12
The genera of Anguinidae ( matoda, Tylenchida) MichaI W. BRZESKI Insfyfuf War~yumicfztra, 96-100 Skierniemice, Poland. SUMMARY The family Anguinidae is redefined ; it includes the genera Anguina Scopoli, 1777 ; Nofhanguincz Whitehead, 1959 ; Afrinn g. 11. ; Orrina g. n. ; Subunguina Paramonov, 1968 ; Difylenchzzs Filipjev, 1936 ; Nofhofylenchus Thorne, 1941, and Dipfenchzzs Khan, Chawla & Seshadri, 1969. Parangzzinu Kirjanova, 1955 and Cynipcznguincz Maggenti, Hart & Paxman, 1974 are synonymised wit.h Angzzinn. Les genres de la famille des Anguinidne (Netnafoda : Tylenchida) L’auteur redéfinit la famille des Angzzinidae ; celle-ci comprend les genres Anguina Scopoli, 1777, Noflzangzzinn Whitehead, 1959, Afrina g. nov., Orrina g. nov., Szzbanguina Paramonov, Nofhofylenchus Thorne, 1941, et. Dipfenchus Khan, Chawla 62 Seshadri, 1968, Dif~lenchzzs Filipjev, 1936, 1969. Paranguznn Kirjanova, 1955 et Cynipanguina Maggenti, Hart & Paxman, 1974 sont c.onsidérés comme synonymes mineurs d’Anguinn. Anguinn poophila Kirjanova, 1952 est synonymisé avec A. ngrosfis Steinbuch, 1799. Nofhtrngzzinn cecidoplasfes Goodey, 1934 est. la seule espèce retenue dans le genre Nofhanguina. Nofhangzzina phyllobin Thorne, 1934 devient l’espèce type d’Orrina g. nov. Afrinn g. nov. comprend A. hypnrrheniae (Corbett, 1966) comb. nov. et A. fzzmefczciens (Cobb, 1932) comb. nov. Dix-sept espkces appartenant aux genres dngzzina ou Paranguinu sont. transférées au genre Szzbanguina. A l’int&ieur des Anguinidae la complexité de la structure de la gonade femelle parait être en relation avec le degré d’adaptat,ion au parasitisme. Paramonov (1962) considered t!he Anguininae a subfamily of Tylenchidae, and c.haracterised this subfamily by the large number of oogonia and sexual dimorphism. He included the genera Anguina Scopoli, 1777, Paranguina Kirjanova, 1955, and Nothnnguina Whitehead, 1959. Later on, the genus Subangnina Paramonov, 1968, was added. While recognising close afflnities between Anguina and Dit,ylenchus Filipjev, 1936, Paramonov (1962, 1970) placed the latter in Tylenchinae. Wu (1967) found it difflcult to separate these two genera and redefined both on the basis of t,he length of crustaformeria in relation to the length of spermatheca. Choi and Loof (1973) also discussed the status of Anguina Revue Nématol. 4 (1) : 23-34 (1981) and Ditylenchus when they regarded Subanguina as synonymous with Anguina. In 1974, the new genus Cynipanguina Maggenti, Hart & Paxman, 1974, was proposed. Siddiqi (1971) recognised the subfamilies Anguininae, Pseudhalenchinae and Syc.hnot,y- chinae within t.he Anguinidae. However, Geraert and Kheiri (1970) studied the two known species of PseudhaZenchus Tarjan, 1958, and found that P. nzinutus, t.he type species, is closer to Tylen- chus Bastian, 1865 s. 1. than to Ditylcnchrzs. Golden [1971) proposed the new subfamily Ditylenchinae to ac.comodate Ditylenchus, Pserrd- halenchus and Diptenchus, while in the Angui- ninae he included Anguina, Paranguina and 23

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Page 1: The genera of Anguinidae (Nematoda, Tylenchida)horizon.documentation.ird.fr/exl-doc/pleins_textes/... · Les genres de la famille des Anguinidne (Netnafoda : Tylenchida) L’auteur

The genera of Anguinidae ( matoda, Tylenchida)

MichaI W. BRZESKI

Insfyfuf War~yumicfztra, 96-100 Skierniemice, Poland.

SUMMARY

The family Anguinidae is redefined ; it includes the genera Anguina Scopoli, 1777 ; Nofhanguincz Whitehead, 1959 ; Afrinn g. 11. ; Orrina g. n. ; Subunguina Paramonov, 1968 ; Difylenchzzs Filipjev, 1936 ; Nofhofylenchus Thorne, 1941, and Dipfenchzzs Khan, Chawla & Seshadri, 1969. Parangzzinu Kirjanova, 1955 and Cynipcznguincz Maggenti, Hart & Paxman, 1974 are synonymised wit.h Angzzinn.

Les genres de la famille des Anguinidne (Netnafoda : Tylenchida)

L’auteur redéfinit la famille des Angzzinidae ; celle-ci comprend les genres Anguina Scopoli, 1777, Noflzangzzinn Whitehead, 1959, Afrina g. nov., Orrina g. nov., Szzbanguina Paramonov, Nofhofylenchus Thorne, 1941, et. Dipfenchus Khan, Chawla 62 Seshadri,

1968, Dif~lenchzzs Filipjev, 1936, 1969. Paranguznn Kirjanova, 1955 et

Cynipanguina Maggenti, Hart & Paxman, 1974 sont c.onsidérés comme synonymes mineurs d’Anguinn. Anguinn poophila Kirjanova, 1952 est synonymisé avec A. ngrosfis Steinbuch, 1799. Nofhtrngzzinn cecidoplasfes Goodey, 1934 est. la seule espèce retenue dans le genre Nofhanguina. Nofhangzzina phyllobin Thorne, 1934 devient l’espèce type d’Orrina g. nov. Afrinn g. nov. comprend A. hypnrrheniae (Corbett, 1966) comb. nov. et A. fzzmefczciens (Cobb, 1932) comb. nov. Dix-sept espkces appartenant aux genres dngzzina ou Paranguinu sont. transférées au genre Szzbanguina. A l’int&ieur des Anguinidae la complexité de la structure de la gonade femelle parait être en relation avec le degré d’adaptat,ion au parasitisme.

Paramonov (1962) considered t!he Anguininae a subfamily of Tylenchidae, and c.haracterised this subfamily by the large number of oogonia and sexual dimorphism. He included the genera Anguina Scopoli, 1777, Paranguina Kirjanova, 1955, and Nothnnguina Whitehead, 1959. Later on, the genus Subangnina Paramonov, 1968, was added. While recognising close afflnities between Anguina and Dit,ylenchus Filipjev, 1936, Paramonov (1962, 1970) placed the latter in Tylenchinae. Wu (1967) found it difflcult to separate these two genera and redefined both on the basis of t,he length of crustaformeria in relation to the length of spermatheca. Choi and Loof (1973) also discussed the status of Anguina

Revue Nématol. 4 (1) : 23-34 (1981)

and Ditylenchus when they regarded Subanguina as synonymous with Anguina. In 1974, the new genus Cynipanguina Maggenti, Hart & Paxman, 1974, was proposed.

Siddiqi (1971) recognised the subfamilies Anguininae, Pseudhalenchinae and Syc.hnot,y- chinae within t.he Anguinidae. However, Geraert and Kheiri (1970) studied the two known species of PseudhaZenchus Tarjan, 1958, and found that P. nzinutus, t.he type species, is closer to Tylen- chus Bastian, 1865 s. 1. than to Ditylcnchrzs.

Golden [1971) proposed the new subfamily Ditylenchinae to ac.comodate Ditylenchus, Pserrd- halenchus and Diptenchus, while in the Angui- ninae he included Anguina, Paranguina and

23

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fil. W. Brzeski

Subanguina. This ha s not been rec.ognised by Hooper (1978) who synonymised Anguininae and Ditylenchinae.

Sumenkova (1975) and Geraert (1976) stated that Nofhofyknchus Thorne, 1941 is probably synonymous wit.11 Difylenchus, although they belong to different superfamilies.

In order to resolve the above confusion, extensive studies have been made of these groups and especially of Anguinu s. 1.

Family Anguinidae Nicoll, 1935

GENERAL MORPHOLOCX

The cuticle of members of this family lias delicate t.ransverse annulat,ions and no longi- t,udinal lines were observed. Lateral field of the genera Difylenchus, Nofhofylenchzzs and Dipfen- clous Khan, Chawla 3t Seshadri, 1969, bas four to six incisures. The number of incisures on swollen adults of the remaining genera was impossible t,o Count..

Stylet is thin and small, wit,h the conus usually shorter than t.he shaft. In some Anguina species the conus may be slightly bent. Knobs small. Cephalic skeleton weakly developed. Stylet protract.ors attac.hed to the knobs and t.he basa1 plate of t.he skelet.on.

Oesophageal median bulb with or wit,hout refract,ive thickenings. Terminal bulb may be offset, slightly overlap int.estine on. dorsal side, or form a dist,inct lobe. The swollen adult,s of some genera may bave irregularly shaped terminal bulb. Sometimes secretions of dorsal oesophageal gland may accumulate in isthmus or proc.orpus giving the appearance of a “storage organ” (Thorne, 1961). However, this depends on t.he physiological activity of a spec.imen and cannot be considered as a taxonomie character. Isthmus may be offset from terminal bulb by deep constriction. Cardia absent and ant,erior intestinal c.ells hyaline.

Female reproduct.ive system proclelphic ; post.- ulerine sac. present, rarely absent.. The few cells usually present in posterior branc.h of the repro- ductive system are remnants of an ancestral paired gonad. One female of Angzzinn fritici was seen wit.h a long posterior gonad, but without. a spermatheca. Vulval lips may be

24

prot.ruded. A few tightly packed cells always present, between uterine sac and crustaformerja. Crust,aformeria of the genera least adapted to parasitism is in t.he form of quadricolumella : four rows of cells with four cells per row. The evolution toward parasit.ism is marked by imrease in the number of çells in t.he crusta- formeria ; the greatest development occurs in Anguina and Nofhangzzinn where it is a multi- nuc1eat.e tube. Spermatheca axial. Oogonia may be arranged in tandem wit,h few cells in circum- ference, or with many c.ells in c.ircumference. In t.he last case the rachis is probably present.

Male with paired spicules, (an aberrant male of Su banguina. picridis had three spic,ules). Gubernaculum not protrusible. Bursa usually envelops most of the tail. Testis similar in st.ruct.ure to ovary. Sperm rounded.

DIAGNOSIS : Tylenchoidea. Cuticle delicately annulated. Deirids usually present, phasmids not observed. Amphidial apertures pore-like, labial. Stylet short, weakly developed. Cephalic skele- t.on not or weakly sclerotizecl. Median bulb present. or absent., with or w&hout. refrac.tive t.hickenings. Cardia lacking, anterior intestinal c.ells hyaline. Female gonad prodelphic, sperma- theca axial, wit,h a sphinc.ter between uterine sac and crustaformeria. Tail of bath sexes conoid. Bursa envelops most. of t.he tail. ‘

SYNONYMS : Nothotylenchinae Thorne, 1941, s. n., Dit,ylenchinae Golden, 1971.

TYPE GENUS

Anguina Sc.opoli, 1777

= Purungzzinn Kirjanova, 1955, s. nov. = Cynipanguina Maggenti, Hart R: Paxman

1974, s. nov. OTHER GENERA

Nofhanguina Whitehead, 1959 Afrina g. nov. Orrina g. nov. Szzba.ngzzina Paramonov, 1965 Difylenchus Filipjev, 1936 Nofhofylenchzrs Thorne, 1941 Dipfenchzzs Iihan, Chawla & Se§hadri, 1969

Revzza Némwfol. 4 (1) : 23-36 (1981)

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Genera of Anguinidae

The proposed diagnosis limits the Anguinidae to the genera with labial amphids, no cardia, an axial spermat,heca, and with a sphincter between uterine sac, and c.rustaformeria. Pres- ence or absence of refrac.tive thic.kenings in median bulb is considered at present as a generic. character. Therefore, the subfamily Nothoty- lenchinae is synonymised with Anguinidae, and Nothotylenchzzs is inc.luded in the lat.ter family. The taxonomie position of other genera of Not.hot.ylenchidae should be furt.her inves- tigated. 1 share the opinions of Sumenkova (1975) and Geraert (1976) t.hat Ditylenchus and Notholyler&us may be identical. However, both

. certainly need more work and regrouping, and 1 prefer not to propose any change at this time.

KEY TO THE GENERA OF ANGUINIDAE

1.

2.

3.

4.

5.

6.

7.

Gonads with many cells in circumference . . . . . 2 Gonads with one or two cells in circumference . . 4 Mrdian bulb large, with refract.ive thickenings . 3 Median bulb fusiform, without thickening . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nothanyuina Crustaformeria a long multinucleate tube ; no muc.ro at tail tip. . . . . . . . . . . . . . . . . . . . Anguina Crustaformeria as four rows of uells, fourteen cells in a row ; tail tip with mucro . . . . . . . . . . . . Afrina hIedian bulb with refrac.tive thickenings . . . . . . . 5 No refractive thickening in oesophageal lumen 7 Crustaformeria as quadricolumella . . . . . . . . . . . 6 Crustaformeria as four rows of ~Ils, eight to twelve cellsinürow . . . . . . . . . . . . . . . . . . . . . Su banguina Posterior uterine sac present ; vagina perpendicular to the body axis . . . . . . . . . . . . . . . . Difylenchus No posterior uterine sac ; vagina oblique to the body axis . . . . . . . . . . . . . . . . , . . . , . Dipfenchus Median bulb fusiform ; oesophageal glands in a terminal bulb offset from intestine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nofhotylenchzzs Rledian bulb absent ; oesophageal glands in a lobe ; dorsal nucleus posterior t.o oesophago-intestinal junction . . . . . . . . . . . . . . . . . . . . . . . . . . . Orrina

Genus Anguina Scopoli, 1777

= Paranguina Kirjanova, 1955, s. nov.

= Cynipanguina Maggenti, Hart & Paxman, 1974, s. nov.

DIAC;NOSIS : Anguinidae. Mature females swollen. Median oesophageal bulb with refractive thick- enings. Isthmus separated from terminal bulb

Revue Nématol. 4 (1) : 23-34 (1981)

by a constriction. C&staformeria, a long multi- nucleate tube composed of more or less hexag- onal cells. Ovary with many oogonia in circum- ference. Testes usually with two flexures. Bursa enclose most of the t,ail, except the tip. Form galls on stems, leaves ancl/or inflorescences of grasses, A. balsamophila is an exception, infesting a dicot,yledon.

TYPE SPECIES

A. trifici (Steinbuch, 1799) Chitwood, 1935

OTHER SPECIES

A. agropyri Kirjanova, 1955 ’ = Paranguina agropyri Kirjanova, 1955

A. agropyronifloris Norton, 1965

A. agrostis (Steinbuch, 1799) Filipjev, 1936 = A. poophila Kirjanova, 1952, s. nov.

A. amsinckiae (Steiner & Sc.ott, 1935) Thorne, 1961

A. australis Steiner, 1940

A. balsamophila (Thorne, 1926) Filipjev, 1936

A. danthoniae (Maggenti, Hart & Paxman, 1974) comb. nov. = Cynipanguina danthoniae Maggenti, Hart

L% Paxman, 1974

A. fzznesta Price, Fisher & Kerr, 1979

A. graminis (Hardy, 1850) Filipjev, 1936

A. microlaenae [Fawcett, 1938) Steiner, 1940

Pa.ranguina and Cynipanguina are synony- mised with Angzzina bec.ause studies of type material and additional collections did not show any differences. The type specimens of the type species of bot,h genera were killed in cold fixa- t,ive, c.ausing much distort.ion. Well fixed and processed spec.imens of A. agropyri, type spec.ies of Paranguina, showed that the additional oesophageal gland described by Kirjanova (1955) was a fixation artefact. Since Kirjanova (1955) in the same paper used both P. agropyri and A. agropyri, no new combination is proposed.

The Stern-like oesophageal extension described as a main differentiating character for Cynipan- guina is also a fixation artefact. The same conditions were seen among species of Angzzina.

25

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M. W. Brzeski

Fig. 1. Anguina tritici. A : oesophageal region, B : head, C : tail of male, D : posterior end of fcmale.

26 Rrvzrc Nthniol. 4 (1) : 23-34 (1951)

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Genera of Anguinidae

Fig. 2. Nothanguina cecidoplastes. A : oesophageal region, B : posterior end of female.

Revue Nématol. 4 (1): 23-34 (1981) 27

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AI. IV. Brzeski

A. poophila is considered synonymous with A. agrostis because st.udies of t*ypes and other collections did not show any differences. Bot,11 were found on the same host.

Adults of different, species of Anguina. are morphologically indistinguishable, although they develop on dif’ferent hosts in widely separated geographical areas. More work is needed before a full revision of this genus cari be presented.

Only those synonyms of the species mentioned in this paper are listed, when they are not included in the exc.ellent, c.hec,k 1ist.s of Tarjan and Hopper (1974, 1977, 1978).

Genus Nothanguina Whit,ehead, 1959

DIAGNOSIS : Anguinidae. Mature females swollen. Median bulb poorly developed, wit.hout refractive thickenings. Isthmus separated from basa1 bulb by a constriction. Crust.aformeria a long muhinucleate t.ube composed of more or less hexagonal cells. Ovary with many oogunia in circumference. Testes reflesed. Bursa does not enclose tail tip. Spicules complicated, no gubernaculum. Forms galls on leaves of grasses.

TYPE AND UNIQUE SPECIES

N. cecidoplastes (T. Coodey, 1934) Whit.ehead, 1959

DIFFERENTIAL DIAGNOSIÇ : This genus is closest t.o Angnina, but differs by having a poorly developed median bulb without refractive t.hick- enings, and by lacking a gubernac.ulum. Bursa smaller than m Angzzina., but since only one species of Nothangzzina is known this cannot be considered as a generic c.haracter.

Genus Afrina g. nov.

DIAGNOSIS : Anguinidae. Mature females swollen. Median bulb with refrac,tive thickenings. Isthmus separated by a c.onstriction from terminal bulb. Crustaformeria formed by four rows of cells, fourteen cells in a row. Test,es often with two flexures. Tail of both sexes conical, tail t.ip mucronate. Bursa notched at

28

the post,erior end, almost reaches t.ip but does net enc.lose mucro. Forms galls on leaves of grasses.

TYPE SPECIES

A. hyparrheniae (Corbet.t, 1966) comb. nov. = Angzzina hyparrheniae Corbett, 1966

OTHER SPECIES

A. tumefaciens (Coob, 1932) comb. nov. = Anguina tztmefaciens (Cobb, 1932) Filip-

jev & Sc.huurmans St.ekhoven, 1941

DIFFERENTIAL DIAGNOSIS : Afrina g. nov. occu- pies an intermediate posit,ion between Anguina and Subangzzina. It. is close to Anguina by oesophageal constriction, many oogonia in circumference, and flexures of testes, while similarity to Subanguina is indic.at.ed by struc- ture of crustaformeria.

Angzzina spermophaga St.einer, 1937 most. probably belongs t.o this genus but no specimens were available and a final clec.ision cannot be made.

Specimens of Afrina g. nov. examined were from southern Afric.a and Australia.

Genus Orpina g. nov.

DIAGNOSIS : Anguinidae. Mature females slightly or not swollen. No median bulb and no refractive thickenings in oesophageal lumen. Oesophageal glands form a lobe overlapping intestine. Nucleus of dorsal gland posterior to oesophago-int.estinal juction. Isthmus short and narrow. Crust,a- formeria in form of quadricolumella. Ovary with few oogonia in circumference. Forms galls on leaves of Solanum elaeagnifolizzm Cav.

TYPE AND UNIQUE SPECIES

0. phyllobia (Thorne, 1934) comb. nov. = Nothanguina phyllobia (Thorne, 1934)

Thorne, 1961

Revue Nt!matol. 4 (1) : 23-34 (1981)

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Genera of Anguinidae

B \\

Fig. 3. Afrina hyparrheniae. A : oesophageal region, B : head, C : posterior end of female, D : posterior end of male, E : spicules and gubernaculum of male.

Revue Nématol. 4 (1) : 23-34 (1981) 29

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M. W. Brzeski

II II

D

Fig. 4. Orrina phyllobia. A : oesophageal regionj B : head, C : tail of male, D : posterior end of female, E : part of female reproductive system.

30 Revue Nématol. 4 (1) : 23-34 (1981)

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Genera of Anguinidae

DIFFERENTIAL DIAGNOSIS: Orrina g. nov. st.ands somewhat separately by having overlapping oesophageal lobe.

0. phyllobia is known from Texas and Arizona only.

The generic name Orrina g. nov. is given in honour of Dr. C. C. Orr, in recognition of bis excellent studies on the biology of this spec.ies.

Genus Subanguina Paramonov, 196s

DIAGNOSIS : Anguinidae. Mature females may or may not be swollen. Median bulb with refractive thickenings. Isthmus separat.ed by constriction. Crustaformeria with four rows of cells, 8-12 cells in a row. Ovary with few oogonia in c.ircum- ference. Testes usually not reflexed. Forms galls on roots, st.ems, leaves and/or inflorescences of plants belonging to many families.

TYPE SPECIES

S. radicicola (Greeff, 1972) Paramonov, 1968

OTHER SPECIES

S. askenasyi (Bütschli, 1873) comb. nov. = Anguina askenasyi (Bütschli, 1873)

Krall, 1971

S. brenani (T. Goodey, 1945) comb. nov. = Anguina brenani (T. Goodey, 1945)

Kirjanova & Krall, 1971

S. calamagrostis (Wu, 1967) comb. nov. = Anguina cala.magrostis Wu, 1967

S. centaurea (Kirjanova SL Ivanova, 1968) çomb. nov. = Paranguina cenlaurea Kirjanova & Iva-

nova, 1968

S. chartolepidis (Poghossian, 1966) comb. nov. = Anguina chartolepidis Poghossian, 1966

S. cousiniar (Kirjanova & Ivanova, 1968) comb. nov. = Paranguina cousiniae Kirjanova $ Iva-

nova, 1968 S. ferrzlae (Ivanova, 1977) comb. nov.

= Anguina ferulae Ivanova, 1977

Revue Nématol. 4 (1) : 23-34 (1981)

S. graminophila (T. Goodey, 1933) comb. nov. = Anguina graminophila (T. Goodey, 1933)

Thorne, 1961 = A. tridomina Kirjanova, 1958, s. nov.

S. kopetdaghica (Kirjanova & Shagalina, 1969) comb. nov. = Anguina kopefdaghica Kirjanova & Sha-

galina, 1969

S. millefolii (Low, 1874) comb. nov. = Anguina millefolii (LOw, 1874) Filipjev,

1936 S. mobilis (Chit & Fisher, 1975) comb. nov.

= Anguina mobilis Chit & Fisher, 1975

S. montana (Kirjanova & Ivanova, 1968) comb. nov. = Paranguina montana Kirjanova & Iva-

nova, 1968

S. moxae (Yokoo & Choi, 1968) comb. nov. = Anguina moxae Yokoo 6% Chai, 1968

S. picridis (Kirjanova, 1944) comb. nov. = Paranguina picridis (Kirjanova, 1944)

Kirjanova & Ivanova, 1968 S. pZa.nfaginis (Hirschmann, 1978) comb. nov.

= Angzzina plantaginis Hirschmann, 1978 S. polygoni (Poghossian, 1966) comb. nov.

= Anguina polygoni Poghossian, 1966 S. varsobica (Kirjanova & Ivanova, 1968)

comb. nov. = Paranguina aarsobica Kirjanova Le, Iva-

nova, 1968

DIFFERENTIAL DIAGNOSIS : Subanguina differs from Ditylenchus by the structure of its crusta- formeria.

Anguina pustulicola Thorne, 1934, and A. klebahni Goffart 1942, may also belong to the genus Suba.nguina. However, the original desc.riptions are not adequate t.o classify them to genus, and specimens were not available for examination. Therefore they are placed in gelzzzs et species inquirenda.

The genus Subanguina contains many sibling spec,ies and detailed biological investigations are necessary before any revision cari be presented.

Genus Ditylenchus Filipjev, 1936

DIAGNOSIS : Anguinidae. Mature females net. swollen. Median bulb with refractive thicke-

31

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M. W. Brzeski

Fig. 5. Subanpzina radicicola. A : oesophageal region, B : head, C : tail of male, D : spicule and gubernac,ulum, E : posterior part of female showing reproductive system.

32. Revue N~matol. 4 (1) : 23-34 (198-l)

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Genera of Anguinidar

nings. Isthmus not separated hy constriction. Crustaformeria in form of yuadricolumella. Ovary wit.h l-2 oogonia in c.ircumference. jTagina perpendic.ular to body axis. Testes usually without, flexures. Tail of both sexes conical, rourided or pointer1 at the end. Soi1 inhabiting fungal feeders or obligatory parasites of higher plants.

TYPE SPECIES '

D. dipsaci (Kühn, 1559) Filipjev, 1936

DTFFEAENTIAL DI,~GNoSIS : Difylenchus differs from Nothotylenchus by the presence of refractive thickenings in median bulb, and from Srtbangui- ncl by the structure of crustaformeria.

This genus probably contains several sibling species.

Genus Nothotylenchus Thorne, 1941

DIAGNOSIS : Anguinidae. Mature females not swollen. Median bulb weakly developed or absent., wit.hout refract.ive thickenings. Isthmus net. separated by a const.ric.tion. Crustaformeria in form of yuadricolumella. Ovary with one or two oogonia in c.ircumference. Vagina perpen- diçular to body axis. Testes usually not reflexed. Bursa short., or estending up to 2/3 of tail length. Soi1 and fresh water nematodes, probably niost, feed on fungi.

TY-PE SPECIES

N. acris Thorne 1941 7

DIFFERENTIAL DIAGNOSIS : It differs from Dity- lerzchus by structure of oesophagus only.

Sumenkova (1975) in her review of t.his genus mentioned eight. groups of species. Di~ylevzchus and Nothotylenchus need very detailed taxonomie stuclies ancl regrouping.

Genus Diptenchus Iihan, Chawla & Seshadri, 1969

DIAGNOSIS : Anguinidae. Female not swollen. Median bulb with refractive thickenings. Isth-

Revue Névnafol. 4 (1) : 2.3-36 (1981)

mus not separated by constriction. Crustafor- meria in form of quadricolumella. Ovary wit.1~ one or two oogonia in circumference. Vagina oblique t.o the body asis, vaginal wa.lls refraçt-ive. No post-vulval uterine sac. Tail of bot11 sexes conical.

TYPE AND UNIQUE SPECIES

D. indicus Iihan, Chawla L% Seshadri, 1969

DIFFERENTIAL DIAGNOSIS : Diptcnchrrs differs from Ditylenchus by structure of t,he vagina and lack of post-vulval ut.erine sac.

Type specimens frorn Dr. Iihan fitt.ed the origmal description, although cardia and phas- mids were net. seen.

D. indicrrs is known from one collection from New Delhi.

Many prople c‘ontribut rd to this lvorlc by sending slides or discussing various aspects of &lguinidae taxonomy. 1 want to express to a11 of t.hem my sinccre appreciat.ion. Most cordial thanks are due to Eino Krall who put. bis collect.ion at. my disposa1 md whose drep interest. in nemnt.ode evolution st.imulated this mark. The following hrlprd in various ways, and t.his papcr would not have bcen completed without. their efforts : .J.G. Baldwin, AF. Bird, R.H. Brown, W.N. Chizhov, R.C. Colbran, E. C&a&, U.J. Hooper, T.S. IVXlOW1, E. KhilIl, P.A.A. LOOf, R. SI~L~O~, E.M.. Noffsinger, 13.C. Norton, C.C. Orr, T.S. Skarbi- lovich, G.1. Solov’eva, D. St.urhan, 13. Stynes and .H. Triant.aphyllou.

REFERISYCES

~~OR~SY, 1. (1976). Evnlution us a hasis for systemu- tization of ntmatorles. Budapest,, hltadémiai RiadO, 488 p.

C~OI, Y.E. k LOOF, P.A.A. (1973). Redescription of Anguina moxae \wokoo 8~ Chai, 1968 (Tylenehida). Nemaiologica, 19 : 185-292.

GERAERT, E. (1976). The femalr reproductive systrm in Deladenus and Hexxtylus with a redefinition of the oviduct in the Tylrnchida (Nematoda). Nema- fologica, ?2 : XST-145.

C;ERAERT, E. & KIIEIRI, A. (1971)). The frmale gonads and oesophageal structure in thc genus l?sezzcZlznlen- ehus (Nematocla : Tylenchida). Nemafologica, 16 : 197-202.

33

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GOLDEN, X.M. (1971). Classitication of t.he genera and higher categories of the ordw Tylenchida (Nema- t.oda). In : Zuckwman, RM., RIai, W.F. Sr Rohde, R.A. (Eds) Plant Parnsific A7r77~afodcs, Vol. 1, New York, Academic. Presa : 191-232.

HOOPER, D.J. (19%). S~.PUctMt? :~rld (;kIssifkXItiOR of nemat.odes. In : Southry, J.F. (Ed.) Planf Nema- fology, London, H.bl.S.0. : 3-45.

HOPPER, B.E. 62 TAR~TAN, A.(:. (1977). Srzpplanwnt for the ycar 1972 fo nomencltrloric~l rompilafion of planf and soil ne7nafodes. Ott.awa, X@c. Canada Res. Uranch, 17 p.

HOPFER, B.E. & TARJXY, X.C. (1!)78). Snp@wlent for fhe year 197s fo n»nleiiclaforiul compilafion of planf nnd soi1 nemafodes. Ott.awva, Agric. Canada Rcs. Branch, 2.1 p.

KIRJANOVA, E.S. (19%). Pyrejnaja ugrica - Parm- guina agropyri Kirjanova, geu. et. sp. n. (Nemato- des). Trzzdy mol. Inst., 18 : 42-52.

PARAMONOV, A.A. (1962). 0~7107.~~ PtogeLnintoZo!lii. Tom 1. &Io&va, Izdat. Xlrad. Nauk SSSR, 480 p.

PARAXIONOV, AA. (19ïOj. Osnoz~y ~itoyelminfologii. Tom III. Moskva, Izdat.. hkad. Nauk SSSR, 253 p.

SIDDTQI, h1.R. (1971). Structure of t.he oesophngus in the classification of the sqwrfamily Tylenchoidea (Nematoda). Intiitrn J. Nc771trtol., 1 :‘?.a-43.

SUMENICOVA, N.I. (1975). i\rrmafody rcrstenij i pochr~y. NeofyZenchoitEen. MOS~VL~, NRIlkR, l!)T p.

TARJXN, A.G. & HOPPF.R, l3.E. (1971). Nomendnforicrf compilation of plunf n77d soi1 7zemafodes. DeLeon Springs, U.S.A. Soc. oï Ncmat.ologist.<, 419 p.

THORNE, CI. (1961). Prinriples of 72crnatolo!~g. New York, McGraw-Hi11 Book C;o.! 553 p.

AccepfP. pour publicnfion le 20 mari 19SO.

\Vu, L.T. ( 1967). Angrzina ctrlnmtrgrosti.s, 0 I~CW- speçies from gras wilh an rmrndi~t.ion of the geueric. cha.ractrrs for the generti Angrrinn Scopoli, 1777 and Difylenchus Filikijev, 1036 (Tylenc.hidae : Nemil- toda). Chn. J. Zool., 45 : lOl13-11110.

NOTE ADDED ON PROO"

Two more genera mere recent.ly proposecl within Anguiniclae :

Hdrrotrizgrrirzn Tchizhov, 1950, cannot. be dif- ferentiat.ed frorn Subcm~gui7za ad rnust be con- siderecl it,s junior synonyin. ,4ngnin.a grcmino- philn is nientioned as t;ype species of Hrtm~- myuinc7, but. this taxon is not c.it.ecl as new combinat.ion.

Saficzn~n~cr Siddiqi, 1980, c-liffers from Dihy- lcncl~us 1:)~ Iiaving lon g oesophageal lobe. No differenws are ment.ioned between Sufiiuwmtc and Ditylrmhrrs species with oesophagus over- lapping int.est.ine, as D. &drrzt:tnr ancl D. ~ulycc- liophayus.

SIDDIQI, h1. R. (1980). Two new nematodr grnrra, Safianernn Anguinidae and Discofglenchus Tylen- chidar, wit.h descriptions of thrrt: new species. Proc. helminfh. Soc. Wnsh., 47 : X5-M.

TCHIZI-IOV, T. N. (1980). 0 t.alisonomit.chrskam statuse nekotorykh vidov roda d72gzrincl Scopoli, 1777. Biul. T‘sfwjum. 171st. Gelmirxf., 26 : 83-W

34 Retw Némnfol. G! (1) : 23-32 (1961)