species should be encouraged, Subanguina, the genus AJi-ina is here proposed as a biology alone doesnot differentiate a genus, when there junior synonym of Subanguina. are no morphological differences. Also the use of bio- Mesoanguina was proposed by Chizhovand Subbotin logical characters should be subject to the same pru- (1985) for M. millefolii (type species) and a dozen other dence as any other character. Generic criteria must be anguinid species (M. amsinckia, M. balsamophila, M. shown to be reasonably constant and their presence be centaureae, M. chartolepidis, M. cousiniae, M. kopetdag- verified in al1 species grouped into a new genus. hica, M. mobilis, M. montana, M. moxae, M. pharangii, Mesoanguina millefoliiis here retransferred to Suban- M. picridis, M. plantaginis and M. varsobica). The new guina because of the arrangementof oocytesin one row.

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Mesoanguina is proposed as a newsynonym of this OTHER SPECIES genus. Al1 species placed in Mesoanguina by Chizhov See list in Brzeski (1981) with the following addition : 1. and Subbotin (1985) are transferred back to Anguina or cecidoplastes (Goodey, 1934) Filip’ev, 1936 Subanguina, according to their taxonomic position in A. = Nothanguina cecidoplastes (Goodey, 1934) Whi- Brzeski (198 1). tehead, 1959 Anguillulina and Anguina had the same typespecies. Chitwood (1935) commented on the status of Anguina Cynipanguina Maggenti, Hart & Paxman, 1974 and recognized itsseniority over Anguillulina. Anguina was accepted in the Officia1 List of Works Approvedas DIAGNOSIS . Available for Zoological Nomenclature, First Instal- ment, 1958, opinion 329, and Anguillulina was con- Anguinidae. Median bulb with valve; isthmus separ- sidered a junior objective synonym of Anguina in theOf- ated from the glandular bulb by a constriction; intestine ficial Index of Rejected and Invalid Generic Names in overlapping the end of the glandular bulb that forms a Zoology, First Instalment, 1958, opinion 341, name 235. stem-like extension.,Oocytesin oneltwo rows; columned Brzeski (1981) proposed to consider Paranguina as a uterus with four long rows of cells (about 14 cells per synonym of Anguina because his examination of speci- row). Testes without flexures, caudal alae leptoderan. mens of the type species of Paranguina revealed that the Mature females swollen.Above ground parasites of additional esophagealgland described by IGr’yanova higher plants. The adults are the cryptobiotic stage was a fixation artifact. Hisconclusions are accepted instead of one of the larval stages as in the other here. Anguinids. Nothanguina was differentiated from Anguina by the lack of a valve in thepoorly developed median bulb and bylack of a gubernaculum.Both characters are not TYPEAND ONLY SPECIES accepted as valid generic characters, and Nothanguina C. danthoniae Maggenti, Hart & Paxman, 1974 is here proposed as a new junior synonym of Anguina. The synonymy of Cynipanguina and Anguina pro- posed by Brzeski (1981) is here rejected because the Sychnotylenchus Riihm, 1956 stem-like esophageal extension of C. danthoniae is not (Name not available in Riihm, 1950; Riihm, 1955) an artifact. It was seen in living specimens, and in both younger and older females as well as in males. Also the = Neoditylenchus Meyl, 1961 (n. syn.) genital system is closer to that of Subanguina. DIAGNOSIS Anguina Scopoli, 1777 Anguinidae. Labial framework six sectored with the six sectors equal or lateral sectors narrower than sub- = Anguillulina (Anguina) Scopoli, 1777 dorsal and sub-ventral ones. Esophagus with procorpus = Anguillulina Gervais & Van Beneden, 1869 generally wide and separated from median bulb by a = Paranguina IGr’yanova, 1955 constriction, rarely thinand without a constriction. = Nothanguina Whitehead, 1959 syn.) (n. Medianbulb with valve,well-defined, fusiformto spherical. Esophageal glands short, pyriform to elon- DIAGNOSIS gate, wide. Position of excretory pore variable from anterior to median bulb to opposite posterior end of Anguinidae. Procorpus generally separated from the glands. Female tail small, cylindroidto broadly rounded median bulb by a constriction. Median bulb with or end. Oocytes in one/two rows. Columned uterus four without valves; isthmus generally separated from the rows of more than four cells (seven to fourteen cells), glandular bulb bya constriction; esophagealglands PUS present. enlarged, generally overlapping intestine. Oocytes in Male with peloderan caudalalae. Spicules long, some- many rows; columned uterus a long multinucleatetube. times longer than tail. Testes usually with flexures; gubernaculum rarely ab- sent; caudal alae long but notreaching tail tip. Mature TYPE SPECIES females swollen. Form galls on above ground parts of higher plants, generally grasses. S. intricati Rühm, 1955

OTHERSPECIES TYPE SPECIES S. abieticolus (Riihm, 1956) n. comb. A. tritici (Steinbuch, 1799) Chitwood 1935 = Ditylenchus abieticolus Riihm, 1956

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S. abietis Riihm, 1955 works are small and documented only in a few species, S. autographi (Riihm, 1956) n. comb. it is not considered that thelip shape can beused for the = D. autographi Riihm, 1956 moment to differentiate genera. S. corniculatus (Massey, 1974) n. comb. = Neoditylenchus corniculatus Massey, 1974 2. Position of the excretoly pore S. dendmctoni (Massey, 1974) n. comb. = N. dendroctoni Massey, 1974 The position of the excretory pore varies in Neodity- S. dendrophilus (Marcinowski, 1909) n. comb. Zenchus from thelevel of the posterior end of the glands = Tylenchus dendrophilus Marcinowski, 1909 to that of the base of the median bulb; it varies in S. erelnus (Riihm, 1956) n. comb. Sychnotylenchus from thebase of the median bulb to the = D. eremus Riihm, 1955 posterior third of the procorpus. The ranges of positions S. gallicus (Steiner, 1935) n. comb. in the two genera overlap slightly; the pore is more = Anguillulina gallica Steiner, 1935 anterior in N. petithi than in S. phloeosini. S. glandarius (Massey, 1974) n. comb. Because the position of the excretory pore varies = N. glandarius Massey, 1974 continuously from species to species with no definite S. glischrus (Riihm, 1956) n. comb. gap between the two nominal genera - with, in fact, a = D. glischrus Riihm, 1956 slight overlap - it is difficult to accept this character as S. major (Fuchs, 1915) n. comb. a generic criterion. = i? major Fuchs, 1915 S. ~zuticiMassey, 1974 3. Esophagus S. ortus (Fuchs, 1938) n. comb. The species traditionnally placed in Neoditylenchus = A. orta Fuchs, 1938 S. ovarius (Massey, 1974) n. comb. (excretory pore in amore posterior location) and in = N. ovarius Massey, 1974 Sychnotylenchus (excretory pore more anterior) exhibit S. panurgus (Riihm, 1956) n. comb. a wide variability in the shapes of the different parts of = D. panurgus Riihm, 1956 the esophagus. S. petithi (Fuchs, 1938) n. comb. Procorpus :In both genera, the procorpus is generally = A. petithi Fuchs, 1938 wide and separated fromthe median bulb bya S. phloeosini Massey, 1969 constriction. It is thinner and lacks a constriction in N. S. pinophilus (Thorne, 1935) n. comb. striatus, N. petithi, and S. ulrni. = A. pinophila Thorne, 1935 S. pityokteinophilus (Riihm, 1956) n. comb. Median buZb :The median bulb varies from a strong = D. pityokteinophilus Riihm, 1956 spherical structure to a weak fusiform swelling. Both S. puniwopus (Massey, 1969) n. comb. shapes occur in either genera : N. ovarius, S. ulmi, etc., = N. puniwopus Massey, -1969 have strong spherical bulbs; N. eremus, S. scolyti, etc., S. rarus (Meyl, 1954) n. comb. have fusiform bulbs. = D. rarus Meyl, 1954 Isthmus :The isthmus is generally thin. It is wider in S. scolyti Massey, 1969 ' N. panurgus and a few other Neoditylenchus. However, S. striatus (Fuchs, 1938) n. comb. many Neoditylenchus have a thinor a very thin isthmus: = Anguillonema striatuln Fuchs, 1938 S. ulnzi Riihm, 1955 N. corniculatus, N. abieticolus, etc. S. yasinskii (Massey, 1969) n. comb. Esophageal glands :The glands are generally long and = N. yasinskii Massey, 1969 enlarged in Neoditylenchus, except in some species like N. striatuswith shorter glands. They are short and squat, pyriform, in Sychnotylenchus, except S. scolyti with COMMENTS longer glands. Neoditylenchus has been differentiated from Sychno- The taxonomicvalue of the shape of esophageal tylenchus by several authors (Rühm, 1956; Meyl, 1961; glands can be questioned in view of the wide range of Goodey, 1963; Massey, 1974; Kakuliya & Devdariani, variation that exists in the related genus Ditylenchus. 1975). The differentiating criteria are discussed below. 4. Relative size of spicules and male tail 1. Relative size of lip sectors In the two genera under consideration, the species- The lip sectors are described only in N. panurgus, S. with the most anterior position of the excretory pore intricati, S. phleosini, S. scolyti. In al1 cases, the illus- have very long spicules, equal to, or longer than thetail, trations show the labial framework, not the lip sectors. while the species withmore posterior excretory pore We are unable tolocate any SEM photograph of the lip have comparatively shorter spicules. sectors. The division between the two groups of species does Because the actual external shape of the lip sectors is not qvite fit the traditional division between Sychnoty- unknown and because the differences in labial frame- le6hÛs and Neoditylenchus. All Sychnotylenchus and the

Revue Nématol. 10 (2) : 163-176 (1987) 173 R. Fortuner & A. R. Maggenti

Neoditylenchus spp. with excretory pore at level or present a derived esophageal region(with elongation of anterior to nerve ring have longer spicules. The rest of glandsthat eventually overlapthe intestine), derived the Neoditylenchus have shorter ones. However, there are genital system (with elongated ovary that may reach the exceptions to this rule : N. pityokteinophilus, N. striatus, esophagus and/orfold upon itself), longer relative length etc. of bursa inrelation to tail (bursa reachingto 2/3 or more In conclusion, some species in the two genera under of tail length). Evolution of each character seems to be discussion have the excretory pore more anterior, eso- independent from the othercharacters. For example, D. phageal glands shortand squat, spicules as long as tail, cyperiwith bursa reaching almostto tail end has esopha- and possibly lateral sectors of the labial framework gus and ovary not derived. narrower than the other sectors. Other species have the These evolutionnary tendanciesin Ditylenchus points excretory pore moreposterior, esophageal glands longer towards the morederived plant parasitic anguinids : and wider, spicules shorter, and al1 sectors oflabial Subanguina, Cynipanguina, and Anguina. Evolution in framework equal. Many species do not fit this dicho- these genera proceeded by furthermodifications of the tomy and present characters of both groups. There is no esophagus (enlarged procorpus and/or isthmus,increa- difference in the biology of these species: they are al1 sed size of glands). In Cyni9aguina the end of the glands associates of bark beetles. are enfolded withinthe anterior end of the intestine and Neoditylenchus is proposed as a new junior synonym becomes acharacteristic " stem " not unlike the stem in of Sychnotylenchus paurodontids. These modifications of the digestive sys- tem maybe related to a betker adaptation to plant parasitism evident in many species of these genera (leaf galls). At the same time, and perhaps correlated to the Chitinotylenchus Micoletzky, 1922 better food supply, modifications of the genital system produce more eggs (rearrangement of the ovary with The nomenclaturalstatus of this genushas been multiple rows of oocytes) that are processed through a recently reviewed by Fortuner (1982). Because many longer columned uterus (twelve or multi-celled). characteristics relevant for generic differentiation are The plant parasitic anguinidsare well adapted to not known for C. paragracilis, type species of the genus, above-ground parasitism on higher plants. However, Chitinotylenchus is considered a genus inquirendum or this ecological nicheproved to be an evolutionnary incertae sedis in Anguininae. dead-end and the main stream of evolution towards phytoparasitism was due to occur within the soi1 eco- system where it probably originated and developed in different nematode groups (belonolaimids, . pratylen- Hypothetical evolution of the anguinids chids, hoplolaimids, and heteroderids). Many species in Ditylenchus, as defined above, seem Some species in the Ditylenchus (0.ausaji, D. emus, to follow a different evolutionaryprocess. In this second . D. equalis, D. microdens, D. nortoni, and D. valveus) are group of species, the median bulb valve regresses and very close to Tylenchidae (esophagusnot evolved, male eventually disappear completely.The evolutionary signi- tail with short bursa). This may indicate acommon ficance (if any) of this regression is not known. origin of the two families Tylenchidae and Anguinidae. A third group of species evolved from hypothetical The basic differences between these two groups is the Ditylenchus-like ancestors when they becameassociated structure of the genital system. In Anguinidae, sperm withbark beetles. Foran unknown reason their tail cells have a greatly enlarged cytoplasm. The evolutio- length, both in male and female specimens, became naryvalue of this modification is not known. The greatly reduced until a different genus, Sychnotylenchus, increase in sperm ce11 size is related with a modification was differentiated with short rounded femaletails, and of the female spermatheca.In Anguinidae, this organ is short male tail, completely enveloped by the bursa. a long tubular structure, much larger than the corres- ponding small rounded pouch of the tylenchids. Ditylenchus is a large complex genus, withspecies that Generic identification show evolutionary tendencies in several directions. A first group of species became specialized parasites To help with the practical identification of the genera ~f zbGve-goiind pzfis of %&leï plailts (D.disaci, E. discussed in hguiridae, a t~?xt!~rky for the idectificz- destructor,D. angustus, and others). These species tion of the valid taxa is proposed in Table 1.

174 Revue Nématol. 10 (2) :163-1 76 (1987) Reappraisal of Tylenchina. 4. Anguinidae

Table 1 Tabular key to the genera in Anguinidae

Procorpus Isthmus Glands OocytesProcorpusGlandsIsthmus CaudalCells of alae Renzarks in Ovay Colunzella

~~~ ~ ~~ ~ ~ ~ Pseudhalenchus No constric. No constric. Overlap 112 rows 4x4 Leptoderan Srnall sperms Ditylenchus No constric. No constric. + 1-overlap 112 rows 4x4 Leptoderan Halenchus No constric. No constric. overlap 112 rows Leptoderan Marine; wide excre- tory duct. Thada No constric. No constric. No overlap 112 rows 4x4 Leptoderan DG0 far from sty- let Pterotylenchus No constric. No constric. Overlap 112 rows 4 X (8-9) Leptoderan Cuticular flaps pre- sent Subanguina + I- + I- + /-overlap 112 rows 4 x (8-12) Leptoderan constric. constric. Cynipanguina No constric. Constric. Stern Several rows 4 X (12-14) Leptoderan Anguina + I- Constric. + 1- overlap Several rows Multi-celled Leptoderan constric. Sychnotylenchus + I- No constric. No overlap 112 rows 4 X (7-14) Peloderan Tai1short cylin- constric. droid

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Accepté pour publication le 20 août 1986.

176 Revue Nématol. 10 (2) : 163-176 (1987)