Systematics of the Trichodoridae (Nematoda) with Keys to Their Species

Systernatics of the Trichodoridae (Nematoda) with keys to their species

Wilfrida DECRAEMER Instituut uoor Dierkunde, Rijksuniversifeif, Ledeganckstraat 35, 9000 Gent, Belgium.

SUMMARY Keys are given for the identification of species in the Trichodoridae, partly based on the keys of Loof (1975) and Siddiqi (1974). Recent additions are included and for al1 species a drawing is given of the structure referred to in the key. The genus Paratrichodorus and the diagnostic value of the subgenera Paratrichodorus, Atlantadorus and Nanidorus are discussed.

RÉSUMB Systématiquedes Trichodoridae (Nematoda) et clés de déterminationdes espèces L’auteur propose de nouvelles clés de détermination des espèces de Trichodoridae fondées sur les clés de Loof (1975) et de Siddiqi (1974), et tenant compte desdonnées les plus récentes. Des illustrations sont fournies pour les caractères les plus importants de chaque espèce. Le genre Paratrichodorus et, la,valeur dessous-genres Paratricho- dorus, Atlantadorus et Nanidorus sont discutés.

The Trichodoridaeare economically, a very Superfamily TRICHODOROIDEA important group of nematodes as ectoparasites (Thorne, 1935) Siddiqi, 1961 on the roots of many crop plants and as vectors of tobacco rattle virus and pea early browning virus. Despite the ease with which nematodes THE GENERA OF THE TRICHODORIDAE(Thorne, can acquire viruses from plants there are rela- 1935) Clark, 1961 tively few vectors. Of some 35 species of Tricho- doridae, at least eleven transmit tobra-viruses Until recently the family Trichodoridae con- (Hooper, 1973),so identification to species level tained only the genus Trichodorus Cobb, 1913. may be very important. Siddiqi (1974) Split it into two genera,: Triclzo- During mystudy on Trichodoridae 1 came dorus and Paratriclzodorus; at the same time across several inadequacies in available descrip- he divided Paratrichodorus into threesubgenera : tionsand keys; thesechanges, togetherwith Paratrichodorus,Atlantadorus and Nanidorus. recentadditions made it necessary to modify Recentlythe monodelphic species (except the Beys of Loof (1975) and Siddiqi(1974). P. roestindicus) were included in two new genera : For al1 the species 1 haveincluded a drawing Monotrichodorus hndr&sy, 1976 for species of the structure referred to in the key. closely related to Trichodorus. and Allotriclzo- dorus Rodriguez-M., Sher & Siddiqi,1978 for Suborder DIPHTHEROPHORINA species closely related to Paratrichodorus (Atlan- Coomans & Loof, 1970 tadorus).

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Genus Trichodorus Cobb, 1913 S. str. 4. - Tailcontour evenly rounded,terminal cuticlenot conspicuously thickened; spicules not setose (Pl. II : Fig. 3) DEFINITION T. aequalis Allen, 1957 - Tail contour not evenly rounded, terminal ' Trichodoridae.Cuticle notstrongly swollen cuticle conspicuously thickened (P. II : upon fixation.. Posterior ventrosublateral oeso- Fig.4) ...... 5 phagealglands not overlapping the intestine, nor does the intestine anteriorly partly overlap 5. - Spicules not setose, almost st,raight (Pl.II : Figs. 6, 7) the oesophagus.Dorsal oesophageal gland T. hooperi Loof, 1973 nucleus usually situated between both pairs of - Spicules setose, curved proximally (Pl. II : ventro-sublateral gland nuclei, near the poste- Figs. 4, 5) rior pair. Female reproductive system didelphic; T. sparsus Szczygiel, 1968 spermathecaepresent; a pair of lateralbody 6. - Excretory pore situated between the 2nd pores present within one body;width from the and 3rd ventromedian cervical papillae

' vulva.Vaginal sderotization well developed. (Pl. 1 : Fig. 8; Pl. II : Fig. 10; tail region + Males conlmon ; diagonalcopulatory muscles copulatory apparatus) T. pakistanensis Siddiqi, 1962 well developed, extending far anteriorly t,o the - Threeventromedian cervical papillae spicular region, and responsible for ventral locatedanterior to theexcretory pore curvature of themale, tail in death. Usually (Pl. 1 : Fig.9) ...... 7 withoutcaudal alae, exceptionally with small 7. - Onchiostyle length 72-75 Pm; body length caudal alae. Spicules arcuate ; suspensor muscles 500-590 pm (Pl. II : Fig. 8 : copulatory conspicuous. Preanal supplements three, excep- apparatus) tionallyfour, more or less evenlyspaced, the T. longistylus Yokoo, 1964 two anterior ones nearly always located anterior - Onchiostyle length 52-54 Pm; body length to the spicules. Number' of ventromedian 570-610 Pm; (Pl. 1 : Fig. 9 : oesophageal region;Pl. II : Fig. 9;tail region + cervical pores often 2 or 3, rarely 1 or O. copulatory apparatus) T. kurunzeensis Yokoo, 1966 - Onchiostyle length 58-60 Pm; body length TYPE SPECIES 510-850 Pm; spicule length 42-45 Pm; (Pl. III : Fig. 13; tail region + copulatory Trichodorusprimitiuus (De Man, 1880) Mico- apparatus) let.zky, 1922. T. cedarus Yokoo; 1964 Syn. Dorylaimus prinzitiuus De Man, 1880. 8. - Onchiostyle length over 130 Pm; (Pl. II : Fig.11 ; tail region f copulatoryappa- KEY TO Trichodorus MALES ratus) T. elegans Allen, 1957 1. - No ventromedian cervical papillae in t.he - Onchiostyle lengthunder 85 Pm ...... 9 onchiost.yle region (Pl. 1 : Fig. 1) ...... 2 - One or two ventromedian cervical papillae 9. - Supplementary papillae situated anterior inthe onchiostyle region (Pl. 1 : Fig. 2, to spicules (Pl. II : Fig. 13) cf. remarlr 1) ...... 13 T. proximus Allen, 1957 - Posteriorsupplementary papillae within 2. - Ventromedian cervical papillae O ; (as in reach of spicules ...... 10 tail region + copulatory apparatus Pl. II : Fig. 15; remark 2) 10. - Spicules irregularlytapering, slightly T. obscurus Allen, 1957 + T. obtusus Cobb, 1913 expanded in distal third (Pl. II : Fig. 12) - Ventromediancervical papillae 1 (Pl. 1 : T. cottieri Clark, 1963 Fig.3) ...... 8 - Spicules regularly tapering, not expanded - Ventromediancervical papillae 2 (cf. in distal third (Pl. III : Figs. 2,3) ...... 11 rem ark 3) 3 3) remark ...... - Ventromediancervical papillae 3 ...... 6 Il. - First and second supplement close to the 3. - Two supplements within reach of spicules; anus, opposite distal half of spicules when spicules striated (Pl. II : Figs., 1.2) completelyretracted (Pl. III : Fig. 3) T. borneoensis Hooper, 1962 T. intermedius Rodriguez-M. & Bell, 1978 - One supplement within reach of spicules; - Second supplementanterior to spicules ...... whencompletely retracted ...... 12 -. . --- . spiculesstriated not - -. - ~. 4 -. - .- . 82 Rerrue Nématol. 3 (1) : 81-99 (1980) Pl. 1. Oesophageal region in Trichodorus (scale = 20 Pm). 1 : T. hooperi (after Loof,, 1973). 2 : T. primitivus. 3 : T. californicus (specimen, courtesy E.M. Noffsinger). 4 : T. lusitanicus (after Siddiqi, 1974). 5 : T. viruliferus (specimen, courtesy D. De Waele). 6 : T. similis. 7 : T. castellanensis (redrawn after Arias - Delgado, Jimenez-Millan CE Lopez-Pedregal, 1965). 8 : T. pakistanensis (after Siddiqi, 1962). 9 : T.lcurumeensis (redrawn after Yokoo, 1966). 10 : T. variopapillatus (after Hooper, 1972). 3

Pl. II. Copulatory apparatus in Triehodorus (scale = 520 Fm). 1-52 : T. borneoensis (after Hooper, 1962). 3: T. aequalis (after Loof, 1973). 4 : T. sparsus (after Loof, 1973). 5 : T. sparsus. 6-7 : T. hooperi (after Loof, 1973). 8 : T. longistylus (after Yoko0;1964):9 : T. kurumeensis--(afterYolroo; 1966). -10 : T.-pakistanensis (after Siddiqi,--1962).-11 : T. lusitanicus (after Siddiqi,1974). 12 : T.cottieri (afterClark, 1963). 13: T. proximus (paratype, courtesy E.M. Noffsinger). 14 : T. obscurus (specimen, courtesy E.M. Noffsinger). 15 : T. obtusus (redrawn after Cobb, 1913). Pl. III. Copulatory apparatus in Trichodorus (scale = 20 Pm). 1 : T. californicus (specimen, courtesy E.M. Noffsin- ger). 2 : T. dilatatus (after Rodriguez-M. & Bell, 1978). 3 : T. intermedius (after Rodriguez-M. & Bell, 1978). 4 : T. cylindricus (specimen, courtesy J.W. Seinhorst). 5 : T. castellanensis (after Arias-Delgado,Jimenez-Millan & Lopez-Pedregal, 1965). 6 : T. variopapillatus (specimen, courtesy A. Coomans). 7 : T. similis. 8 : T. uiruliferus (one specimen, courtesy J.W. Seinhorst). 9: T. similis. 9 : T. intermedius (after Rodriguez-M. & Bell, 1978). 10 : T. uelatus (after Hooper, 1972). 11 : T. lusifanicus (after Siddiqi, 1974). 12 : T. primitivus. 13 : T. cedarus (after Yokoo, 1964). Pl. IV. Vaginal region in Trichodorus (scale = 10 Fm). 1 : T. elegans (redrawn after Allen, 1957). 2-4 : T. variopapillatus (specimens, courtesy P.A.A. Loof). 5: T. velatus (redrawn after Hooper, 197'2). 6 : T. similis. 7-9 : T. cylindricus (specimens, courtesy J.W. Seinhorst, D. De Waele). 10-12 : T. similis (specimens, courtesy J.W. Seinhorst). 13 : T. cedarus (redrawn after Mamiya, 1967). 14 : T. intermedius (redrawn aft,er Rodriguez-M. & Bell, 1978). 15 : T. cottieri (redrawn after Clark, 1963). 16-22 : T. primitivus. 23 : T. prozimus (redrawn after Allen, 1957). 24-28 : T. sparsus (specimens, courtesy P.A.A. Loof, J.W. Seinhorst; 24-25 : specimens from Koningstadten, Germany, 26 : from Oosterbeek, Netherland, 28 : from Hundsdorf, Germany).29 : T. aequalis (specimen, courtesy E.M. Noffsin- ger). 30 : T. pakistanensis (redrawn after Siddiqi, 1962). 31-33 : T. viruliferus (specimens, courtesy J.W. Seinhorst, D. De Waele). 34 : T. longistylus (redrawn after Yokoo, 1964). 35 : T. borneoensis (redrawn after Hooper, 1962). 36 : T. hooperi (paratype, courtesy P.-A..A; Loof). 37. : T. lusitanicus (redrawn- after Siddiqi, -1974). 38 ,: T. californicus (paratype, courtesy E.M. Noffsinger). 39 : T. obscurus (specimen, courtesy E.M. Noffsinger). 40 : T. dilatatus (redrawn after Rodriguez-M. LE Bell, 1978). Systematics of the Trichodoridae

12. - Two verysimilar species (synonymous? ventromedian cervical papillae; (Pl. III : cf. remark4). Third supplement 1.1 body , Fig. 13 : tail region + copulatoryappa- ’ widthanterior to second supplement ratus) (Pl. III : Fig. 2) T. cedarus Yokoo, 1964 T. dilatatus Rodriguez-M. & Bell, 1978 - Thirdsupplement 0.8-1.0 bodywidth anteriorto second supplement(Pl. III : Remarlrsto the key of Trichodorus nal les Fig. 1; Pl. 1 : Fig. 3 : oesophageal region) T. californicus Allen, 1957 1.In the original figure of T. cedarus 13. - One ventromedian cervical papilla located Yokoo, 1964 two ventromedian cervical papillae behind the excretory pore (Pl. 1 : Fig. 7; arelocated within the onchiostyleregion. Pl. III : Fig. 5 : tail region + copulatory However, since thetype specimens arebadly apparatus) fixed we cannot be certain that in well fixed T. castellanensis Arias et al., 1965 - No ventromedian cervical papilla located animals no cervical papillae would be found posteriorto the escretory pore (Pl. 1 : within the onchiostyleregion. Consequently I Fig.2) ...... 14 mentioned both possibilities within the key. 14. - ‘Spicules distallyvery slender so that Mamiya (1967) observed no cervicalpapillae gubernaculum mainly appears ventral to within the onchiostyle region in other specimens spicules (Pl. III : Fig. 12; Pl. 1 : Fig. 2 : than the type specimens. oesophageal region) T. primitiuus (De Man, 1880) Micoletzky, 1922 2. As Loof (1975) pointed out, the status of . - Gubernaculum never ventral to spicules . . 15 T. obtusus Cobb, 1913 has never been clear. 15. - Spicules transverselystriated; tail with As noredescription is available we canonly small caudal alae, narrow part of spicules rely on the original drawing and description of with “bristles” (Pl. III : Fig. 4) a male without ventromedian cervical papillae. T. cylindricus Hooper, 1962 - Spicules not striated; caudal alae absent . 16 Thetrichodorid collected byThprne in 1931 from the type locality was described by Allen 16. - Spicules constrictednear middle, with capitulumrather broad, only slightly (1957) as T. obscurus. Both species were very offset (Pl. III : Fig. 8) ...... 17 similar, the only difference pointed out by Loof - Spicules not constricted nearmiddle, with (1975) is that thespicules are much less strongly rounded offset capitulum (Pl. III : Fig. 6) . 19 curvedin T. obscurus. 17. - Two ventromediancervical papillae present (Pl. 1 : Fig. 4; Pl. III : Fig. 11 : 3.The characters used until now to dis- tail region + copulatory apparatus) tinguish the four species : T. borneoensis, T. lusitanicus Siddiqi, 1974 T. aequulis, T. sparsus and T. lzooperi (cf. Loof, - Threeventromedian cervical papillae 1975) thickened terminal cuticle and tail contour present(Pl. 1 : Fig.9) ...... 18 arein my opinion not so apparent especially 18. - Spicules stout,ventrally expanded in between T. borneoensis andthe other species. distal third (Pl. III : Fig. 10) T. uelatus Hooper, 1972 Therefore 1 used other cl1aracteristic.s inthe - Spicules moreslender, notventrally key to distinguish T. borneoensis. from the expandedin distal third (Pl. III : Fig. 8; ’ other species. Pl. 1 : Fig. 5; oesophageal region) T. viruliferus Hooper, 1963 4. Inthe males both species have to be , distinguished by a very small difference in the 19. - Onchiostylelength 35-45 [lm;spicule length 35-40 pi; threeventromedian location of supplementarypapillae. Inthe cervical papillae (remarlr 5; Pl. 1 : Fig. 6 : holotype male of T. dilatatus the distance Pl. III : Figs. 7,9; tailregion + c.opulatory between the anteriorsupplementary papilla apparatus) and the second supplementary papilla is 42 pm, T. similis Seinhorst, 1963 - Onchiostylelength 41-58 Pm; spicule i.e. 1.1 times the bodywidth at this level. In length 40-45 Fm; spicules slender; two or four males from Allen’s collection of T. califor- threeventromedian cervical papillae rzicus the distance between both papillae (measu- (remarlr 5 : Pl. 1 : Fig. 10; Pl. III : Fig. 6; red dongthe ventral line) varied between copulatory apparatus) 35-41 pm i.e. 0.8-1.0 times thebody width. T. uariopapillatus Hooper, 1972 - Onchiostylelength 58-60 Pm; spicule Consequently theonly difference mentioned length-42-45 Pm; spicules stout;three between T. dilatatus and T. californicus (cf.

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Rodriquèz-M. & Bell, 1978) becomes negligible. 7. - A pair of ventrosubmedian pores behind T.' californicus shows some variationin the the vulva; sclerotization of vagina appearing parallel to the vaginal lumenin lateral position of the lateral cervical pore in relation view; onchiostyle length58-62 pm (Pl. IV : to thesingle ventral cervical pore, the excretory Fig. 13) pore and the nerve ring. 1 observed the lateral T. cedarus Yokoo, 1964 cervicalpore varying between a position just - No ventrosubmedian pores hehindthe behind the nerve ring, i.e. 12.5 pm anterior to vulva but a pair of lateral pores ...... 8 the ventral c,ervical pore to a position slightly 8. - Sclerot.ization of vagina appearing as more or less triangular,prominent, pieces in (3.5 Pm) behind theexcretory porei.e. about lateral view ...... 9 26 pm behind the nervering. - Sclerotization of vagina differently struc- 5. Themeasurements for T. similis an'd turedand smaller (Pl.IV : Fig.15) ; onchiostyle length 57-68 pm; body length T. variopupillatus include persona1 measure- 700-1.010 pm ments, that extend the range so far published. T. cotfieri Clark, 1963 9. - Onchiostyle length 62-75 pm; body length 680-1.130 pm;sclerotization of vagina KEY TO Trichodorus FEMALES large, vaginal region rounded(Pl. IV : Fig. 14) T. intermedius Rodriguez-M & Bell, 1978 . 1. - Onchiostyle length 140-150 pm (Pl.IV : - Onchiostyle length 66 pm;body length Fig. 1 : vulva region) 1.170 pm; sclerotization of vagina smaller, T. elegans Allen, 1957 vaginal region cylindrical (Pl, IV: Fig. 39) - Onchiostyle lengthunder 100 pm ...... 2 T. obscurus Allen, 1957 2. - Sclerotized pieces at .vagina rather close 10. - Sclerotized ring of vagina broad; in lateral to one anot.her (i.e. aperture of the sclero- view sclerotization of vagina appearing as tized ririg rather small (Pl. IV : Fig. 2) . . 3 tworods parallel to the vaginallumen; - Sclerotized pieces at vagina more clearly threepairs of lateralpores; onchiostyle separated (i.e. aperture of the sclerotized length 44-54 pm (Pl.IV : Figs 16-22 : ring large) (Pl. IV : Fig. 21) ...... 10 vaginal region) 3. - Sclerotization of vaginain lateral view T. prinzifiuus (De Man, 1880) Micoletzky, 1922 appearing as very large quadrangular to - Sclerotized ringnarrower; sclerotization roundedtriangular pieces; t.wo pairs of of vagina different in lateral view (Pl. IV : lateral pores (one pair just anterior tohalf Fig.30) ...... 11 way between base of oesophagus and 11. - Sclerotization of vagina in lateral view vulva, one pair half to one bodywidth appearing as two .small triangular pieces behind the vulva) ; vaginal area rounded (Pl. IV : Fig.24) ...... 12 (elongated when contracteddueto - Sclerot,ization of vaginalarger (Pl. IV : fixation) (Pl. IV : Figs. 2-4) Figs 37-40) ...... 18 T. uariopapillafus Hooper, 1972 12. - Vagina more or less rhomboid with typical - Sclerotization of vaginasmaller ...... 4 interna1 differentiation; onchiostyle length 4. - Onchiostyle lengthunder 50 pm ...... 5 38-48 pm; body length620-800 pm (Pl. IV: - 0nc.hiostyle length over 50 pm . : ...... 7 . Figs 31-33 : vaginalregion) 5.' - Sclerotization of vagina oblique in lateral T. uiruliferus Hooper, 1063 view; onchiostyle length 42-50 pm; vagina - Shape of vaginadifferent ...... 13 with large lobes distally (Pl. IV : Fig. 5) 13. - Vagina barrel-shaped or elongatedwith T. uelatus Hooper, 1972 parallelwalls (Pl. IV : Figs 23, 24) ...... 14 - Sclerotization of vagina roundedtrian- - Vagina taperingtowards the uterus gular in lateral view (Pl. IV : Fig. 6) ; one . (Pl.IV: Figs. 35,36) ...... 17 pair of lateral pores ...... 6 14. - Two pairs of lateral pores ...... 15 6. - Onchiostyle length 35-45 pm;vagina - One pair of lateral pores ...... 16 barrel-shaped, completely surroundedby 15. - Body lengthover 1,120 pm; onchiostyle constrictor muscles (Pl. IV : Figs. 7-9) length 49-70 pm; (Pl. IV: Fig. 23 : vaginal T. cylindricus Hooper, 1962 region) - Onchiostyle length 36-44 pm;vagina T. proximus Allen, 1957 typicallyrounded to rhomboid-shaped, - Bodylength under 1,120 Fm (Pl.IV : towards uterus surrounded by fine cons- Figs. 24-38 + 29 : vaginal region) trictor muscles (Pl. IV : Figs. 6, 10-12) T. sparsus Szczygiel, 1968

~ -~ .- - -_ . T.,similis Seinhorst, 1963 - .. ~~ ....~- T. . aequalis-- Allen,-. 1957

88 Revue Nématol. 3 (1.) : 81-99 (1980) Systetnatics of the Trichodoridae

16. -'Onchiostylelength 39-46 pm;vaginal small difference inshape of the sclerotized sclerotizationdot-like; body length 520- vaginalring (Pl. IV : Figs 38, 39)and in the 1,220 pm (Pl. IV : Fig. 30 : vaginal region) T. pakistanensis Siddiqi, 1962 distancebetween the sclerotized pieces in - Onchiostyle length 73-75 pm;vagina lateral view although 1 cl0 not know if these elongated (Pl. IV : Fig. 34) ; body length differences will be found in a larger number of 570-640 pm specimens and in thetypemmaterial of T. obscurus T. longistylus Yokoo, 1964 (the difference invaginal shape : roundedor 17. - Sclerotization of vagina in lateralview elongated is probably due to fixation). reniform to pyriform; onchiostyle length 53-57 pm; body length650-920 pm (Pl. IV: Fig. 35 : vaginal region) T. borneoensis Hooper, 1962 Genus Paratrichodorus Siddiqi, 1974 - Sclerotization of vagina rounded; onchiostyle length 53-60 Pm; body length 710- 1,100 pm;(Pl. IV : Fig. 36 : vaginal VALIDITYOF SUBGENERA region) T. hooperi Loof, 1973 Siddiqi (1974) distinguished anlongthe species 15. - Sclerotization of vagina in lateral view as of the genus Paratrichodorus threesubpoups rounded equilateral triangles; onchiostyle for which he proposed three subgenera : Para- length 41-44 pm; (Pl. IV : Fig. 37) trichodorus,Atlantadorus and Nanidorus. T. lusitanicus Siddiqi, 1974 - Sclerotization of vagina different; onchio- The basic diagnostic characters in the identi- style length over 50 pm (cf. remarks 2,3).. 19 fication of Paratrichdorus and Atlantadorus 19. - Sclerotization of vaginareniform with are the.posterior ventrosublateral oesophageal longest axisparallel to body Wall in lateral glands overlapping the intestine 'in Paratricko- view; onchiostyle length 55-50 pm; body dorus and an anterior overlap of the intestine length 720-1,250 pm; (Pl.IV : Fig. 35; dorsally anddorsolaterally over the posterior vaginal region) T. californicus Allen, 1957 part of the oesophagus in Atlantadorus (cf. Sid- - Sclerotization of vaginareniform with diqi, 1974; key on p. 270). longestaxis parallel to vaginal Wall in Otherdiagnostic characters inthe identifi- lateral view; onchiostyle length 61-69 pm; body length 720-1,030 pm (Pl. IV: Fig. 90; cation for the males of Paratrichodorus and vaginal region) Atlantadorus are : T. dilatatus Rodriguez-M. & Bell, 1975 - the numberof pairs of postcloacal ventrosub- medianpapillae : one pairin Paratriclmdorus Remarks to the key of Trichodorus fernales and usually two pairs in Atlantadorus (Siddiqi, 1974; Loof, 1975). 1. T. castellanensis Arias, Jimenez & Lopez, - the presence of lateral cervical pores located 1965, T.kurumeensis Yokoo, 1966 and T.obtusus near the level of the excretory pore in Atlanta- Cobb, 1913 cannot be identified (cf. Loof, 1975). 1 dorus, and their usual absence in Paratrichodorus 2. Females of T. dilatatus are morphologi- (Siddiqi,1974). cally very similar to T. californicus (cf. Rodri- In females the subgenera are characterized on guez-M. & Bell, 1978). They have a comparable the base of the vulva-shape : a short longitudinal shape of the vagina but slightly differ in the slit in Paratrichodorus and a pore-like opening shape of thetriangular sclerotization of the in Atlantadorus. vulva : in T. californicus the longestaxis is The subgenus Nanidorus is distinguished from parallel tothe body Wall andin T. dilatatus the othersubgenera mainly bythe posterior the longest axis is parallel to the vaginal Wall. position of the excretory pore near or posterior 3. According to Allen (1957)females of to the base of the oesophagus. The male speci- T.obscurus cannot be distinguished from thoseof mens differ from theother subgenera bythe T. californicus. 1 have studied a few spekimens very long spicules and the presence of only one of T. californicus from Allen's collection and a ventromedianpapilla; the females bythe few specimens of T. obscurus fromBeltsville absence of lateraland caudal pores andby collected by Schindler in 1957. 1 noticeda the shape 'of the vulva : a short transverse slit.

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Discussion it was seen opposite the middle of the posterior oesophageal bulb. In a population of Parafrichodorus (probably The absence of cervicalpores is a character a new species, close to P. feres) specimens were that is also found in the subgenus Paratricho- found with posterior ventro-sublateral oesopha- dorus. gealglands overlapping theintestine either slightly or distinctly, together with an anterior Conclusion overlapping by the intestinedorsally and dorsolaterally over the oesophageal bulb. Both cha- ' Since nlost differences between thethree racters were howeverused as important dia- subgeneraare very small and since 'eventhe gnosticcharacters for distinguishing Paratri- maindiagnostic charact,er may be doubtful 1 chodorus from Atlantadorus; as they have been prefer not to split tip the genus Paratrichodorus found together in the same specimens they are into subgenera. no longer diagnostic. In Nanidorus the posterior ventrosublateral oesophageal glands may over- DEFINITION lap the'intestine; the intestine does not overlap the oekophagus anteriorly. Trichodoridae. Cuticle strongly swollen when Thenumber of pairs of postcloacalventro- treated with acid fixative. Posterior subventral submedian papillaeis not useful in distinguishing oesophageal glands overlap the intestine (except the subgenera (especially Atlantadorus) since of in P. nanrrs) orintestine anteriorly partly the five known species of At1anta.doru.s (P. atlan- overlapsoesophageal bulb or bothtypes of ficus, P. unemones P. grandis, P. pachydermus, overlapoccur. Females without lateral body P. porosus) only P. atlanticm is described with poreswithin one body width from the vulva; twopairs of postcloacalventrosubmedian pa- vaginal sclerotization wealdy developed ; sper- pillae. The other species of Atlantadorus possess mathecae present or absent. Males with caudal

only one pair as in Paratrichodorus and Nanido- alae; copulatory muscles not extending anterior , rus. to bursa, ride taiI straight in death. Suspensor The pair of lateral cervical pores in Afla.nta- muscles inconspicuous. Two supplements within dorus is notalways located near the level of reach of caudalalae, well separatedfrom the the excretory pore. 1 observed in P. pachyder- anterior third whichis often reduced or only one rnzzs and P. anenzones the pair of lateral cervical supplement present. One or two pairs of large pores at some distance anterior to the ventro- ventrosubmedianpostcloacal papillae. Usely median cervical pore. In P. grandis and P. poro- one ornoventromedian cervical papillae, sus it lies atthe level of theventromedian rarely three. cervic,al pore. In Nanidorus cervkal poresare absent. In Paratriclzodorrzs lateral cervical pores TYPE SPECIES were not described.Siddiqi (1974) considered 1 the pair of lateral cervical pores as very rarely Paratrichodorustunisiensis (Siddiqi,1963) presentin the subgenus Paratrichodorus. Siddiqi,1974 The difference in vulva-shapebetween the subgenerais very difficult tointerpret. The Syn. Trichodorustunisiensis Siddiqi, 1963 vulva opening is always very small in trichodo- rids. In lateral view the vulva shape may be KEY TO Paratrichodorus MALES (cf.remark 1) difflcult to determinee.g. in P. anemones the funnelshaped vulva (PI. VI1 : Fig. 13) may be 1. - Males with a single ventromediansup- plement (Pl. V : Fig. 1); no ventromedian considered as a large pore ; it is however, difflcult c.ervica1 pores; posterior subventral oeso- to distinguish it from a short longitudinal slit phagealglands usually overlapping the asis found in thesubgenus Paratrichodorus. intestine;intestine anteriorly not overlapping oesophageal bulb (Pl. V : Fig. 3) 2 ln Nanidorus theexcretory pore is ' not - Males with two or threeventromedian alwaysnear or posterior tothe base ,of the supplements; ventromedian cervical pores oesophagus.. In some. specimens 0f.P.(N.) minor .. ~. - present .or absent ._..~. . ._...... _.... ._...... 3

90 ReuLze Nématol.(1980) 3 (1) : 81-99 1 c

10 8

l i \\

w Pl. V. Paratrichodorus males(scale = 20 Pm). P. nanus (afterSiddiqi, 1963). 1 : tail + copulatoryapparatus. 2 : anterior body region. P.. minor (after Siddiqi, 1963). 3 : oesophageal region; 4 : tail + copulatory apparatus. 5 : P. tunisiensis : oesophageal region (after Siddiqi, 1963). 6 : P. pachydermus, posterior part of oesophageal region 7 : Paratrichodorus cf. teres, posterior part of oesophageal region. 8 : P. acaudatus (after Sidiqi, 1960) tail + copulatory apparatus. 9 : P. mirzai (after Siddiqi, 1960). 10 : P. teres (after Kuiper & Loof, 1961). 11 : P. Zobatus (after Colbran, 1965). 12 : P. tunisiensis (after Siddiqi, 1963). 13 : P. alleni (after Andrhssy, 1968). 14 : P. allius (after Jensen, 1963). W. Decraemer

92 Systematics of the Trichodoridae

7. - Testis degenerate, without sperm; onchios- P. fansaniensis Siddiqi,1974; P. westindicus tylelength 47-52 Pm; spide length Rodriguez-M., Sher & Siddiqi, 1978 and P. obesus 45-52 pm (remark 2) (Pl. V : Fig. 10 : tail region + copulatoryapparatus) Razjivin & Penton, 1975. P. teres (Hooper, 1962) Siddiqi, 1974 2. Males of P. , teres (Syn. P. flevensis - Testiswith well developed sperm...... 8 Kuiper & Loof, 1962) can show a body length 8. - Spicule length over 50 pm smaller Lhan. 3QQ..pm,this being in contradiction P. Zobatus (Colbran, 1965) Siddiqi, 1974 with the key of Loof (1975). Spicule length 50 pm or less...... 9 Distance second supplementanusto about 45 % of spiculumlength; spicule length 50 pm (Pl.V : Fig.13) KEY TO Paratrichodorus FEMALES P. alleni (Andrhssy, 1968) Siddiqi, 1974 Distance second supplement to anus about 1. - Intestineanteriorly partly overlapping 90 of spiculum length ...... 10 posterior oesophageal bulb dorsally and Spicule length 31 pm; onchiostyle length dorsolaterally,posterior subventral oeso- 42 pm (Pl. V : Fig. 14 : tail region + copu- phageal glands not overlapping the intes- latoryapparatus) tine(Pl. VI1 : Fig. 1) ...... P. allius (Jensen,1963) Siddiqi, 1974 - Intestineanteriorly not overlapping the Spicule length 42-44 pm;onchiostyle oesophagus; posteriorsubventral oeso- length 39-41 pm (Pl. VI: Fig. 1 : tail region phagealglands usually clearly, or rarely + copulatory apparatus) slightly,overlapping thé intestine (Pl. P. rhodesiensis (Siddiqi & Brown, 1965) Siddiqi, 1974 VI1 : Fig. 3) ...... ).. 11. - Two ventromedianpreanal supplements, - Intestine anteriorly overlapping posterior both within spicular range; spicule length oesophageal bulb and posterior subventral 36-39 pm; onchiostyle length 43-48 pm oesophageal glands overlapping the intes- (Pl. VI : Fig. 2 : tail region + copulatory tine (Pl. VI1 : Fig. 8); onchiostyle length apparatus) 50-60 Pm; one pair of lateral pores (5.5-8.2 P. porosus (Allen,1957) Siddiqi, 1974 timesthe body width posterior to the - Three ventromedian preanal supplements, vulva) anterior supplement well anterior to head Paratrichodorus sp.(cf. teres\ of retracted spicules ...... 12 2. - Spermathecapresent ...... 3 12. - Onchiostyle length more than 60 pm. .... 13 - Spermathecaabsent ...... 4 - Onchiostylelength under 60 pm...... 14 3. - One pair of subventral body pores behind 13. - Spicule length 79-87 pm; spicules straight the vulva (Pl. VI1 : Fig. 12); onchiostyle with roundedoffset capitulum; onchiostyle length 64-70 Pm; body length 760-1130 pm length 68-82 pm (Pl. VI: Fig. 3 :tail region P. atlanticus (Allen,1957) Siddiqi, 1974 + copulatory apparatus) - Lateral body pores anterior and posterior P. atlanticus (Allen,1957) Siddiqi, 1974 to vulva0-2 on each body side; onchiostyle - Spicule length 39-43 pm; spicules curved, length 42-53 pm; body length 650-970 pm capitulumnot offset;onchiostyle length (Pl. VI1 : Fig. 13 : vaginal region) 64-72 pm (Pl. VI : Fig. 4 : tail region + P. anemones (Loof, 1965) Siddiqi, 1974 copulatory apparatus) 4. - Onchiostyle length 65-72 pm; body length P. grandis Rodriguez-M. & Bell, 1978 900-1,040 pm.. (Pl. VI1 : Fig. 17 : vaginal 14: - Tail short, round; spicules fine, 46-53 pm region) long (Pl. VI: Fig. 6); distance second prea- P. granais Rodriguez-M. & Bell, 1978 na1 supplementto anus 38-64 % of spi- - Qnchiostylelength under 65 pm...... 5 culum length Ventromedian pores nearvulva present P. anemones (Loof,1965) Siddiqi, 1974 5. - - (2pairs) (Pl. VI1 : Fig. 14); onchiostyle Tail longer, narrower (trapezoid); spicules length 39-47 pm; body length 420-570 pm broaderwith different shape, 40-45 Pm P. porosus (Allen,1957) Siddiqi, 1974 long(Pl. VI : Fig. 5); distance second - No ventromedian body pores near vulva; preanalsupplement to anus 75-94 y. OC lateral body pores posterior to vulva 2-5 spicule length on each bodyside; onchiostyle' length P. pachydermus (Seinhorst, 1954) Siddiqi, 1974 44-54 pm;body length 690-1,020 pm (Pl.VI1 : Fig.2 : partgenital system; Rernarks io the key of Paratrichodorus males Fig. 1 : posteriorend of oesophagus) P. pachydermus (Seinhorst, 1954) Siddiqi, 1974 1. Males areunlmown in P. acutus (Bird, 6. - Caudal pores present (Pl. VI1 : Fig. 4) .... 7 1967)Siddiqi, 1974; P. renifer Siddiqi, 1974; - Caudal pores absent (Pl. VI1 : Fig. 5) .... 14

Revue. Nhzatol. 3 (1) : 81-99 (1980) 93 W. Decraemer

f 7. - Onchiostyle length over 80 pm;body 15. - Tail acute (Pl. VI11 : Fig. 2); onchiostyle length 910-950 pm sclerotized vaginal length 22-26 pm; body length 375-610 Fm ring2 pm thick (Pl. VI1 : Fig. 6) P. acutus (Bird, 1967) Siddiqi, 1974 P. acaudatus (Siddiqi, 1960) Siddiqi, 1974 - Tailhemispherical ...... 16 -- Onchiostyle lengthunder 80pm ...... 8 16. - Posteriorsubventral oesophageal glands S. - Spermathecapresent (hwo vcrysimilar more or less overlappingthe intestine species; see remark 3) ...... 9 (slightly or clearly)(Pl. VI11 : Fig. 4)17 - No spermatheca ...... 10 - Non-overlapping base cf oesophagus. .... 18 9. - Oesophageal overlap one body width long 17. -. Excretorypore oppositeoesophageal (Pl. VI1 : Fig. 3); one lateral body pore base (Pl. VI1 : Fig. 4) ; onchiostyle lengt,h postvulvar; onchiost.yle length 47-52 pm; 27-42 pm (mean 32 pm);body length body length 630-1,035 pm (Pl. VI1: Fig. 7 : 440-1,530 pm (mean 650 pm)(Pl. VI11 : vaginal region) Fig.3 : vaginal region) P. lobatus (Colbran, 1965) Siddiqi, 1974 P. minor (Colbran, 1956) Siddiqi, 1974 - Oesophageal overlap shorter thanone body . - Excretory pore more anteriorly situated; width; two lateral body pores; onchiostyle onchiostylelength 30-34.5 pm; . body length 48-51 pm;body length 900- length 475-500 pm 1,100 pm (Pl. VI1: Fig. 10 : vaginal region) P. obesus (Razjivin & Pentan,1975) P. tunisiensis (Siddiqi, 1963) Siddiqi, 1974 Rodriguez-M. & Bell, 1978 10. - Onchiostyle lengthunder 35 Pm;lateral 18. - Excretory pore opposite anterior part of body ppres absent;body length 490- intestine (Pl. VI11 : Fig. 5); onchiostyle

610 pm !Pl. VI1 : Fig. 11 : vaginal region) 1e.ngt.h , 21-23 Fm;body length 490- P.' mirzai (Siddiqi, 1960) Siddiqi, 1974 600 pm; sclerotization of vagina in lateral - OnChiostyle length over 35 pm ...... 11 viewsmall, inconspicuous (Pl. VI11 : 11. - Lateralbody pores present; onchiostyle Fig. 6) length 41-52 pm;body length 770- P. 'nanus (Allen, 1957) Siddiqi, 1974 1,230 pm; uteri without sperm (Pl. VI1 : - Excretory pore 0pposit.e base of oesopha- Fig. 15;vaginal region) gus; onchiostylelength 31-34 pm;body P. teres (Hooper, 1962) Siddiqi, 1974 length 420-560 pm; sc,lerotization of - Lateralbody pores absent ...... 12 vaginainlateral view kidney-shaped (Pl. VI11 : Fig.7) 12. - Vaginal sclerotizationappearing as two P. renifer Siddiqi, 1974 bold dot,s in lateral view (Pl. VI1: Fig. 18) P. rhodesiensis (Siddiqi & Brown, 1965) Siddiqi,'1974 - Vaginal sclerotizationsmaller in lateral Remarks to the key of Paratrichodorus females view ...... 13 13. - Body length 640-780 pm;onchiostyle 1. P. roestindicus has.a doubtful position length 37-42 pm (Pl. VIT : Fig. 16 : vaginal (cf. Rodriguez-M., Sher & Siddiqi,1978). region) 2. Females are unknown in P. alleni (An- P. alZius (Jensen, 1963) Siddiqi, 1974 - Body length 470-650 pm;onchiostyle drassy,1968) Siddiqi, 1974. length 35-40 pm (Pl. VI1: Fig. 19 : vaginal 3. P. lobatus (Colbran,1965) Siddiqi, 1974 rcgion) and P. trznisiensis (Siddiqi,1963) Siddiqi, 1974 P. tansaniensis Siddiqi, 1974 stronglyresemble eac,h other. P. 1oba.tu.s only 14. - Genital systcmmono-prodelphic (Pl. VI11 : differs from P. Sunisiensis bythe length of Fig. 1); vulva at 60-65 % (cf. remark 1) P. westindicus Rodriguez-M, Sher & Siddiqi, 1978 the oesophagealoverlap, i.e. one bodywidth - Genital systemdidelphic ...... 15 long in P. lobatus andsllorter than one body

Pl. VII. Paratrichodorus females (scalc = 20 pm). 1 : P. pachydermus : posterior part of oesophageal region. 2 : P. pachyderrnus : reproductive system. 3 : P. lobalus: posterior part ocsophagus (after Siddiqi, 1974). 4 : P. mirzai, tail (after Siddiqi, 1960). 5 : P. renifer : tail (after Siddiqi, 1974). 6 : P. acaudatus : vaginal region (after Siddiqi, 1960). 7 : P. lobatus :vaginal region (after Colbran, 1965). 8 : Paratrichodorus sp. (cf. teres) : posterior part oesophageal region. 9 : P. tunisiensis : posterior part oesophagus (after Siddiqi, 1963). 10 : P. tunisiensis: vaginal region (after Siddiqi, 1963). 11 : P. mirzai : vaginal region (after Siddiqi, 1960). 12 : P. atlanticus : vulva, ventral view (after Allen, 1957). 13 : P. anemones : vaginal region (specimen, courtesy P.A.A. Loof). 14 : P. porosus : vaginal region (after Allen, 1957). 15 : P. teres : vaginal region (after Hooper, 1962). 16 : P. allius : vaginal region (after Jensen, 1963).17 : P. grandis : vaginal region (after Rodriguez-M. & Bell, 1978). 18 : P. rhodesiensis : vaginal ...... region (after Cicldiqi&-Brown, 1965). 19; P. tansaniensisi vaginal region. (after. Siddiqi,_. .- 1974). ,. . ..

94 Revue Ndmafol. 3 (1) : 81-99 (1980) Systerrzatics of the Trichodoridae

10 Pm 17

Revue Nérnatol. 3 (1): 82-99 (2980) 95 W. Decraemer

another, especially due to t6e variable position Pl. VIII. Paratrichodorus females (scale = 20 Fm). Of the gland The gland 1 : P. westindicus : reproductivesystem(after Rodri- nucleus is aslarge as or usuallysomewhat guez-M., Sher & Siddiqi 1978).2 : P. acutus: tail (after larger than the nuclei of the posterior ventrola- Bird, 1967). 3 : P. minor : vaginal region (after Allen, teral oesophageal glands;its positionvaries 1957). 4 : P. minor : anteriorbody region (after Siddiqi, 1974). 5 : P. nanus : anteriorbody region ' from the level Of the anterior ventrolateral ' (afterSiddiqi, 1974). 6 : p. nanus : vaginal region Oesophageal gland nudei to the level Of the (after Siddiqi, 1963).7 : P. renifer : vaginal region - .~ pQsterior venJrolatera1 oesophageal.gland n9:Je-i. - -(after Slddiqi, 1974). - - ---

96 Revue Nématol. 3 (1) : 81-99 (1980) Syste~naticsof the Trichodoridae

DEFINITION KEY TO Monofrichodorus FEMALES, basedon Rodriguez-M., Sher & Siddiqi(1978) Trichodoridae. Cuticle not swollen upon fixation.Posterior ventrosublateral oesophageal - Paired lateral cervical pores present immediately glandsnot overlapping the intestine. Female behind the amphid opening M.vangundyi Rodriguez-M., Sher & Siddiqi, 1978 reproductive system monodelphic, prodelphic ; - Paired lateral cervicalpores absent with flexure at the oviduct; postvulvar uterine M. monolzystera (Allen, 1957) Andràssy, 1976 sacpresent. Vagina anteriorlydirected, more than one-half body width, with strong muscula- Remark ture;vaginal sclerotization well developed. Lateral advulvar body pores present.Vulva a Both species are also very similar in females; transverse slit at about 80 y0 of body length. they possess a comparable shape of the vagina Males with well developed diagonal copulatory andvaginal sclerotization. As faras 1 Bnow muscles.Spicules long, slender,setose. Three theobservation of lateral cervicalpores in ventromedian preanal supplements, at least one females of M. vangundyi Rodriguez-M., Sher Ist anteriorto retracted spicule.Paired ventrola- Siddiqi,1978 isunique and until now only ' teral p,ostcloacal papillae present. Single ventral lmown in males (however not found in such a cervicalpapilla anterior toexcretory pore. far anterior position). Inboth sexesasingle caudal pore present.

TYPE SPECIES 'Genus Allotrichodorus Rodriguez-M., Sher & Siddiqi, 1978 Monotrichodorusmonoliystera (Allen,1957) AndrBssy,1976 DEFINITION Syn. Trichodorusmonohystera Allen,1957 Trichodoridae. Cuticle swollen upon fixation. Other species Intestineanteriorly overlapping oesophageal Monotrichodorusvangundyi Rodriguez-M., bulb dorsally and dorsolaterally. Female repro- Sher & Siddiqi,1978. ductivesystem monodelphic, prodelphic with flexure atoviduct; postvulvar uterine sac KEY TO Monotrichodorus MALES, basedon present. Spermatheca conspicuous. Vagina more Rodriguez-M., Sher & Siddiqi, (1978). than half bodywidth in length with strong musculature; vaginal sclerotizationconspicuous. - Two preanal supplements within the range of the Vulva a transverse slit at about 85 %. In males retracted spicules caudalalae present. Ventral cervical papilla, M. vangundyi Rodriguez-M., Sher & Siddiqi, 1978 - One preanalsupplement within the range of the if present, anterior to excretorypore. Spicules retracted spicules transversely striated, elongate, slender, setose ; M. ~nonol~ystera(Allen, 1957) Andràssy, 1976 capitularextension having a dorsal groove or depressionleading intoproximal opening of Remark spicule.Three ventromedian supplementary papillaewithin range of retractedspicules. Both species are very silnilar (synonymous?); Large paired lateroventral postcloacal papillae the only differencelies in the position of the present.Caudal pores present. second preanal supplement, i.e. within the range of the retracted spicules or not. This difference is ratlzer mail to separateboth species using TYPE SPECIES males, since in theoriginal description of M. mo- nolzyslera Allen (1957) the second supplement is Allotrichodoruscampanullatus Rodriguez-M., described atabout the level of theproximal Sher & Siddiqi,1978 end of the spicules (see also Fig. 1C in Rodriguez- Other species A. guttatus Rodriguez-M., Sher M., Sher & Siddiqi,1978). & Siddiqj, 1978

Revue Nénzatol. 3 (1) : 81-99 (1.980) 97 W. Decraemer

KEY TO Allotriclzodorus MALES CLARK, W. C. (1963). A new species of Trichodorus (Nematoda : Enoplida)from Westland, ’ New - One vent>romediancervical papilla; onchiostyle Zealand. N. Z. JI. Sci., 6 : 414-417. length 45-57 Fm; spicule length 46-59 ym; body length COBB, N.A. (1913). New nematodegenerà found 540-690 ym inhabitingfresh water and non-brackish soils. A. campanullatus Rodriguez-M., Sher & Siddiqi, 1978 J. Wash.Acad. Sci., 3 : 432-444. - No vent.romedian cervicalpapilla; onchiostyle length 65-72 Pm; spicule length 65-76 ym; body length COBB, N. A. (1920). One hundred new nemas. Contrib. 670-740 ym Sci.Nematol., 9 : 217-343. A. guttatus Rodriguez-M., Sher & Siddiqi, 1978 COLBRAN,R. C. (1956).Studies of plantand soil nematodes 1. Two new species from Queensland. Qd. J. agric. Sci., 13 : 123-126. KEY TO Allotrichodorus FEMALES, based on COLBRAN, R. C. (1965).Studies of plantand soil Rodriguez-M., Sher & Siddiqi (1978) nematodes 9. Trichodorus lobatus n. sp. (Nematoda : - Onchiostyle length 46-55 ym; body 1engt.h 630-680 Trichodoridae), a stubby-root nematode associated Pm; vaginal sclerotization appears as two small bells withcitrus and peachtrees. Qd. J. agric.anim. in lateral view Sci., 22 : 273-276. A. campanullatus Rodriguez-M., Sher & Siddiqi, 1978 COOMANS,A. & LOOF,P. A. A. (1970).Morphology - Onchiostyle length 63-71 ym; body length 650-720 andtaxonomy of Bathyodontina(Dorylaimida). Pm; vaginal sclerotization appears as two rather un- Nematologica, 16 : 180-196. equaldrops in lateral view ESSER,R. P. (1971).A compendium of thegenus A. guttatus Rodriguez-M., She,r & Siddiqi, 1978 Trichodorus (Dorylaimoidea : Diphtherophoridae). Proc. Soil Crop Sci.Soc. Fla., 31 : 244-253. ACKNOWLEDGEMENTS GOODEY,T. (1951). Soiland freshwater nematodes. London,Methuen, 390 p. 1 thank Prof. Dr A. Coomans and Mr D. De Waele HOOPER,D. J. (1962).Three new species of Tricho- for reading the manuscript. 1 am also very grateful .to dorus (Nematoda : Dorylaimoidea) and observations Dr E. Geraert for the fruitful discussions and for rea- on T. minor Colbran, 1956. Nematologica, 7 : 273-280. ding of the manuscript. HOOPER,D. J. (1963). Trichodorusuiruliferus n. sp. (Nematoda : Dorylaimida). Nematologica, 9 : 100- 204. REFERENCES HOOPER,D. J. (1972). Two new species of Trichodorus (Nematoda : , Dorylaimida) from England. Nema- ALLEN,M. W.(1957). A review of thenematode tologica, 18 : 59-65. genus Trichodorus withdescriptions of ten new HOOPER,D. J. (1973). Nematodes. In : Gibbs, A. J. species. Nematologica, 2 : 32-62. (Ed.) Viruses and invertebrates, Amsterdam, Nether- ,ALLEN,M. W. (1957).Supplement : Dorylaimoidea lands,North Holland Publishing Co : 212-228. dexribed since the monographs on Dorylaims and HOOPER, D.J., KUIPER,K. & LOOF, P. A. A. (1964). Dorylaimoidea were publishedin 1936 and 1939. Observationson the identity of Trichodorus teres In : Thorne, G. (Ed.) A monographof the nematodes Hooper, 1962 and T. fleuensis Kiuper & Loof, 1962. of thesuperfamily Dorylaimoidea, 2nd edition. Nematologica, 9 : 646. Capitaroologica, 8 : 262-263. JENSEN,H. J. (1963). Trichodorus allius, a new species ANDRASSY,1. (1968).The scientific results of the of stubby-rootnematode from Oregon (Nemata : Hungarian soil zoological expedition to the Brazza- Dorylaimoidea). Proc.helnzinth. Soc. Wash., 30 : ville-Congo. 31. Nematodenaus Grundwasser. 157-159. Ann. Uniu.Sci. Budapest, 9-10 : 3-26. KUIPER,K. & LOOF,P. A. A. (1962). Trichodorus ANDRASSY,1. (1976). Evolutionas a basis for the flevensis n. sp. (Nematoda : Enoplida),aplant systematization of nematodes. London,San Fran- nematodefrom a newpolder soil. Versl.Meded. cisco & Melbourne, PitmanPublishing, 288 p. PlZiektenh-.Dienst Wageningen, 136 : 193-200. ARIAS-DELGADO,M., JIMEPI’EZ-MILLAN,F. & LOPEZ- LOOF,P. A. A. (1964). Free-living and plant parazitic PEDREGAL,J. M. (1965). Tres nuevas especies de nematodesfrom Venezuela. Nematologica, 10 : nematodos posibles fitoparasitos en suelos espanoles. 201-300. Publnes Inst. Biol.apl. Barcelona, 38 : 47-58. LOOF,P. A. A. (1965). Trichodorusanemones n. sp. BIRD, G. W. (1967). Trichodorus acutus n. sp. (Nema- with a note on T. teres Hooper, 1962 (Nematoda : todea : Diphtherophoroidea)and a discussion of Enoplida). Versl.Meded. PlZiektenk. Dienst Wage- allometry. Can. J. Zool., 45 : 1201-1204. ningen, 142 : 132-136. CLARK, W. C. (1961). A revisedclassification of the LOOF,P. A. A. (1973). Taxonomybf Trichodorus aequa- orderEnoplida (Nematoda). N. 2. J1. Sci., 4 : lis-complex (Dipht.herophorina). Nematologica, 19 : . . . 1_2’3-1_5_0..- - -.- - - . . 49-6f?. - . --- .. -- 98 Revue Ndmntol. 3 (1) : 81-99 (1980) Systematics of the Trichodoridae

LOOF, P. A. A. (1975).Taxonomy of Trichodoridae. SIDDIQI,M. R. (1960). Two new species of the genus In : Lamberti, F., Taylor, C. E. & Seinhorst, Trichodorus (Nematoda : Dorylaimoidea) from In- J. W. (Eds). Nematode vectors of plant viruses. dia. Proc.helminth. Soc. Wash., 27 : 22227. Londonand New York,Plenum Press : 103-127. SIDDIQI, M. (1962). Trichodoruspakistanensis LORDELLO,L. G. E. & ZAMITH, A. P. L. (1958). R. n.sp. (Nematoda : Trichodoridae)with observa- Notasôbre O gênero Trichodorus Cobb, 1913 com descriçao de Trichodorusbucrius n. sp. (Nematoda, tions on T. porosus Allen, 1957, T. mirzai Siddiqi, Dorylaimoidea). AnaisAcad. bras.Cienc., 30 : 1960 and T. minor Colbran, 1956, fromIndia. 103-105. Nem.atologica, 8 : 193-200. MAMIYA, Y. (1967). Descriptive notes on three species SIDDIQI,M. R.(1963). Trichodorus spp.(Nematoda : of Trichodorus (Dorylaimida : Trichodoridae)from Trichodoridae) from Tunisia and Nicaragua. Nema- forestnurseries in Japan. Appl. En€. Zool., 2 : tologica, 9 : 69-75. 61-68. SIDDIQI,M. R. (1974). Systematics of the genus Tri- DE M~N,' J. G. (1880). Die einheimischen, frei in der chodorus Cobb, , 1913 (Nematoda : Dorylaimida), reinen Erde und im siissen Wasser lebenden Nema- withdescriptions of three new species. Nemafolo- : toden. Tijdschr.ned. dierk. Ver., 5 1-104. gica, 19 : 259-278. DE MAN, J. G. (1884). Die frei in der reinen Erde und im siissenWasser lebenden Nematoden der nieder- SIDDIQI,M. R. & BROWN,K. F. (1965). Trichodorus larzdischen Fauna. Leiden,Brill, 206 p. rhodesiensis and Amphidelustrichurus, twonew MICOLETZKY,H. (1922). Die freilebenden Erd-Nema- nematode species fromcultivated soils of Africa. toden. Arch.Naturgesch., A 87 : 1-650. Proc.helminth. Soc. Wash., 32 : 239-242. RAZJIVIN,A. A. & PENTON,G. (1975). A new species SZCZYGIEL,A. (1968). Trichodorus 'sparsus sp. n. t of thegenus Trichodorus (Nematoda)from the (Nematoda,Trichodoridae). Bull. Acad. pol. Sci. rhizosphere of the sugar canein Cuba. 2001.Zh., Cl. II Sér. Sci. biol., 16 : 695-698. 54 : 1082-1083. THORNE,G. (1935). Notes on free-living andplant- RODRIGUEZ-M,R. & BELL,A. H. (1978). Three new parasitic nematodes, II. Proc. helminth. Soc. Wash., : species of Trichodoridae(Nematoda Diphthero- 2 : 96-98. phorina) with observations on the vulva in Para- trichodorus Siddiqi, 1973. J. Nematol., 10 : 132-141. TIXORNE,G. (1939). A monograph of the nematodes of RODRIGUEZ-M,R., SHER, S. A. & SIIYDIQI,M. R. thesuperfamily Dorylaimoidea. Capitazoologica, (1978).Systematics of the monodelphic species of 8 : 1-261. Trichodoridae (Nematoda : Diphtherophorina) with YEATES,G. W. (1967).Studies on nematodesfrom descriptions of a new genus and four new species. dunesands 4. Diphtherophoroidea. N. 2. J1. Sci., J. Nematol., 10 : 141-152. 10 : 322-329. SEINHORST,J. W. (1954). On Trichodoruspachyder- mus n. sp.(Nematoda : Enoplida). J. Helminth., YOKOO,T. (1964). On thestubby root nematodes 28 : 111-114. from the Western Japan., Agric. Bull. Saga Univ., 20 : 57-62. SEINI-IORST,J. W. (1963). A redescription of the male of Trichodorus prirnitivus (de Man), and the descrip- YOKOO,T. (1966). On a new stubbyroot nematode tion of a newspecies X. similis.Nematologica, (Trichodoruskurumeensis n. sp.) fromKyushu, 9 : 125-130. Japan. Agric. Bull. SagaUniv., 23 : 1-6.

Accepté pour publication le 10 octobre 1979.

Revue Némafol.3 (1) :81-99 (1980) 99