Revision of the Genus Paratylenchus Micoletzky, 1922 and Descriptions of New Species

JOURNAL OF NEMATOLOGY VOLUME 8 APRIL 1976 NUMBER 2

Revision of the Genus Paratylenchus Micoletzky, 1922 and Descriptions of New Species. Part III of Three parts--Gracilacus;

D. J. RASKI ~

Abstract: Part 1II covers species with female stylet length ~41 #m which are cottsidered by this author to coutprise the genus C;racilacus Raski, 1962. Seven new species of (;racilacus are descrihed and further observations given on 14 other species. ParatyleTwhtts slrenzkei (Volz, 1951) Oostenbrink, 19f~} is transferred to species i~qttirendae. A key to the species of Gracilacus is in- cluded. Key Word: taxonomy.

This report completes the present re- it was accepted by Allen and Sher (1) and vision of the genus Paratylenchus and the by Geraert (6), and rejected by Thorne related genus Gracilacus. It includes those and Malek (12) and by Golden (7). The species in which the females bear a stylet six new species with long stylets (described Longer than 41 ttm. since 1962) and tlle seven species (described The genus Gracilacus Raski, 1962 (10) in this paper) contribnte some new infor- was proposed for those species of Paraty- mation which supports the concept of this lenchus previously described as ltaving a group of nematodes as representing a sep- female stylet greater than 48 t~m and for arate and distinct taxon. five new species. Other characters used to The species described by Colbran (3) distinguish Gracilacus from Paratylenchus as Paratylenchus mutabilis indicates a closer were: (i) swollen females found in some relationship between these long-stylet spe- species, (ii) most juveniles with elongate cies and Cacopaurus pestis Thorne, 1943 stylet, and (iii) excretory pore mostly in because of the cuticular ornamentation on region of metacorpus (opposite isthmus in swollen females of P. mutabilis. On the some species). Gracilacus was distinguished other hand, some species judged to belong from the genus Cacopaurus Thorne, 1943 to Gracilacus are found to have what ap- by the elongation of its body posterior to pears to be fourth-stage juveniles with very vulva (tail very short and blunt in Caco- reduced stylets, a characteristic most fre- paurus) and a finely annulated cuticle with- quently found in species of Paratylenchus out ornamentation. with female stylets 38/zm or less. This char- This concept was not accepted by Sid- acteristic indicates relationship with Paraty- diqi and Goodey (l l) who synonymized lenchus. Gracilacus with Paratylenchus. That action of synonymy has been variously received; It is my opinion that these long stylet species are more closely related to Caco- paurus than to Paratylenchus. Gracilacus differs in tail shape from the short, bhmt Received for publication 1 July 1975. 'The kind cooperation of many hematologists, including tail of C. pestis. In addition, senile, swollen I. Andr~.ssy, M. W. Brzeski, R. C. Colbran, A. l)e(;risse, females of C. pestis change to brownish, J. C. Edward, R. P. Esser, E. Geraert, A. M. Golden, D. J. Hooper, W. R. Jenkins, P. A. A. Loof, W. F. Mai, empty cuticular remnants which persist for G. Merny, R. H. Mnlvey, B. W. Ngtmdo, J. W. Seinhorst. long periods in the soil. Such forms have A. R. Seshadri, S. A. Sher, (;. Swarnp, A. C. Tarjan, G. Thorne and W. M. Wouts, who sent type specimens. not yet been reported for any species of identified material, and new collections, is gratefully ac- Gracilacus. So far, none of the species of knowledged. The technical assistance of N. Jones, U. K. Mehta, H. Chang, and J. Pinochet, especially in mounting Paratylenchus with stylet 38 /~m or less are and measuring specimens, was most valuable in this study. known to form swollen females as found Professor of Nematology, Department of Nemalolog2/', Uni- versity of California, Davis 95616. in Gracilacus and Cacopaurus.

The JOURNAL OF NEMATOI.OGY for January (8: 1-96) was issued 15 January 1976. 97 98 Journal of Nematology, Volume 8, No. 2, April 1976 Therefore, it is concluded that Gracil- syn. Paratylenchus micoletzkyi acus should be recognized as a separate Edward, Misra and Singh, 1967 genus. Gracilacus robusta MATERIALS AND METHODS (Wu, 1974) n. comb. syn. Paratylenchus robustus The study was based on more than 550 slides holding approximately 2,100 speci- Wu, 1974 mens. Most o1: the specimens examined were Endings of specific names are changed mounted in glycerin, but the precise meth- throughout to conform with feminine ods of killing, fixing, and processing to de- gender of Gracilaeus. hydrated glycerin are not known. After specimens were collected, they were sep- Gracilacus aonli arated from soil by a combined gravity- (Misra and Edward, 1971) n. comb. screening and mist-extraction technique, syn. Paratylenctms aonli killed by gentle heat, and fixed in 5% Misra and Edward, 1971 formaldehyde. The specimens then were Type specimens have not been accessible passed through F.A.A., 2.5% glycerin in and judgment of G. aonli is based on the 30% alcohol, and 5% glycerin in g0% pnblished description. The authors (8) con- alcohol. They were allowed to dehydrate sider G. aonli to be most closely related to from the last solution to glycerin and then G. goodeyi and G. straeleni. The head of were mounted in dehydrated glycerin. G. aonli is described as "truncate conoid," Gracilacus Raski, 1962 hut in the illustrations it appears to be more rounded in outline. G. aonli also is Diagnosis (emended): Paratylenchidae. descrihed as having oval spermatheca, but Individuals small, less than 0.50 mm. Most there are no detailed drawings indicating juveniles with elongate stylet but some spe- its precise structure. In some species, the cies with reduced stylet in fourth-stage spermatheca is oval and is about the same juvenile. Female slender to swollen, with a in length as in width, whereas the elongate, stylet 41-119 ~m in length. Body elongate ovoid spermatheca of G. goodeyi and of G. posterior to vulva. Cuticle finely annulated, pandata is very long, 2 to 3 times as long without ornamentation (except G. mu- as the corresponding body diam. The tabilis). Excretory pore generally in region shorter stylet (44-56 /~m) and conical head of metacorpus near valve or further an- of G. goodeyi appear to be distinctive from terior, but may be near nerve ring. Male G. aonli (stylet= 55-65 ~m). slender, active; styler absent. Caudal alae The distinction of G. aonli from G. lacking or at most represented by slight, stroeleni seems to be based on the spherical thickened, cuticular evaginations. Ovary spermatheca of G. slraeIeni as contrasted single. Testis one. with the oval spermatheca of G. aonli. Also Type species: Gracitacus epacris the esophagous is shorter in G. straeleni (Allen and Jensen, 1950) (b = 3.0-4.5 vs. 2.5-3.0 in G. aonli) and the Raski, 1962 tail more rounded in G. aonli (finely syn. Cacopaurus epacris rounded to acute in G. straeleni). Allen and Jensen, 1950 Paratylenchus epacris Gracilacus abietis (Allen and Jensen, 1950) (Eroshenko, 1974) n. comb. Siddiqi and Goodey, 1963 syn. Paratylenchus abietis Eroshenko, 1974 Olher changes: Type specimens have not been accessible Gracilacus crenata (Corbett, 1966) so judgment is based on the published de- n. comb. scription. The evidence indicates the syn. Paratylenchus crenatus Cor- author's diagnosis of relationship with G. bett, 1966 straeleni is correct (5). The characteristics Gracilacus micoletzkyi used to distinguish G. abietis from G. (Edward, Misra and Singh, 1967) n. comb. straeleni are not entirely convincing. The Revision of Paratylenchus Part III--Gracilacus: Raski 99 smooth head of G. abietis may be a de- rounded than those of most females seen pendable character (as it is in Paraty- in other collections. In all other respects, lenchus profectus) although it is not com- these specimens fit G. acicula. mon in these genera. Other species have been considered to have the head smooth Gracilacus acuIenta (Brown, 1959) or without amndation, but occasionally Raski, 1962 fine head annulation is apparent. Tile cor- Two slides with 35 female and 4 male pus, described and illustrated as being nar- paratypes were loaned by the Canadian row in the anterior half and then swelling National Collection of Nematodes for this to nearly twice that diameter in posterior study. These are not from the type locality half, also seems distinctive but has not been but from grass sod in Gatineau Park, Que- noted or used heretofore. The position of bec, which was reported by Brown (2) as hemizonid relative to the excretory pore is one of the distribution records of G. not dependable as a specific character in aculenta. These specimens conform closely other species. It has been found to vary with the original description. from anterior to posterior in various speci- Five new collections, identified as G. mens of the same population. The range aculenta, are reported here. One is from of 'V' values in the paratypes (76-82) is soil about the roots of a mixture of plants more anterior than the composite for all (including cactus) near tile tide line at Bay- the populations of G. straeIeni reported in side, Texas. Dimensions are as follows: this paper. Until more collections of this 7 females: L = .28 (.22-.31) mm; a = nominal species are made, and more speci- 23 (19-26); b = 2.7 (2.4-3.0); V = 74 mens are available for examination, the (73-76); stylet = 51 (48-53) t~m. diagnosis is acknowledged as given and G. 3 males: L --- .33 (.31-.34) mm; a = 28 abietis is considered a distinct species. (27-30); c = 12 (10-14). Another collection is from a tea planta- Gracilacus acicula (Brown, 1959) Raski, 1962 tion, Balanoor Estate, Balehonnur District, Karnataka State, India. The dimensions Two slides identified by G. L. Brown are: as G. acicula were loaned by the Canadian 5 females: L = .25 (.24-.26) ram; a = National Collection of Nematodes for this 21 (17-25);b = 2.4 (2.3- 2.5);c = 15;V = study. One slide had I3 females, I male and 75 (74-76); stylet = 54 (53-56) /~m; pro- 1 juvenile (labelled paratypes) from pasture rhabdion = 48 (47-50) ~m; excretory pore sod at Met Bleu, Ontario. The second slide = 76 (69-89)/~m. contained 5 females from meadow sod near 3 males: L = .27 (.26-.28) mm; a = 26 Nesbitt, Manitoba, which was reported in (23-30); c = 12 (11-13); spicules = 16 the original description as a record of its (15-18) ~m; gubernaculum = 4 (3-4) /~m; distribution. T = 32 (26-44); excretory pore = 59 This species also has been found in soil (54-64) ~m. with mixed roots from several plant species The Texas sample also held specimens in a nursery near Winter Park, Florida. of P. minutus and the Balanoor Estate There are 32 females and 9 males in that sample held specimens of P. neoambly- collection. Two of the females retained the cepkalus. fourth-stage cuticle with very short pro- The other three collections, identified rhabdions attached. No juveniles were as G. aculenta, inclnded 12 females, 1 male found. The juvenile described by Brown and 1 juvenile from soil about the roots (2) as having a stylet 34 ttm long probably of sagebrush at German Flats, Salina, Utah; is the third stage. 9 females from grassy soil at Gobbler One other collection, identified tenta- Spring, Williams, Arizona; and 2 females, tively as G. acicula, has three females from 3 males in natural grassland at Matador, soil in a greenhouse at Michigan State Uni- Saskatchewan, Canada; 5 females, 1 male versity, Ingham County, Michigan. Unfor- from strawberry at Strzeloe, Poland. tunately these are considerably flattened Some of these specimens, including and some characters are difficult to discern. some labelled paratypes, have an incon- The tails are more slender-conoid and finely spicuous lateral vulvar membrane which is 100 Journal of Nematology, Volume/¢, No. 2, April 1976 not clearly present in most of the speci- goodeyi. According to Geraert (6), the mens. In all other respects, they resemble elongate spermatheca of G. goodeyi is an G. aculenta and are so considered in this important character differentiating it from paper. A small lateral vulvar membrane closely related species. It may be 3 times as indicates close relationship of G. idalima long at the corresponding body diameter. with G. aculenta, which differs in stylet The identification of G. goodeyi as having length [58 (54-62) t~m for G. aculenta vs. an elongate spermatheca is at variance with 83 (75-88) t~m for G. idalima]. the published description by Oostenbrink A composite of the dimensions of all (9). He described the spermatheca as round. five collections is: This character must be confirmed in other Female: L = .29 (.24-.34) mm; a = 23 type specimens since the paratypes at Uni- (17-28); b = 2.7 (2.3-3.0); c = 12 (10-15); versity of California, Davis, are useless. V = 72 (69-76); stylet = 57 (,t8-68) ~m; Another character which distinguishes excretory pore = 73 (59-89) t~m. G. goodeyi is the conoid head which pre- Male: L = .31 (.26-.35) ram; a = 27 sents a narrow tapering outline with fine (23-30); c = 13 (10-15); spicules = 18 but definite annules and is rounded at the (15-21) t~m; gubernaculum = 4 (3-5) /zm; apex. This appearance may be more pro- T = 33 (26-44); excretory pore = 66 nounced in mature, swollen females. The (54-77) tLm. tail of G. goodeyi also is more often sub- Gracilacus epacris acute to finely rounded as contrasted with (Allen and Jensen, 1950) that of the related species, G. straeleni, Raski, 1962 which has an acute or pointed tail outline. It is doubtful that the male described G. epacris from three new locations in by Oostenbrink is indeed Two California has been added to the UCNS G. goodeyi. other species of were re- Collection, Davis. Two females and 1 ju- Paratylenchus ported in that type collection of soil about venile were found in soil about the roots of a pear tree near Arnheim. One of those California laurel, UmbeIlularia californica could be males of which bear Nutt., near Fairfield, Solano County. A P. hamatus, a stylet. A similar situation was found in a single female was found in soil about the collection from Austria in which one male roots of U. californica near Shell Beach, of was found with 9 females Marin County. Three females and I ju- Paratylenchus of G. 1 female of venile were found in soil about the roots goodeyi, P. microdorus, and 1 female which appears to be of Quercus sp. near Livermore, Alameda P. neo- The male has no stylet but County. This is the same locality in which amblycephalus. does have fairly well-defined caudal alae, a 17 females, 8 males and 4 juveniles were characteristic which is unique for a male found in soil about the roots of Platanus of Paratylenchus. Geraert (6) reports a racemosa Nutt. The measurements and de- male of without stylet. So many scriptions of juveniles reported by Raski G. goodeyi collections are mixtures of species that it is (10) were based on these specimens from P. very difficult to associate specimens of one racemosa but a reference to the host source sex with the appropriate opposite sex. was not inclnded in that report. Mono-specific collections are invaluable for Gracilacus goodeyi resolving such problems. (Oostenbrink, 1953)Raski, 1962 The nine collections mentioned previ- Two slides with 10 females labelled as ously include 8 females from strawberry paratypes of P. goodeyi are on deposit in soil near Gliwice-Labedy, Poland; 6 females the U(]NS Collection, Davis. Three other from grass and weed soil, Laffrey, France; slides with six females labelled as /'. 5 females from grass and weed soil, Digne, goodeyi from Arnheim, The Netherlands France; 6 females, Jembloux, Belgium; 9 (and apparently from the type collection females from grass soil, Innsbruck, Austria; also) are in the UCNS Collection. Unfor- 14 females in soil at roots of Lolium, tunately, all the specimens on the five slides Agrostis and Festuca, at Kirton, Lincoln- are flattened beyond recognition. shire, England; 2 females from old meadow Nine other collections from six different soil, Limburg (Gronsveld), The Nether- countries have been identified as G. lands; 1 female from arable soil, Ruurlo, Revision of Paratylenchus Part III--Gracilacus: Raski 101

The Netherlands; 9 females from soil in ~Lm. Head similar to that of female. Tail garden at queen's palace, Soestdijk, The conoid; tip subacute to rounded. Develop- Netherlands. Except for stylet length, all ing gonad 9-11 tLm long (about 5/zm wide), these specimens fit the description of G. 49-60 ttm tYom terminus. .¢oodeyi. Several females have stylets which Another collection of 4 females and are shorter (the shortest being 44 /tin) than 21 juveniles from a peach nursery at the minimum reported in the type collec- Chateauneuf du Luc, near Valence, France tion. Thus the range of stylet length is was also identified as G. macrodora with 44-56/tm. dimensions as follows: 4 females: L = .32 (.31-.34) mm; a = Gracilacus intermedia Raski, 1962 24 (21-27); b = 2.6 (2.4-2.7); V = ~-' A collection of 6 females identified as (10-2~ 77 (74-78); stylet = 81 (75-86); G. intermedia was made from soil about prorhabdion = 74 (68-80) ~tm; excretory roots of Pinus ponderosa Dougl. and pore = 82 (79-86) ttm. Chamaebatia [oliolosa Benth. near Volcano, 3 juveniles (fourth stage ?): L = .31 Anlador County. Also present were speci- (.30-.34) ram; a = 23 (22-24); b = 4.2 (4.1- mens of G. straeleni. Other collections in- 4.4); stylet = lacking; excretory pore = 70 clude a single female of G. intermedia from ~tm. Head as in female of G. macrodora. California laurel, Umbellularia californica Excretory pore opposite posterior bull). Nutt., near Fairfield, Solano County, to- Gonad 43-46 /tin long, 74-76 ~m from gether with G. epacris and G. straeleni; 2 terminus. Tail slender conoid; tip rounded. females from grapevine near Napa, Napa No specimens of P. longistylosa were County; 2 females, 2 juveniles from prunes available for study and the conclusion that in Sonoma County; and 1 female from it is synonymous with G. macrodora is cypress, Cupressus sp., on the University of based on the published description. The California Berkeley campus together with diagnosis of P. longistylosa (4) indicates Para tylench us neoa rn blycephalus. that it is most closely related to P. macro- phallus, a conclusion which is doubtful. In Gracilacus ivorensis appearance and most measurements, it is (Luc and de Guiran, 1962) n. comb. more closely related to G. macrodora. syn. Paratylenchus ivorensis Range of total length (given as .27-.31 mm Luc and de Guiran, 1962 for P. longistylosa) is the only major dif- Paratypes, 7 females and 1 male, of this ference noted [for G. macrodora L = .34 species were available for study. The speci- (.33-.39) ram], and this is not sufficient for mens conform closely with the description, specific differentiation. but I noted a greater range of variability Gracilacus mira Raski, 1962 in female tail shape than was illustrated. The tails of some females end in an ex- Specimens identified as G. mira have tremely fine, acute terminus and one fe- been collected from five new localities in male has a rather bluntly rotmded tail California. One female was collected fronl terminus. soil about roots of Arctostaphylos sp. in Gracilacus macrodora Mix Canyon near Vacaville, Solano County. (Brzeski, 1963) n. comb. The collection also held specimens of syn. Paratylenchus macrodorus Paratylenchus neoamblycephalus and G. Brzeski, 1963 straeleni. Fifteen females and 7 juveniles Paratylenchus Iongistylosa were collected from soil about roots of Cali- Dement'eva, 1972 n. syn. fornia juniper, .luniperus cali[ornica Cart., and 3 females from soil about roots of Eight females and 3 juvenile paratypes Rhus trilobata Nutt. in Joshua Tree Na- of G. macrodora were available for study. tional Monument, San Bernardino County. The female paratypes fit the description Eight females of Paratylenchus variabilis precisely. The juveniles were measured as Raski, 1975 also were found in the Rhus follows: (3 second stage ?): L = .25 (.20-.30) trilobata sample. Three females and 1 ju- ram; a = 20 (18-21); b = 2.3-2.4; stylet = venile were collected about roots of Tar- 45 (43-49) /tin; prorhabdion = 38 (37-39) tarian cherry on Mahaleb rootstock near 102 Journal of Nematology, Volume 8, No. 2, April 1976 Linden, San Joaquin County. Four females baldacci Raski, 1975 and Paratylenchus were collected about prunes in Tehama nainianus Edward and Misra, 1963. County and 1 female from peach near Six swollen females were recovered di- Modesto, Stanislaus County. rectly from roots of Laurus nobilis L. near Bari together with 5 swollen females GraciIacus mutabilis of Rotylenchulus macrodoratus Dasgupta, (Colbran, 1969) n. comb. Raski and Sher, 1968. syn. Paratylenchus mutabilis Colbran, 1969 Gracilacus robusta (Wu, 1974) Four slides with 10 females, 3 males, n. comb. and 4 juveniles from the type locality (two syn. Paratylenchus robustus of the slides labelled paratypes), were avail- Wu, 1974 able for this study. The adults are precisely Specimens of G. rob usta were not avail- as described by Colbran. The juveniles, two able for this study. According to tile de- of which were in the process of molting, scription and illustration (14), G. robusta appear to be third stage. Dimensions of the has a number of characters to support a juveniles are as follows: L = .23-.25 toni; relationship with G. straeleni. The range a = 17-18; b = 2.5-2.7; stylet = 33-37 /,m; of total length, 'V' value, position of ex- prorhabdion = 26-29/~m; excretory pore = cretory pore opposite isthmus, and rounded 69 /,m. Head similar to that of adult fe- head with fine annules are all similar to males with distinctly rounded submedian G. straeleni. However, the presence of a lobes not set off by constriction. Excretory spermagonium is unique for this genus and pore opposite isthmus. Tail conoid, sub- also for Paratylenchus. Because ot~ the acute to bluntly rounded. One specimen spermagonium and the greater stylet showed a conspicuous parasite similar to length [55 (44-66) /,m for G. straeleni vs. the one illustrated in G. pandata n. sp. 58-73 /,m for G. robusta], this species is ac- Gracilacus oostenbrinki cepted as distinct and separate. (Misra and Edward, 1971) n. comb. Gracilacus steineri (Golden, 1961) syn. Paratylenchus oostenbrinki Raski, 1962 Misra and Edward, 1971 syn. Paratylenchus steineri G. oostenbrinki is diagnosed by the Golden, 1961 authors (8) as being related to G. peratica, Three slides with single female para- G. intermedia and G. mira. Its description, types were available for this study. In addi- however, indicates it is more closely related tion, a single slide with 4 females and 1 to G. macrodora. The greater size of G. male, identified as G. steineri, was sent by macrodora (.33-.39 mm vs..25-.28 mm for R. C. Colbran from soil around a native G. oostenbrinhi) and position of excretory legume at Western Ridge, corner of west pore (near stylet knobs in G. mira vs. op- Road, Mt. Tamborine, Australia. These posite nerve ring in G. oostenbrinhi) is evi- have the following dimensions: dence these are two separate and distinct 4 females: L = .29 (.28-.31) mm; a = species. 26 (24-29); b = 2.4 (2..8-2.5); V = 76 Gracilacus peratica Raski, 1962 (76-77); styler = 73 (68-75) /,m; excretory pore = 86 (75-96)/~m. Four new records of this species were 1 male: L = .35 ram; a = 33; c = 15; from samples taken in Italy. Five females, spicules = 17 /,na; gubernaculum = 4 tim; 2 males, and 9 juveniles were identified T = 32; excretory pore --- 86 /,m. from soil collected near Pescara. Thirteen The stylet of these specimens is longer females, 2 males, and 5 juveniles were but overlaps the range described for G. found in soil about roots of olive near steineri. In other respects these specimens Molise, Campobasso. Three females were and G. steineri are considered conspecific. found in soil about roots of Paulsen grape hybrid 1103 in the government nursery Gracilacus straeleni near Palermo, Sicily. Also present in that (de Coninck, 1931) n. comb. sample were specimens of Paratylenchus syn. Procriconema straeleni Revision of Paratylenchus Part IlI--Gracilacus: Raski 103

de Coninck, 1931 form closely with G. straeleni. Therefore, Paratylenchus audriellus it is concluded G. audriella is a synonym Brown, 1959, n. syn. of G. straeleni. Other collections from the ParatyIenchus straeleni U. S. A. and Europe show similar varia- (tle Coninck, 1931) Oostenbrink, 1960 tions with a longer styler more typical of Paratylenehus sarissus G. straeleni. The specimens are strong evi- Tarjan, 1960 dence that G. straeleni is widely distributed and has a variable tail shape and a variable There are no original type specimens stylet length. of G. straeleni extant. Geraert (6) reported A composite of all the previous identifi- a collection from moss anti soil from the cations indicates a species with the follow- type locality, Baraque Michel, Liege, Bel- ing ranges of dimensions: gium, in which he found one female. He Female: L = .37 (.28-.44) ram; a = 23 briefly described that specimen and desig- (16-30); b = 3.5 (3.0-4.5); c = 12 (10-14); nated it as neotype. Eight collections, V = 82 (79-85); styler = 55 (44-66) ttm. identified as G. straeleni, from Europe Male: L = .37 (.35-.40) mm; a = 31 were studied in this review. One collection (28-35); c = 12 (10-15); spicules = 22 from sphagnum moss at Gribsee-Moor, Den- (21-24) /tin; gubernaculum = 4 (4-5) t~m; mark held 11 females and 1 male. No other T = 27 (24-35); excretory pore = 83 species of Paratylenchus or Gracilacus were (79-90) /tm. present. This is the first record of a male of New distribution records in California G. straeleni from Europe. The dimensions include: Cypress, Cupressus sp., near for that population are as follows: Bolinas, Matin County; California laurel, 5 females: L = .39 (.37-.44) mm; a = Umbeltularia cali[ornica Nutt., near Fair- 24 (22-25); b = 3.6 (3.4-4.0); c = 13 field, Solano County, near Willits, Men- (12-13); V = :,2 1~,-4~) 81 (80-82); stylet = docino County, and on the American River 60 (57-63) /~m; prorhabdion = 47 (45-49) near Auburn, Placer County; soil on Uni- ttm; excretory pore = 93 (88-98) /~m. versity of California Berkeley campus; 1 male: L = .36 ram; a = 30; c = 10; Pinus attenuata Lemmon and Arctostaphy- spicules = 21 t~m; gubernaculum = 5 /~m; los sp., Moraga Ridge, Contra Costa T = 28; excretory pore = 82 #m. County; Black sage, Artemesia arbuscula Two female and 2 male paratypes of G. Nutt., Badger Canyon; Quercus sp., Mt. sarissa were studied. Also 14 female and 5 Diablo, Contra Costa County; Quercus sp., male paratypes of G. audriella were loaned near Monticello, Napa County; Quercus sp., from the Canadian National Collection of and Pinus sp., Hobo Gulch Camp, Trinity Nematodes for this review. On the basis of County; Quercus sp., near Vacaville, Solano the synonymy of these two species with County; Quercus sp., Tecate area, San G. straeleni, 31 collections from the U. S. A. Diego County; Quercus sp., near San Lucas, and Canada, 11 of which have 1 or more Monterey County; Quercus sp., near Mill- males, have been identified as G. straeleni. brae; Camellia japonica L., Huntington Females and males also were found in 1 Gardens, Los Angeles County; Opuntia sp., collection from New Zealand. locality unknown; grassy soil, Asilomar; The decision of Geraert (6) that G. Pinus ponderosa Dougl. and Charnaebatia sarissa is a synonym of G. straeleni is ac- [oliolosa Benth., near Volcano, Amador cepted. Geraert further judged that G. County; Aesculus californica (Spach) Nutt., straeleni is not so different from G. au- in Mix Canyon, near Vacaville, Solano driella and noted that G. audriella has a County; and redwoods and U. cali]ornica, more sharply pointed tail and shorter stylet. on a stream bed in Henry Cowell Redwoods Brown (2) stressed the hook-like termi- State Park, near Felton, Santa Cruz County. nus of the tails in both males and females Distribution records in other states in- of G. audriella as a distinguishing char- clude: Grey birch roots and soil, Lafayette acter. In fact, the paratypes show conoid, Campground, Franconia Notch, New sharply-pointed tails on some females as Hampshire; Quercus sp., Great Smoky Na- well as typical hook-like processes on others. tional Park, Tennessee; pecan grove, By- In all other respects, these paratypes con- ron, Georgia; Pinus sp., 9.7 km south of

Revision ot7 Paratylenchus Part III-Gracilacus: Raski 105

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Body rather broad, narrowing only slightly tan from terminus; suggests second stage. to rounded head with slightly projecting, No juveniles with elongate stylet present. rounded submedian lobes as in female. Two slender females in last molt present Stylet small; knobs only slight swellings. inside a swollen female. Each molted cuticle Body contents coarsely granular through- with prorhabdion 9 /zm long. out; details of internal morphology ob- Holotype: Female, collected by C. M. scured. Tail bluntly rounded. Developing Heald on January 22, 1973, slide number gonad appears to be 9-10 /~m long, 65-93 1446, UCNS Collection, Davis, California. 106 journal of Nematology, Volume 8, No. 2, April 1976 A llotype: Male, same data as holotype, rounded anteriorly to same level as lateral slide number 1447, UCNS Collection, Davis, lobes; produce a somewhat flat or truncate California. anterior surface. Sclerotization light; stylet Paraiypes: 40 females, 104 swollen fe- guiding apparatus appears as two dark males, 8 males, 87 juveniles, same data as lines slightly curved at level of retracted holotype, deposited as follows: I1 females, stylet tip. Stylet slender; knobs rounded; 43 swollen females, 1 male, 45 juveniles in apparently slightly anteriorly directed. UCNS Collection, Davis, California; 22 fe- Length of stylet shaft plus knobs 9 /~m males, 53 swollen females, 3 males, 37 ju- (7-9 ~m in paratypes). Excretory pore an- veniles in USDA Nematode Collection, terior to base of stylet (Fig. 9). Hemizonid Beltsville, Maryland; 4 females, 2 swollen inconspicuous, immediately posterior to ex- females, 1 male, 5 juveniles in National cretory pore. Esophago-intestinal valve Nemotode Collection, IARI, New Delhi, small, rounded. Ovary outstretched (Fig. India; 1 female, 2 swollen females, 1 male 13); spermatheca inconspicuous (Fig. 10), each at Nematology Department, Roth- without sperm. Vulva lips slightly rounded amsted Experimental Station; Plantenziek- and protruding; lateral vulvar membranes tenkundige Dienst, Wageningen, The lacking. Tail long, slender-conoid; annules Netherlands; UCNS Collection, Riverside, very fine but visible; subacute, to finely California. rounded terminus. Body annules average about 1.0 ~m in width; lateral field with Type host: Mesquite, Prosopis sp. 3 incisures equally spaced; center line less Type locality: Weslaco, Texas. distinct than outer 2 (Fig. 11-12). Allotype (male): L --- .32 mm; a = 28; Diagnosis: This species is most closely c = 12; spicules = 18 /~m; gubernaculum related to G. aculenta from which it differs = 5/ml; T -- 37; excretory pore = 63 #m. in longer stylet [58 (51-66) /zm for G. acu- Body an open 'C' shape after fixation. Body lenta] and length of esophagus (b = 2.4- narrows gradually anteriorly to rounded 4.1 in G. aculenta). head. Submedian lobes slightly rounded, Distribution: 9 females, 3 males and 2 not set off; lateral area slightly depressed; juveniles found in soil about roots of Mes- give concave anterior outline. Sclerotization embryanthernum sp. at Dillon Beach, Matin light. Stylet lacking, esophagus degenerate; County, California; 7 females, 2 males and only remnants identifiable (Fig. 14). Excre- 3 juveniles from soil about strawberry, tory pore immediately anterior to hemi- Noyo, Mendocino County, California. zonid which extends about 3 body annules Gracilacus costata n. sp. in length. Testis outstretched; anterior (Fig. 9-15) third with large developing cells; posterior portion filled with small sperm. Spicules Paratypes (13 females): L = .31 (.29- slightly curved; cloacal sheath short without .34) mm; a = 25 (22-29); b = 2.4 (2.3-2.6); projecting posterior margin. Tail slender c -- 11 (11-12); V = ~'~ (12-ls~ 70 (67-72); conoid; annules visible almost to acute stylet -- 77 (70-87) /~m; prorhabdion = 69 terminus (Fig. 15). Body annules average (63-78) #m; excretory pore = 69 (56-79) about 1.0 /~m in width; lateral field with 3 /zm. incisures equally spaced; center line less Paratypes (4 males): L = .34 (.32-.35) distinct than outer 2. ram; a = 28 (26-31); c = 11 (11-12); spic- Juveniles (3 fourth stage ?): L = .28 ules = 20 (18-22) ~m; gubernaculum = 5 (.26-.30) ram; a = 18 (15-22); b = 3.6 (3.6- (4-6) /~m; T = 37 (33-41); excretory pore 3.7); stylet = 11 (10-13)/~m; prorhabdion = 66 (62-72) t~m. = 7 (6-7) /~m; excretory pore -- 69 (64-73) Holotype (female): L -- .31 mm; a = /~m. Head rounded with submedian lobes 26; b = 2.3; c ~- 11; V = 17 71; stylet = or lips as in female; rounded but not set 79 ~tm; prorhabdion ~- 70 ~m; excretory off. Stylet very weakly developed; knobs pore = 64 t~m. Body strongly curved al- small swellings. Excretory pore at level of most a closed 'C' shape after fixation. Body isthmus. Developing gonad not distinct. narrows gradually anteriorly to a rounded Tail subacute to rounded. head. Submedian lobes or lips not set off; Holotype: Female, collected by L. H. Revision of Paratylenchus Part IiI--Gracilacus: Raski 107

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IZlG. 9-19. 9-15) Oracilacus costata n. sp. 16-19) Gracilacus solivaga n. sp. Wang on May 21, 1967, slide number 1442, holotype except these were collected on UCNS Collection, Davis, California. May 15, 1967, deposited as follows: 8 females, 13 males and 10 juveniles in UCNS AlIotype: Male, same data as holotype, Collection, Davis, California; 1 female, 1 .slide number 1443, UCNS Collection, Davis, male each at USDA Nematode Collection, C.alifornia. Beltsville, Maryland; National Nematode Paratypes: 11 females, 8 males, 9 ju- Collection, IARI, New Delhi, India; Nema- veniles, same data as holotype plus 2 re- tology Department, Rothamsted Experi- males, 10 males, 1 juvenile, same data as mental Station, Harpenden, England; 108 Journal o/Nematology, Volume 8, No. 2, April 1976 Plantenziektenkundige Dienst, Wagen- Holotype: Female, collected by M. T. ingen, The Netherlands; UCNS Collection, Hutchinson on March 22, 1962, slide num- Riverside, California. ber 1448, UCNS Collection, Davis, Cali- Type habitat: Noncultivated loam soil. fornia. Type locality: Mt. Yangmin, Taipei, Paratypes: 5 females, same data as halo- Taiwan. type, deposited in UCNS Collection, Davis, Diagnosis: This species is most closely Cali fornia. related to G. solivaga n. sp. from which it Type habitat: Upper jungle soil. differs in its greater length [L = .24 (.21- Type locality: Mooloya, Hewaheta, Sri .25) mm for G. solivaga] and slender-conoid Lanka. subacute tail (tail mostly deformed in G. Diagnosis: This species is most closely solivaga). G. costata is also related to G. related to G. costata from which it differs latescens from which it differs also in larger in its smaller size [.31 (.29-.34) mm for G. size [L = .27 (.22-.29) mm for G. latescens] costata] and its variously misshapen tail and subacute tail (tail is bluntly rounded (uniformly subacute in G. costata). in G. latescens). Also present in this soil sample were 13 Distribution: 11 females, 2 males and 20 females, 2 males, and 1 juvenile of Paraty- juveniles from soil about Manihot utilis- lenchus minutus. sima Pohl., at the property of Rafael n. sp. Navarro A. in Colombia, South America. Gracilacus pandata (Fig. 20-24) Gracilacus soIivaga n. sp. Paratypes (10 females): L = .38 (.33- (Fig. 16-19) .42) mm; a ---- 26 (23-32); b ---- 3.1 (2.8- Paratypes (5 females): L = .24 (.22-.25) 3.2); c = 11 (9-12); V = 2z (19-31) 74 ram; a = 24 (22-25); b = 2.1 (2.0-2.1); c = (70-76); stylet = 66 (63-70) /~m; prorhab- ?; V = 14 (13-,4~ 73 (71-75); stylet = 71 dion = 55 (53-59) /~m; excretory pore = (69-75) /zm; prorhabdion = 64 (60-69) tim; 11)3 (94-119) ttm. excretory pore = 54 (52-56)/tm. Paratypes (3 males): L = .39 (.36-.42) Holotype (female): L = .25 ram; a = mm; a = ?; c = 10; spicules -- 20 (19-22) 25; b = 2.1; c = ?; V ~ a4 71; stylet = 74 #m; gubernaculum = 4 (4-5); T = 36 ttm; prorhabdion = 69 ~m; excretory pore (34-40); excretory pore -- 89 (83-94)/~nl. = 52 ttm. Body an open 'C' shape after Holotype (female): L = .40 mm; a = fixation. Body gradually narrows anteriorly 28; b = 3.1; c = 11; V = 29 75; stylet = to a rounded head. Sclerotization very light; 68 ~m; prorhabdion = 55 ttm; excretory styler guiding apparatus appears as two pore = 102 #m. Body only slightly curved slightly arcuate, darker lines at tip of re- ventrad after fixation. Body gradually nar- tracted stylet. Styler slender; slightly arcuate rows slightly to rounded head. Submedian ventrad (Fig. 17). Knobs small, rounded; lobes small rounded; protrude only very slightly posteriorly directed. Stylet shaft slightly beyond round contour of head. plus knobs 5 /~m (6-7 in paratypes). Ex- Sclerotization light; styler guiding appa- cretory pore about 2 body widths anterior ratus appears as two dark, slightly curved to stylet knobs. Hemizonid immediately lines at tip of retracted stylet. Stylet slender, posterior to excretory pore. Isthmus slender, slightly curved; knobs large, rounded, relatively short. Esophago-intestinal valve slightly posteriorly directed. Excretory pore small, rounded. Ovary outstretched; sperm- at level of anterior end of isthmus (op- atheca indefinite; no sperm seen (Fig. 19). posite nerve ring in some paratypes). Hemi- Vulva inconspicuous; lateral vulvar mem- zonid 2 body annules in length at level of branes lacking. Tail long, slender-conoid; excretory pore. Esophago-intestinal valve narrows gradually; annules distinct to mis- small, rounded. Ovary outstretched; sper- shapen; lobed terminus [paratypes vari- matheca narrow, elongate (7 x 25 /tm); ously misshapen, two subacute-conoid (Fig. filled with large sperm (Fig. 22). Vulva in- 18)]. Body annules fine; less than 1 ttm in conspicuous; only a slight notch in ventral width. Lateral field with 3 incisures (Fig. outline. Lateral vulvar membrane rounded, 16), center one lighter than outer 2. prominent. Tail long, slender-conoid end- Male: Unknown. ing in narrow, finely rounded, almost acute Revision of Paratylenchus Part III--GraciIacus: Raski 109 l:l ~,~/~.y l i /i,! ,,i/t 25 // i t ~! 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• \ / ' "k :. fly \~.5..7 7::-:"" 27 28 20 22 FIG. 20-28. 20-24) Gracilacus pandata n. sp. Fig. 20) Two specimens ot: Bacillus penetrans present on anterior end of female. 25-28) Gracilacus parvula n. sp. terminus. Body annules average 1.3-1.8 /~m short, inconspicuous, without projection of wide. Lateral field with 4 incisures (Fig. posterior margin. Tail slender, conoid; 21); inner 2 lighter than outer 2 [several terminus finely rounded (Fig. 24). paratype females had conspicuous, infec- Holotype: Female, collected by F. E. tions of Bacillus penetrans (syn. Dubosqia Caveness on August 31, 1964, slide number penetrans) on cuticle, especially on anterior 1399, UCNS Collection, Davis, California. end (Fig. 20)]. Allotype: Male, collected by F. E. Cave- Allolype (male): L = .36 mm; a = ?; ness on July 10, 1964, slide number 1400, c = 10; spicules -- 18 /~m; gubernaculum UCNS Collection, Davis, California. = 4/nn; T = 31; excretory pore = 87 /~m. Paratypes: 26 females, same data as holo- Body an open 'C' shape after fixation. Body type, and 1 female, 3 males, same data as narrows gradually to slender, rounded allotype, deposited as follows: 22 females, head. Sclerotization very light; styler lack- 3 males in UCNS Collection, Davis, Cali- ing (Fig. 23). Testis outstretched. Spicules fornia; 1 female each in National Nema- slightly curved, ventrad. Cloacal sheath tode Collection, IARI, New Delhi, India; 110 Journal of Nematology, Volume 8, No. 2, April 1976 USDA Nematode Collection, Beltsville. .29) mm; a = 22 (22-23); b = 2.6 (2.0-3.2); Maryland; Nematology Department, Roth- stylet = 40 (38-42) /,m; prorhabdion ~--- 32 amsted Experimental Station, Harpenden, (30-34) /~m; excretory pore = 70 (61-80) England; Plantenziektenkundige Dienst, /,m. Head with distinct set-off lips as in Wageningen, The Netherlands; UCNS Col- female. Stylet knobs large, backwardly di- lection, Riverside, California. rected. Excretory pore opposite isthmus. Type host: Grapefruit (holotype); water Gonad about 21 /,m long, 66-68 /~m from yam, Dioscorea alata L. (allotype). terminus. Tail bluntly rounded. Type locality: 1 mile north of Katsina Holotype: Female, collected May 15, Twp., Katsina Province, Nigeria (holo- 1967, slide number 1449, UCNS Collection, type); 8 miles southwest of Jebba, Ilorin Davis, California. Province, Nigeria (allotype). Paratypes: 4 females, 11 juveniles, same Diagnosis: This species is most closely data as holotype, deposited in UCNS Col- related to G. goodeyi from which it is dis- lection, Davis, California. tinguished by its longer stylet (63-70 tzm vs. 48-56 tzm for G. goodeyi) and rounded Type habitat: Noncultivated loam soil. head (conoid in G. goodeyi). Type locality: Mt. Yangmin, Taipei, Taiwan. Gracilacus parvula n. sp. Diagnosis: This species is most closely (Fig. 25-28) related to G. mutabilis but differs in the set-off protruding lips of G. parvula (in G. Paratypes (4 females): L = .35 (.34-.37) mutabilis the head outline is continuous, mm; a = 28 (27-29); b = 3.3 (3.2-3.4); c not set off, but with submedian lobes = 16; V = 25 ~,oa-2s) 79 (78-80); stylet = rounded anteriorly). The stylet is longer in 58 (56-60) /,m; prorhabdion = 46 (45-48) G. mutabilis [58 (56-60) /~m vs. 48-57 /,m /~m; excretory pore = 91 (86-96) /,m. for G. mutabilis]; stylet knobs of G. parvula Holotype (female): L = .35 mm; a = larger than in G. mutabilis. 28; b = 3.3; c -- 17;V = 28 79); stylet -- Additional collection: After this manu- 59 ~m; prorhabdion = 47 t,m; excretory script was submitted for publication but pore = 89 /,m. Body an open 'C' shape before it was set for printing, another col- after fixation. Head narrows with rounded lection of this species was provided by outline, distinct annules, to constriction C. S. Huang. The collection held 10 males, setting off rounded, protruding submedian 5 juveniles, and 7 females from soil about lobes or lips, truncate or flattened at an- roots of Sunki orange rootstock near Chiayi, terior surface. Sclerotization light; stylet Taiwan in May 1975. Measurements of guiding apparatus appears as two dark males and females and a description of the lines, slightly arcuate, near tip of retracted male by C. S. Huang are: styler (Fig. 26). Styler slender; knobs rel- 12 females (Chiayi): L = .35 (.32-.37) atively large, directed slightly posteriad. ram; a = 24 (19-30); b -- 3.4 (3.0-3.8); c = Excretory pore opposite iqhmus. Hemi- 16 (15-19); v = 82 (81-84). zonid 3 body annules in length immedi- 9 males (Chiayi): L = .31 (.29-.35) ram; ately anterior to excretory pore. Length of a = 29 (27-30); c = 14 (11-16); spicules = stylet shaft plus knobs 12 t,m (11-12 txm 19 (17-20) /,m. in paratypes). Isthmus slender, relatively Male (Chiayi): Body shape after fixa- short. Esophago-intestinal valve small, tion variable. Some an open 'C' shape; lobate. Ovary outstretched; spermatheca others tightly curved; tail overlapping small, ovate, without sperm. Vulva incon- body. Body gradually narrows anteriorly, spicuous; lateral vulvar membranes dis- then more abruptly at rounded head. tinct, rounded (Fig. 27). Tail conoid; nar- Sclerotization delicate. Stylet lacking. Hemi- rows gradually to bluntly rounded terminus zonid anterior to excretory pore; occupies (Fig. 28). Lateral field with 4 incisures about 3 body annules in length. Testis out- about equally spaced; inner 2 lighter than stretched. Cloacal opening with prominent outer 2 (Fig. 25). sheath and hooked posterior margin. Spic- Male: Unknown. ules slender, curved. Caudal alae not ob- Juvenile (4 third stage ?): L -- .26 (.25- served, Tail narrows gradually with slender Revision of Paratylenchus Part III-Gracilacus: Raski 111 conoid shape; terminus finely rounded, al- (72-92) /~m; excretory pore = 90 (80-110) most acute. #m. Paratypes (2 males): L = .34-.35 mm; Gracilacus enata n. sp. a ---- 33-34; c = 10; spicules = 17-18 /zm; (Fig. 29-35) gubernaculum = 3 /~m; T = 31-38; excre- Paratypes (16 females): L = .31 (.25- tory pore = 75-80 ffm. .35); a = 26 (23-28); b = 2.3 (2.1-2.4); c Holotype (female): L = .35 ram; a = = 12 (11-13); V = ,7 (1~-.~0) 78 (74-80); sty- 27; b = 2.4; c = 12; V = ~7 77; stylet = let = 94 (82-104) ffm; prorhabdion = 83 104 ffm; prorhabdion = 92 /zm; excretory

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FIG. 29-40. 29-35) Gracilacus enata n. sp. 36-40) Gracilacus teres n. sp. 112 Journal of Nematology, Volume 8, No. 2, April 1976 pore = 97 /ma. Body only slightly curved at level of isthmus. Gonad 39 (29-50) t~m ventrally after fixation. Body gradually nar- long, 70 (65-77) ~m from terminus. Tails rows anteriorly; ends in a constriction or slender, conoid; some narrow abruptly near depression setting off sub-median lobes or terminus and give hook-like or digitate ap- lips which are rounded and protrude for- pearance. Terminus finely rounded to ward. Sclerotization light; stylet guiding acute. apparatus appears as two dark lines, slightly Holot)~pe: Female, collected by A. Li- curved, near tip of retracted stylet (Fig. 29, zana on September 26, 1968, slide number 30). Stylet slender, with rounded knobs. 1444, UCNS Collection, Davis, California. Length of stylet shaft plus knobs 12 /~m A llotype: Male, same data as holotype, (8-14 tim in paratypes). Excretory pore slide number 1445, UCNS Collection, anterior to styler knobs. Ovary outstretched Davis, California. (Fig. 32); spermatheca spherical (spherical Paratypes: 24 females, 2 males, 18 ju- to slightly ovate in paratypes). Rounded veniles, same data as holotype, deposited as vulvar lips protrude slightly. Lateral vulvar follows: 17 females, 2 males, 12 juveniles membranes absent. Tail slender conoid; an- in UCNS Collection, Davis, California; 3 nules distinct up to narrow, acute terminus. females, I juvenile in USDA Nematode Col- Body annules average less than 1 t~m in lection, Beltsville, Maryland; 1 female, 4 width. Lateral field with 4 incisures equally juveniles in National Nematode Collection, spaced and distinct (Fig. 31). IARI, New Delhi, India; I female, 1 ju- Allotype (male): L = .34 ram; a = 36; venile in Nematology Department, Roth- c --- 9; spicules = 17 ~m; gubernaculum = amsted Experimental Station, Harpenden, 4 /~m; T = 31; excretory pore = 75 t~m. England; 1 female each in Plantenziekten- Body an open 'C' shape; curved dorsally kundige Dienst, Wageningen, The Nether- after fixation. Body gradually narrows an- lands, and UCNS Collection, Riverside, teriorly to a slender rounded head. California. Sclerotization light; stylet lacking (Fig. 33). Type host: Custard apple, or Chirimoya Hemizonid at level of excretory pore. Testis var. Concha Temarana, Annona cherimola outstretched; sperm small. Testicular gland Mill, prominent. Spicules curved slightly; cloacal Type locality: Quillota, Chile. sheath short, without protrusion of pos- Diagnosis: This species is most closely terior margin. Caudal alae not observed. related to G. teres n. sp. from which it dif- Tail slender, conoid; terminus finely fers in the longer stylet [74 (72-75) /~m for rounded, almost acute (Fig. 35). Body an- G. teres] and acute tail (rounded in G. nules average less than 1 tim in width. Ieres). Lateral field with 4 incisures equally spaced and distinct (Fig. 34). Gracilacus teres n. sp. Juveniles (4 second stage ?): L = .23 (Fig. 36-40) (.18-.29) ram; a --- 20 (18-23); b = 2.6 (2.1- 3.2); styler = 49 (45-51) t~m; prorhabdion Paratypes (14 females): L = .31 (.29-.35) = 41 (38-43) tan; excretory pore = 70 (64- mm; a = 21 (19-25); b = 2.6 (2.4-2.7); c = 77) ~m. Body narrows at head to set-off 13 (10-15); V = 18 ~t~-20) 74 (72-75); sty- protruding lips or submedian lobes similar let ~ 74 (69-83) /zm; prorhabdion --- 68 to female. Stylet slender; distinct and well- (64-77) t~m; excretory pore = 77 (70-85) developed knobs directed posteriad. Ex- /~m. cretory pore at level of isthmus. Gonad 7 Paratypes (6 males): L = .35 (.29-.38) (6-8) t~m long, 59 (55-65) ~m from terminus. mm; a = 30 (29-31); b = 3.9; c = 16 (15- Tail slender, conoid; ends in finely rounded 17); spicules -- 18 (16-19) /~m; guber- terminus. naculum = 4 /zm; T = 35 (34-37); excre- Juveniles (6 fourth stage): L -- .31 (.27- tory pore ~ 71 (55-80)/~m. .34) mm; a = 24 (21-27); b = 3.3 (3.1-3.6); Holotype (female): L = .30 ram; a = excretory pore = 81 (66-89) ttm. Body nar- 21; b = 2.4; c = 13; V = 20 74; stylet = rows at head to set-off protruding lips or 7.'~ tzm; prorhabdion = 67 /~m; excretory submedian lobes similar to females. Stylet pore = 78 ~m. Body after fixation an open lacking. Procorpus elongate. Excretory pore 'C' shape; more strongly curved posterior Revision of Paratylenchus Part III--Gracilacus: Raski 113 to vulva. Body very gradually narrows an- 84) ~m. Head rounded similar to female. teriorly, with distinct annules up to slight Styler much reduced, knobs slight swellings. constriction which sets off small, rounded, Excretory pore near posterior end of meta- submedian lobes or lips. Lateral lips corpus. Developing gonad 16-4l t~m long, rounded, at same level with submedian 72-104 /~m from bluntly rounded terminus. lobes, give tri-lobed outline which is Holotype: Female, collected by R. D. rounded (almost truncate outline in some Sharma in 1973, slide number 1450, UCNS paratypes). Sclerotization of head, light; Collection, Davis, California. styler guiding apparatus appears as two Allotype: Male, same data as holotype, dark straight lines near tip o[ retracted slide number 1451, UCNS Collection, styler (Fig. 39). Stylet slender; curves dorsad Davis, California. more than curvature of body; knobs small, Paratypes: 14 females, 6 males, 19 ju- rounded. Length of styler shaft plus knobs veniles, same data as holotype, deposited as 6 /~m (5-7 t~m in paratypes). Dorsal gland follows: 6 females, 1 male, 17 juveniles in orifice not visible because esophageal hnnen UCNS Collection, Davis, California; 2 fe- coils. Excretory pore at level between stylet males, 1 male, 1 juvenile in National Nem- knobs and valve of median bulb. Small atode Collection, IARI, New Delhi, India; hemizonid at level of excretory pore. 2 females, 1 male in USDA Nematode Col- Isthmus slender, elongate. Esophago-in- lection, Beltsville, Maryland; 1 female, 1 testinal valve distinct, small, lobate. Ovary male, 1 juvenile in Nematology Depart- outstretched; spermatheca inconspicuous, lnent, Rothamsted Experimental Station, without sperm (Fig. 40). Vulva inconspicu- Harpenden, England; 1 female, 1 male each ous without lateral vulvar membranes. Tail at Plantenziektenkundige Dienst, ~:agen- conoid; narrows gradually, with distinct ingen, The Netherlands, and UCNS Collec- annules almost to bluntly rounded termi- tion, Riverside, California. nus. Body annules average 1.0-1.5 btm ira Type host: Theobroma cacao L. width. Lateral field with 4 incisures, equally Type locality: Linhares, Espirito Santo, spaced about 1 t~m apart, inner 2 lighter Brazil. than outer 2 (Fig. 38). Diagnosis: This species is most closely Allotype (male): L = .38 mm; a = 31; related to G. steineri and G. enata n. sp. It c = 15; spicules = 18 ~tm; gubernaculum differs from G. steineri in its set-off, pro- = 4 ~tm; T = 36; excretory pore = 80 ~m. truding lips (bead rounded in G. steineri) After fixation, body coils tightly, especially and longer stylet [67 (65-69) sLm for G. in posterior portion. Body gradually nar- steineri]. It differs from G. enata by its rows anteriorly, then narrows more rounded tail (acute in G. enata) and shorter abruptly near lips to set-off small protrud- stylet [94 (82-104) /~m in G. enata]. ing lips or submedian lobes which are some- Distribution: Four other collections, all what like holotype head but lips do not from T. cacao, were identified as this spe- protrude so distinctly. Sclerotization deli- cies. One is from Linhares, the type locality cate; stylet lacking (Fig. 37). Hemizonid 2 of G. teres; two others are from the same body annnles posterior to excretory pore. general area of Espirito Santo; and the Testis outstretched, surrounded in pos- fourth is from Mutuipe at Fazenda do Dr. terior portion by prominent testicular Rni. gland. Spicules curved slightly. Cloacal SPECIES INQUIRENDAE sheath distinct, without hook-like extension o[ posterior margin. Tail narrows gradually Paratylenchus strenzkei (Volz, 1951) to conoid outline, with bluntly rounded Oostenbrink, 1960 terminus (Fig. 36). Body annules average syn. Hemicycliophora 1.0-1.5 ~m in width. Lateral field very ( = Procriconema) strenzkei faint, with 4 incisures equally spaced; in- Volz, 1951 ner 2 lighter than outer 2. There are no type specimens of P. stren- [uveniles (5 fonrth stage ?): L = .34 zkei. Volz (13) described his specimens as (.30-.36) ram; a = 21 (17-25); b = 4.1 (3.9- juveniles, but undoubtedly the vulva was 4.2); stylet = 16 (15-19) t~m; prorhabdion mistaken for the anus, and they were in- = 13 (11-15)~tm; excretory pore --- 78 (72- deed females. 114 Journal of Nematology, Volume 8, No. 2, April 1976

In 1952, Volz sent one specimen for 14. Spermatheca ovoid to ovoid-elongate ...... 15. examination which he had identified as P. Spermatheca spherical ...... 17. 15. Female head conical with rounded apex; strenzkei. This was a juvenile with meas- stylet = 48-56 #m ...... urements as follows: L = .19 ram; a = 15; ...... goodeyi (Oostenbrink, 1953). b = 2.4; stylet = 42 t~m; prorhabdion = Female head rounded or a truncated cone; 32 /am. Adhering to the specimen was part stylet >55 #m ...... 16. 16. Female length = .25-.31 mm; styler = 55-65 of the previous cuticle with a molted #m; male length = .28-.34 mm ...... prorhabdion measuring 30 t~m. There is no ...... aonli Misra and Edward, 1971. way of determining whether the juvenile Female length = .38 (.33-.42) ram; stylet = was third- or fourth-stage. 66 (63-70) #rn; male length = .36-.42 mm Geraert (6) probably was correct in ...... pandata n. sp. 17. Posterior edge of annules on females crenate judging this species to be synonymous with ...... crenata (Corbett, 1966). P. straeIeni, but in the absence of desig- Posterior edge of annules on females smooth 18. nated type specimens from the type locality, 18. Female head a truncated cone ...... it is best to assign this species to species ...... marylandica (Jenkins, 1960). inquirendae. Female head rounded ...... 19. 19. Female head smooth; corpus swells markedly in posterior half abietis (Eroshenko, 1974). KEY TO SPECIES OF GRACILACUS Female head with fine annulations; corpus 1. Lateral field with 2 incisures ...... 2. without marked swelling in posterior half 20. Lateral field with 3 incisures ...... 3. 20. V = 73-77; b = 2.8-3.2 ...... Lateral field with 4 incisures ...... 8. .... ivorensis (Luc and de Guiran, 1962). 2. Female stylet = 68-76 /tin ...... V = 79-85; b = 3.0-4.5 ...... peperpotti Schoemaker, 1963...... straeleni (de Coninck, 1931). Female stylet = 100-119 /tin ...... 21. Female tail bluntly rounded ...... elegans Raski, 1962...... epacris (Allen and Jensen, 1950). 3. Lateral vulvar membranes small ...... Female tail subacute to finely rounded ...... 22...... idalima Raski, 1962. 22. Female stylet = 97 (91-102) /am ...... Lateral vulvar membranes lacking ...... 4...... anceps (Cobb, 1923). 4. Female tail bluntly rounded ...... 5. Female stylet (92 /am ...... 23. Female tail subacute to finely rounded or 23. Female tail subacute to rounded; stylet = deformed ...... 6. 89 (83-92) #m .... intermedia Raski, 1962. 5. Female stylet = 58 (48-68) #m; b = 2.7 (2.3- Female tail finely rounded, almost acute; sty- 4.1) ...... aculenta (Brown, 1959). let = 77 (70-85) #m .... mira Raski, 1962. Female stylet = 75 (63-80) /~m; h = 2.2 (1.7- 24. Female stylet = 41-46 #m ...... 2.5) ...... latescens n. sp. micoletzkyi (Edward, Misra and Singh, 1967). 6. Female stylet = 61-69 tun ...... Female stylet ~>63 #m ...... 25...... acicula (Brown, 1959). 25. Female head rounded to conoid but sub- Female stylet ~>69 jam ...... 7. median lobes not set off ...... 26. 7. Female length = .24 (.21-.25) ram; tail Female head with submedian lobes set off or mostly deformed, some subacute ...... protruding ...... 28...... solivaga n. sp. 26. Female stylet = 67 (65-69) #m ...... Female length = .31 (.29-.34) mm; tail sub- ...... steineri (Golden, 1961). acute ...... costata n. sp. Female stylet ~72 /am ...... 27. 8. Lateral vulvar membranes present ...... 9. 27. Female length = .25-.28; excretory pore at Lateral vulvar membranes absent ...... 24. isthmus, opposite nerve ring ...... 9. Female head with submedian lobes or lips ...... oostenbrinki Misra and Edward, 1971. prominent and/or protruding ...... 10. Female length = .33-.39; excretory pore Female head conoid to rounded ...... 12. near knobs o[ styler ...... 10. Female stylet= 92 (86-95) #m ...... macrodora (Brzeski, 1963)...... peratica Raski, 1962. 28. Female stylet = 74 (69-83) /am; tail bluntly Female stylet (60 /~m ...... 11. rounded ...... teres n. sp. 11. Female styler = 48-57 #m; knohs delicate; Female stylet= 94 (82-104) #m; tail acute submedian lobes prominent but not set ...... enata n. sp. off ...... mutabilis (Colbran, 1969). LITERATURE CITED Female stylet = 56-60 #m; knobs strongly developed; submedian lobes set off by 1. ALLEN, M. W., and S. A. SHER. 1967. Tax- slight constriction ...... parwda n. sp. onomic problems concerning the phytopara- 12. Excretory pore at isthmus or more posterior 13. sitie nematodes. Annu. Rev. Phytopathol. 5: Excretory pore near valve of median bulb 247-264. or more anterior ...... 21. 2. BROWN, G. L. 1959. Three new species of the 13. Spermatheca absent; spermagonium present, genus Paratylenchus from Canada (Nema- round ...... robusta Wu, 1974. toda: Criconematidae). Proc. Helminthol. Soc. Spermatheca present ...... 14. Wash. 26:1-8. Revision of Paratylenchus Part III-Gracilacus: Raski 115

3. COLBRAN, R. C. 1969. Studies of plant and 9. OOSTENBRINK, M. 1953. A note on Paraty- soil nematodes. 14. Five new species of Ty- lenchus in the Netherlands with the descrip- lenchorhynchus Cobb, Paratylenchus Mico]et- tion of P. goodeyi n. sp. (Nematoda: Cri- zky, Morulaimus Sauer and Hemicycliophora conematidae). Tijdschr. Plantenziekten. 59: de Man (Nematoda: Tylenchoidea). Queensl. 2O7-216. J. Agric. Anita. Sci. 26:181-192. 10. RASKI, D. J, 1962. Paratylenchidae n. faro. 4. DEMENT'EVA, S. P. 1972. Thornenema uui- with descriptions of live new species of formis, sp. n., and Parat,vIenchus longistylosa, Gracilacus n. g, and an emendation of Ca- sp. n., new species of plant nematodes from copaurus Thorne, 1943, Paratylenchus Mi- the rhizosphere of tobacco. IZV Akad. Nauk. coletzky, 1922 and Criconematidae Thorne, Mold. SSR Ser. Biol. Khim. Nauk. 4:62-64. 1943. Proc, Hehninthol. Soc, Wash. 29:189-207. ,6. EROSHENKO, A. S. 1974. A new nematode 11. SIDDIQI, M. R., and J. B. GOODEY. 1963. The from coniferous [orests in the Far East of status of the genera and subfamilies of the the USSR. Parazitologiya 8:220-222. Criconematidae (Nematoda); with a comment 6. GERAERT, E. 1965. The genus Paratylenchus. on the position of Fergusobia, Nematologica Nematologica 11:301-334. 9:363-377. 7. GOLDEN, A. M. 1971. Classification of the 12. THORNE, (.., and R. B, MALEK. 1968. Nem- genera anti higher categories of the Order atodes of the Northern Great Plains. Part 1. Tylenchida (Nematoda). Pages 191-232, in Tylenchida (Nemata: Secernentea). S. D. B. M. Zuckerman, W. F. Mai and R. A. Agric. Exp. Stn. Tech. Bull. 31:1-111. Rohde, eds. Plant parasitic nematodes. Aca- 13. VOLZ, P. 1951. Untersuchungen iiber die demic Press, New York. Mikrofauna des Waldbodens. Zool. Jahrb. 79 8. MISRA, S. L. and J. C. EDWARD. 1971. Two (5/6):514-566. new species of the genus Paratylenchus with 14, WU, L. Y. 1974. Paratylenchus robustus n, sp. description of their larval stages and a note (Paratyleuchinae: Nematuda) from forest soil on P. nawadus a synonym of P. nainianus. in Ontario. Can. J. Zool. 52:1423-1425. Allahabad Fmr. 45:345-351.