REVISION OF THE GENUS PRIONCHULUS COBB, 1916 (NEMATODA: MONONCHINA). I. PRIONCHULUS MUSCORUM (DUJARDIN, 1845) COBB, 1916 AND RELATED SPECIES

Transcription

1 ANNALES ZOOLOGICI (Warszawa), 2003, 53(3): REVISION OF THE GENUS PRIONCHULUS COBB, 1916 (NEMATODA: MONONCHINA). I. PRIONCHULUS MUSCORUM (DUJARDIN, 1845) COBB, 1916 AND RELATED SPECIES GRAŻYNA WINISZEWSKA 1 and ANDRIJ SUSULOVSKY 2 1 Museum and Institute of Zoology, PAS, Wilcza 64, Warszawa, Poland 2 State Museum of Natural History, Theatralna str. 18, L viv 79008, Ukraine Abstract. This paper deals with six species belonging to the genus Prionchulus Cobb, 1916 which are characterized by having spacious buccal cavity, lower position of dorsal tooth, relatively large and clearly separated subventral denticles, female genital branches without spermatheca and without valvular apparatus between uterus and oviduct. Description of male of P. muscorum (Dujardin, 1845) is given. P. ctenodentatus (Tysowski, 1915) is recognized as a valid species and a neotype is designated. P. fagi Zell, 1985 is redescribed on the basis of the type material. P. zelli nom. nov. is proposed for P. bastiani Zell, 1985, nec Mononchus bastiani De Man, 1876 (syn. P. muscorum (Dujardin, 1845)) and is redescribed from type material and specimens from Ukraine and Poland. P. oxycercus Zell, 1985 is considered a synonym of P. zelli. Two new species: P. ferox sp. nov. and P. olexandri sp. nov. are described and illustrated. The first one is characterized by the relatively small body dimensions; relatively long pharynx; rounded lip region; protruding cephalic and labial papillae; buccal cavity spacious and ovate with funnel-shaped base; lower position of dorsal tooth apex; close arranged sclerotizations, rounded, drop-shaped or arcuate semicircular, with smooth surface; tail conical and slightly ventraly bent; hyaline part of tail conical, longer than wide with slightly rounded tip. P. oleksandri sp. nov. is characterized by a medium size of body; truncate lip region with not protruding cephalic papillae; thick-walled and cylindrical buccal cavity with funnel-shaped base; lower position of dorsal tooth and amphid; well separated sclerotizations of pars refringens vaginae, which are arcuate drop-shaped with smooth surface; conical and regularly tapering tail with cylindrical and weakly rounded tip; well expressed hyaline part of the tail. Key words. Morphology, nematodes, new species, neotype, Prionchulus, taxonomy. INTRODUCTION This is the third paper (Winiszewska 2002, Susulovsky and Winiszewska 2002) in a series of publications on the taxonomy of the species belonging to the genus Prionchulus Cobb, This genus encompass several species related to P. muscsorum (Dujardin, 1845), which are characterized by having spacious buccal cavity, relatively lower position of dorsal tooth and large and clearly separated subventral denticles. Female genital branches are without spermatheca and without valvular apparatus between uterus and oviduct. Detailed study of these species, coming from different regions of Europe showed that a review of this group is needed. In this paper revision of five species: P. muscsorum (Dujardin, 1845), P. ctenodentatus (Tysowski, 1915), P. fagi Zell, 1985, P. bastiani Zell, 1985 and P. oxycercus Zell, 1985 is made, and two new species: P. ferox sp. nov. and P. olexandri sp. nov. are described. MATERIAL AND METHODS Ukrainian specimens used for these descriptions were extracted by modified Baermann funnel method, relaxed by gentle heat and fixed in cold TAF. Polish specimens were extracted by decantation and sieving method with final separation on extraction sieves with filters, fixed in hot 4% formaldehyde. All specimens were processed to pure glycerine by slow evaporation method and mounted on permanent slides in glycerine with paraffin as a support for the cover slide.

2 560 G. WINISZEWSKA and A. SUSULOVSKY TAXONOMY Prionchulus muscorum (Dujardin, 1845) Cobb, 1916 (Figs 1 19) Oncholaimus muscorum Dujardin, 1845: 237; Mononchus muscorum: Bastian 1865: 103; Mononchus (Prionchulus) muscorum: Cobb 1916: 196; Prionchulus muscorum: Wu and Hoeppli 1929: 42; Mononchus bastiani de Man, 1876: 184; syn. by Clark 1960: 194; Prionchulus medius Eroshenko, 1975: 166; syn. by Andrássy 1985: 17. Description. Measurements: see Table 1. Female. Relaxed specimens arcuate more curved ventrad at posterior end. Body tapering towards both ends. Maximum body width at the level of vulva or anterior ovary. Cuticle 3.8±0.5 ( ) µm thick, with faint transverse striae. Cuticular pores small, numerous, arranged along body (excluding tail) in four rows on each side: one subdorsal, one subventral and two sublateral. Each pore is connected with small glandular cell trough the very thin duct. Tail with five pores present on each side: two or three subdorsal, one or two lateral and none or one subventral. Lip region rounded, slightly offset by a depression, almost the same width as adjacent body. Labial and cephalic papillae small, conical, protruding. Cephalic papillae of the same size or somewhat smaller than labial papillae. Amphid cup-like, its aperture 5.9±0.7 ( ) µm wide, located at the level of anterior end of buccal cavity, 14.6±1.0 ( ) µm from head end. Buccal cavity cylindrical, 1.8±0.05 ( ) times as long as wide, with funnel-shaped base. Dorsal wall (at a level of dorsal tooth apex) 2.8±0.5 ( ) µm thick. Dorsal tooth of medium size, its apex located 12.1±0.6 ( ) µm from anterior end of the sclerotized buccal cavity; its upper edge 5.8±0.6 ( ) µm long is perpendicular to the body axis or slightly bent and pointing forward. Subventral denticles, in number, relatively large, forwardly pointing, separated, sometimes arranged irregularly, covering 61.8±4.7 ( )% of the ventral wall. Distance between first and last denticle 23.5±2.2 ( ) µm. Pharynx cylindrical and muscular. Excretory pore weakly marked. Rectum almost straight, 0.8±0.03 ( ) Character Population I Population II Population III n 25 females 2 males 2 males 5 females L (µm) 2710±167 ( ) 2651, , ( ) a 27.9±1.8 ( ) 30.1, , ( ) b 4.2±0.1 ( ) 4.1, , ( ) c 15.5±1.1 ( ) 16.8, , ( ) c 3.1±0.3 ( ) 2.5, , ( ) V (%) 63.7±1.2 ( ) 63.9 ( ) G1 (%) 11.7±1.1 ( ) 13.3 ( ) G2 (%) 11.4±1.1 ( ) 12.4 ( ) Buccal cavity length (µm) 51.6±1.4 ( ) 52.9, , ( ) Buccal cavity width (µm) 29.3±0.8 ( ) 29.2, ,9, ( ) Position of tooth apex (%) 23.5±0.8 ( ) 22.9, 24.0, ( ) Length of ventral wall (µm) 37.9±0.9 ( ) 38.6, , ( ) Excretory pore from anterior end (µm) 222±8.6 ( ) 232, , ( ) Pharynx length (µm) 638±26.3 ( ) 655, , ( ) Lip region height (µm) 12.3±0.9 ( ) 12.8, , ( ) Lip region width (µm) 48.2±2.2 ( ) 47.9, , ( ) Body width at pharynx base (µm) 85.9±7.3 ( ) 78.0, , ( ) Maximum body width (µm) 97.9±9.4 ( ) 88.2, , ( ) Anal body width (µm) 57.0±3.7 ( ) 62.8, , ( ) Vagina length (µm) 40.3±3.2 ( ) 37.3 ( ) Rectum/cloaca length (µm) 47.2±3.3 ( ) 54.5, , ( ) Spicules length (µm) 87.9, , 86.0 Lat. guid. pieces length (µm) 20.6, , 22.1 Tail length (µm) 176±16.2 ( ) 158, , ( ) Hyaline part of tail length (µm) 8.0±1.4 ( ) 5.0, , ( ) Table 1. Morphometrics of Prionchulus muscorum (Dujardin, 1845) from Ukraine (mean ± standard deviation, with range in parentheses, standard deviation was calculated when n>5).

3 PRIONCHULUS MUSCORUM AND RELATED SPECIES Figures 1 8. Prionchulus muscorum (Dujardin, 1845). (1) Entire female, (2) entire male, (3) female anterior region, (4 5) female gonad (anterior), (6) vaginal region, (7, 8) female tails. 1 4, 6 8 Popul. I; 5 Popul. III. Scale bars: 1, 2 = 200 µm; 3, 6 = 20 µm; 4, 5, 7, 8 = 50 µm.

4 562 G. WINISZEWSKA and A. SUSULOVSKY Figures Prionchulus muscorum (Dujardin, 1845). (9) Male posterior region, (10 12) male spicules, (13) aberrant male anterior region, (14 19) diversity of females tail shape in the same population. 9, 10, popul. I; popul. II. Scale bars: 9, = 50 µm; = 20 µm.

5 PRIONCHULUS MUSCORUM AND RELATED SPECIES 563 times the anal body diameter. Tail conical, ventrally bent, its distal 1/3 part more curved ventrad, hookshaped. Terminal part of tail slightly cylindrical, with weakly rounded tip. Hyaline area of tail 1.4±0.3 ( ) times wider than long. In some specimens hyaline part of tail may be swollen or hook-shaped bent subventrally (figs 7, 8). Some spherical glandular bodies present at the proximal part of tail just behind the anus. Genital branches almost symmetrical, anterior branch 317±40.4 ( ) µm, posterior 310±40.5 ( ) µm long. Ovaries on alternate sides of intestine, well developed, anterior ovary 208±27.3 ( ) µm, posterior 215±33.8 ( ) µm long, not reaching the uterus-oviduct junction. Gravid females with maximal 12 oocytes in one row. Oviduct with well marked pars dilatata 58.5±0.8 ( ) µm in width (without egg). Uterus-oviduct junction in the shape of a strong contraction. Uterus with thick walls, in the shape of relatively long, often curled tube. Pars proximalis vaginae longer than wide, with straight walls, surrounded by strong circular muscles, making up 41.3±4.4 ( )% of corresponding body width. Pars refringens vaginae with two very close rounded drop-shaped or semispherical sclerotizations (7.9±0.9 ( ) µm long and 5.0±0.4 ( ) µm wide). These sclerotizations have a characteristic granular structure. Pars distalis vaginae short with rounded walls. Vulva a transverse slit with occasionally protruding lips. Intra-uterine eggs, maximum two in one female, measuring µm (n=6). The eggshell thick ( µm) with shallow folds. Male. General appearance similar to female. Testes full with oval-shaped spermatozoa. Anterior testis 403 µm, posterior 299; 332 µm long, reflexed. Vas deferens and ductus ejaculatorius separated by a constriction. Spicules stout, weakly ventrally curved, with rounded head and bifurcate terminus, times the cloacal body width. Lateral guiding pieces with arcuate edges and bifurcate terminus. This furcation is short and symmetrical. Gubernaculum well developed, bifurcate, µm long. Ventromedian supplements in number, conical and regularly arranged. Between the first supplement and excretory pore there are ventromedian cuticular pores, in number. Tail similar to that of female. A small caudal papilla presents on ventral side of tail somewhat behind the anus. One of the males (fig 13) has the misshapen buccal cavity: dorsal sector is narrower than usually, particularly in upper part; dorsal tooth is malformed, lacking of typical for this species pointed apex; a small additional denticle is located below the basis of dorsal tooth; alone the junction of dorsal and subventral walls short longitudinal strips are visible, each of them have about seven transverse plates, which create denticulate structure. Material examined. 1. Ukraine: (I). Zakarpats ka prov., Vynogradiv distr., N. Klynovo Vill, Julivs ki Mts., hornbeam-oak forest, , leg. A. Susulovsky; (II). Zakarpats ka prov., Vynogradovo, Chorna Gora, oak forest, moist litter, , leg. A. Susulovsky; (III). L viv Prov., Yavoriv Distr., Ivano-Frankove, Roztochya National Reserve, beech-oak forest, litter, , leg. A. Susulovsky; 2. France, forests around Paris (Arpin 1979: Bois de la Tour, Mont-Morin, La Tillaie). Remarks. Our specimens of females correspond well with those described by Clark (1960). They correspond also with specimens from France, which were described in series of publications (Arpin et al. 1984, Samsoen et al. 1984), where the characters used to separate the two related species P. muscorum and P. punctatus were discussed. In two populations of Prionchulus from Ukraine in which the females were recognized by us as P. muscorum, four males were found. Their general appearance is similar to female, however differs essentially from the description given by Mulvey (1967). We consider specimen described by Mulvey as a male of P. muscorum belonged to another species. Diagnosis. P. muscorum is characterized by medium body length; rounded lip region; protruding labial and cephalic papillae of equal size; spacious and cylindrical buccal cavity with funnel-shape base; low position of dorsal tooth; amphid aperture located at a level of anterior end of buccal cavity; slightly cylindrical terminal part of tail, with weakly rounded tip; hyaline part of tail well expressed, longer than wide. Females have a pars proximalis vaginae with straight walls; pars refringens vaginae with two very close rounded drop-shaped or semispherical sclerotizations with characteristic granular structure. Males have stout and weakly ventrally curved spicules; relatively long lateral guiding pieces with arcuate edges and bifurcate terminus; presence of ventromedian cuticular pores between first the supplement and excretory pore. Prionchulus ctenodentatus (Tysowski, 1915) Mulvey, 1967 (Figs 20 28) Mononchus ctenodentatus Tysowski, 1915: 90. Description (population from Poland, 4 females). Measurements: see Table 2. Female. Relaxed specimens arcuate, more curved ventrad at posterior and. Body slightly tapering towards both ends. Maximum body width at a level of vulva. Cuticle smooth, relatively thick (2.6 ( ) µm) especially in posterior part of body; subcuticle with faint transverse striae, more prominent in caudal region. Cuticular pores of medium size, not numerous, arranged along the body (excluding tail) in four rows on each side: one subdorsal, one subventral and two sublateral. In pharyngeal region they could be found more densely than in a rest of the body. Tail with one pair of subventral, one pair of sublateral and one or two pairs of subdorsal pores. One pair of subdorsal pores always

6 564 G. WINISZEWSKA and A. SUSULOVSKY Character Ukraine original description (Tysowski 1915) Neotype Poland The same population as neotype France n 1 female 3 females 10 females L (µm) ( ) 2256±107 ( ) a ( ) 24.8±1.4 ( ) b ~ ( ) 4.2±0.2 ( ) c ( ) 19.3±1.7 ( ) c ( ) 2.4±0.2 ( ) V (%) ( ) 65.0±1.2 ( ) G1 (%) ( ) 13.4±1.5 ( ) G2 (%) ( ) 12.2±2.1 ( ) Buccal cavity length (µm) ( ) 44.5±1.9 ( ) Buccal cavity width (µm) ( ) 26.1±0.7 ( ) Position of tooth apex (%) ( ) 23.2±1.0 ( ) Length of subventral ridge (µm) ( ) 34.0±1.8 ( ) Excretory pore from anterior end (µm) ( ) Pharynx length (µm) ( ) 532±18.4 ( ) Lip region height (µm) ( ) 7.8±0.9 ( ) Lip region width (µm) ( ) 38.3±1.8 ( ) Body width at pharynx base (µm) ( ) 74.5±3.5 ( ) Maximum body width (µm) ( ) 91.1±5.0 ( ) Anal body width (µm) ( ) 48.7±2.5 ( ) Vagina length (µm) ( ) 30.4±1.5 ( ) Rectum length (µm) ( ) 37.5±2.8 ( ) Tail length (µm) ( ) 117±7.5 ( ) Table 2. Morphometrics of females of Prionchulus ctenodentatus (Tysowski, 1915) (mean ± standard deviation, with range in parentheses). opens into extremely big, elongated, glandular cells lying in the proximal part of the tail. Lip region offset by a weak depression. Head rounded. Labial and cephalic papillae conical, broadly rounded, slightly protruding beyond the body outline. Cephalic papillae of the same size as labial ones or somewhat more prominent. Amphid cup-like, relatively small, its aperture 4.9 ( ) µm wide, located at a level of dorsal tooth apex, 18.1 ( ) µm from head end. Buccal cavity wide, barrel-shaped or cylindrical, 1.8 ( ) times as long as wide, with nearly flattened base. Dorsal wall (at a level of dorsal tooth apex) 1.9 ( ) µm thick. Dorsal tooth of medium size, its apex located 10.7 ( ) µm from anterior end of the sclerotized buccal cavity, its upper edge, 4.7 ( ) µm long, little bent and pointing forward. Subventral denticles in number, of medium size, arranged mainly regularly and very closely, but clearly separated from each other, pointed forwards, covering 53.1 ( )% of the ventral wall. Distance between first and last denticle 19.9 ( ) µm. Pharynx cylindrical and muscular. Excretory pore weakly marked. Rectum with thick walls, almost straight, 0.9 ( ) times the anal body diameter. Tail plump, ventrally bent, regularly tapering. Tail terminus bent ventrad with broadly rounded tip. Hyaline part of tail well expressed, somewhat wider then long. The border between cytoplazmatic and hyaline part visible as right line, perpendicular to body axis. Genital branches almost symmetrical, anterior branch 224 ( ) µm, posterior 222 ( ) µm long. Ovaries on alternate sides of intestine, well developed, anterior ovary 146 ( ) µm, posterior 152 ( ) µm long, not reaching the uterus-oviduct junction. Gravid females with maximum 9 oocytes in one row. Oviduct very short, pars dilatata 43.8 ( ) µm in width (without egg). Uterus-oviduct junction weakly marked. Uterus with thick walls, straight and very short (48.6 ( ) µm long). Pars proximalis vaginae longer than wide, with straight contours and almost parallel walls, making up 36.8 ( )% of corresponding body width. Pars proximalis vaginae and pars refringens vaginae are surrounded by strong circular muscles. Pars refringens vaginae with two, very weakly refractive sclerotizations (4.8 ( ) µm long and 3.8 ( ) µm wide). Pars distalis vaginae relatively long (6.6 ( ) µm), thick walled. Vulva a transverse slit lying in a very small depression. Male. Unknown.

7 PRIONCHULUS MUSCORUM AND RELATED SPECIES Figures Prionchulus ctenodentatus (Tysowski, 1915). (20, 28) Female anterior region, (21 23, 27) female tails, (24) vaginal region, (25) entire female, (26) female genital system. 20, 21, neotype female; 22 female from the same population as neotype; 23 popul. from France; 27, 28 after Tysowski Scale bars: 20, 24 = 20 µm; 21 23, 26 = 50 µm, 25 = 200 µm.

8 566 G. WINISZEWSKA and A. SUSULOVSKY Population from France (10 females). Specimens of this population in general appearance are similar to those described above. They differ in somewhat smaller and less numerous cuticular pores, arrangement of pores on tail (more laterad). Intra-uterine eggs oval with smooth shell, measuring µm. Remarks. Mononchus ctenodentatus was described by Tysowski (1915) on the basis of several females (in description he used plural form: mature specimens appeared in April ) found in moss on roof in village Poturzyca (at present Ukraine). The original description is very laconic. It contains the basic dimensions (see table 2) as well as drawings of head region and tail (see figs 27, 28). After the first description this species has not been noted once again. Andrássy (1958) in revision of Mononchidae recognized P. ctenodentatus as a species with uncertain status. Mulvey (1967) considered it as synonym of P. muscorum. Specimens, which have been found in Poland and in France, fit the description and drawings given by Tysowski. Since no fresh specimens of this species were found in the rich material collected in Poturzyca in spring 2002, we have decided to designate the neotype from among females, which were found in eastern part of Poland. Type designation. On the basis of Art. 75 of the International Code of Zoological Nomenclature a female specimen from Poland is herewith designated as neotype. Neotype and two females are deposited at the Museum and Institute of Zoology PAS, Warszawa one female at the State Museum of Natural History, L viv. Neotype locality and habitat. Poland, Orły, moos on the roof, , leg. A. Słojewska. Other locality. France, les Guichards, Grimmia pulvinata (Hedw.) Sm., 10.VII.1977, slide no Diagnosis. P. ctenodentatus is characterized by medium body length; broadly rounded labial region; spacious and barrel-shaped or cylindrical buccal cavity with almost flattened base; lower position of dorsal tooth; amphid aperture located at a level of dorsal tooth apex; plump tail with broadly rounded tip, with extremely big, elongated, glandular cells lying in its proximal part; short oviduct and uterus; pars refringes vaginae with very weakly refractive sclerotizations; pars distalis vaginae relatively long and thick walled. Prionchulus fagi Zell, 1985: 66. Prionchulus fagi Zell, 1985 (Figs 29 38) Description. Measurements: see Table 3. Female. Relaxed specimens arcuate more curved ventrad at posterior end. Body tapering towards both ends. Cuticle 2 3 µm thick, with faint transverse striae. Character P. fagi P. bastiani P. oxycercus Holotype Paratypes Holotype Paratypes Holotype Paratypes n 1 female 6 females 1 male 1 female 14 females 1 female 3 females L (µm) a b c c 2.9* 2.9* 2.5* 2.4* 2.7* V (%) Buccal cavity length (µm) 55* * 55* Position of tooth apex (%) 24* 23 25* 24* Length of subventral ridge (µm) 43* 42* 41* 44* 44* Excretory pore from anterior end (µm) 242* 207* Pharynx length (µm) 685* * 570* Lip region height (µm) 12* 12* 13* 11* 10* Lip region width (µm) 56* 50* 49* 50* 53* Vagina length (µm) 50* 48* 47* Rectum/cloaca length (µm) 50* 50* 56* 45* 50* Spicules length (µm) 114 Lat. guid. pieces length (µm) 29 Tail length (µm) Hyaline part of tail length (µm) 9* 10* 9* 9* 8* Table 3. Morphometrics of Prionchulus fagi Zell, 1985, P. bastiani Zell, 1985 and P. oxycercus Zell, 1985 from original descriptions (Zell 1985); with (*) dimensions added after examination of type material were marked.

9 PRIONCHULUS MUSCORUM AND RELATED SPECIES Figures Prionchulus fagi Zell, (29) Entire female, (30) female buccal cavity, (31) female anterior region, (32) vaginal region, (33) female tail, (34) terminal part of female tail. 29, holotype female, 30 paratype female, (35) entire male, (36) male anterior region, (37) male posterior region, (38) male spicule, gubernaculum and lateral guiding piece paratype male. Scale bars: 29, 35 = 500 µm, 30-32, 34, 36, 38 = 20 µm, 33, 37 = 50 µm.

10 568 G. WINISZEWSKA and A. SUSULOVSKY Lip region flattened in outline, wider than adjacent body. Labial and cephalic papillae relatively large, conical, protruding, located almost at the same level. Cephalic papillae bigger than labial ones. Amphid cup-like, its aperture about 5 µm wide, located at a level of anterior end of buccal cavity, µm from head end. Buccal cavity thinwalled, ovate, with almost flat base. Dorsal wall at a level of dorsal tooth apex, 3 µm thick, below the tooth narrower (2 µm thick). Dorsal tooth of medium size, its apex located µm from anterior end of the sclerotized buccal cavity; its upper edge 7 µm long, perpendicular to the body axis. Subventral denticles 14 in number, medium sized, forwardly pointing, separated, sometimes arranged irregularly, covering 53 59% of the ventral wall. Distance between first and last denticle µm. Pharynx cylindrical and muscular. Excretory pore weakly marked. Rectum almost straight. Tail conical, almost straight, only its distal 1 /3 part weakly curved ventrad. Terminal part of tail slightly cylindrical, with weakly rounded tip. Hyaline area of tail relatively long. Anterior genital branch µm, posterior µm long. Ovaries on alternate sides of intestine, well developed, with maximum 20 oocytes in one row; anterior ovary µm, posterior µm long, not reaching the uterus-oviduct junction. Oviduct with well marked pars dilatata. Pars proximalis vaginae longer than wide, with triangular contours, surrounded by strong circular muscles. Pars refringens vaginae with two very close drop-shaped to rounded drop-shaped sclerotizations. Vulva a transverse slit. Intra-uterine eggs rounded, maximum three in one female, measuring µm. The eggshell thick with shallow folds. Male. General appearance similar to female. Testes full with spermatozoa. Anterior testis 260 µm, posterior 297 µm long, reflexed. Vas deferens and ductus ejaculatorius separated by a constriction. Spicules stout ventrally curved, with rounded head and bifurcate terminus, 1.8 times the cloacal body width. Lateral guiding pieces with arcuate edges and bifurcate terminus. This furcation is short and symmetrical. Gubernaculum well developed, bifurcate. Ventromedian supplements 16 in number, conical and regularly arranged. Tail similar to that of female. A small caudal papilla presents on ventral side of tail somewhat behind the anus. Remarks. Type material is flattened; internal structures are faintly visible. However a few characters can be used to separate this species from other representatives of Prionchulus. After the detailed study of those characters a new diagnosis for this species has been proposed. Material examined. Holotype female (slide QHA-F (1)), paratype female and paratype male (slide: QHA- L-1178 (3)), Landessammlungen für Naturkunde, Karlsruhe, Germany. Diagnosis. P. fagi is characterized by large body length; oblate lip region; cephalic papillae bigger than labial ones; spacious and oval buccal cavity, with almost flat base and relatively thin walls; shape of tail which is almost straight in first 2 /3 length; long and massive spicules. Prionchulus zelli nom. nov. (Figs 39 60) Prionchulus bastiani Zell, 1985: 64, nec Mononchus bastiani De Man, 1876 (see remarks below); Prionchulus oxycercus Zell, 1985: 63. syn. nov. Description. Measurements: see Table 3 and 4. Female. Relaxed specimens arcuate more curved ventrad at posterior end. Body tapering towards both ends. Maximum body width at a level of vulva. Cuticle 3.7±0.6 ( ) µm thick, with faint transverse striae more visible in caudal region, subcuticule is also striated. Cuticular pores very small, numerous, arranged along body (excluding tail) in four rows on each side: one subdorsal, one subventral and two sublateral; between base of buccal cavity and nerve ring are two lateral pores. Each pore is connected with small vacuolated cell trough the very thin duct. Tail with five pores present on each side: two subdorsal, one or two lateral and one subventral. Lip region rounded, slightly offset by a depression, almost the same width as adjacent body. Labial and cephalic papillae of the same size, small, conical, protruding. Amphid cup-like, its aperture 6.1±0.8 ( ) µm wide, located at a level of anterior end of buccal cavity, 14.5±1.0 ( ) µm from head end. Buccal cavity thick-walled, 1.7±0.06 ( ) times as long as wide, with funnel-shaped base, wide and oval in upper part, below to basis of dorsal tooth become narrower and cylindrical. Dorsal wall (at a level of dorsal tooth apex) 2.7±0.4 ( ) µm thick. Dorsal tooth of medium size, its apex located 13.9±0.8 ( ) µm from anterior end of the sclerotized buccal cavity, its upper edge 6.1±0.6 ( ) µm long, little bent and pointing forward. Subventral denticles 11 in number, relatively large, pointing forward, separated, sometimes arranged irregularly, covering 59.7±4.5 ( )% of the ventral wall. Distance between first and last denticle 24.4±1.6 ( ) µm. Pharynx cylindrical and muscular. Excretory pore weakly marked. Rectum thick-walled, almost straight, 0.7±0.08 ( ) times the anal body diameter. Tail conical, ventrally bent, regularly tapering. Tail terminus narrow, elongated, slightly cylindrical or conical, with weakly rounded tip, in many cases bent dorsally. Hyaline part of tail very unclear and faintly visible, the cytoplasmatic part forming a thin filamentous structure, which attains almost the tail terminus. Genital branches almost symmetrical, anterior branch 286±45.0 ( ) µm, posterior 272±48.1( ) µm long. Ovaries, on alternate sides of intestine, well developed, anterior ovary 182±40.4 ( ) µm, posterior 180±47.4 ( ) µm long, not reaching the uterusoviduct junction. Gravid females with maximum 15

11 PRIONCHULUS MUSCORUM AND RELATED SPECIES 569 Character Ukraine Population I Population II Population III Germany (type locality) Poland Czech Republic n 11 females 9 females 6 females 4 females 3 females 1 female L (µm) a b c c V (%) G1 (%) G2 (%) Buccal cavity length (µm) Buccal cavity width (µm) Position of tooth apex (%) Length of ventral wall (µm) Excretory pore from anterior end (µm) Pharynx length (µm) Lip region height (µm) Lip region width (µm) Body width at pharynx base (µm) Maximum body width (µm) Anal body width (µm) Vagina length (µm) Rectum length (µm) Tail length (µm) 2575±176 ( ) 29.4±2.3 ( ) 4.1±0.2 ( ) 16.0±1.0 ( ) 2.9±0.2 ( ) 64.6±1.3 ( ) 10.2±1.5 ( ) 10.3±1.8 ( ) 53.7±1.4 ( ) 31.0±1.1 ( ) 25.9±1.3 ( ) 40.9±1.3 ( ) 211±9.0 ( ) 632±29.7 ( ) 12.0±1.4 ( ) 49.5±1.8 ( ) 80.7±6.9 ( ) 88.3±10.5 ( ) 56.4±5.4 ( ) 37.4±3.1 ( ) 42.1±4.3 ( ) 162±15.1 ( ) 2449±179 ( ) 29.1±1.4 ( ) 4.0±0.1 ( ) 15.3±0.6 ( ) 3.1±0.2 ( ) 64.9±1.0 ( ) 9.1±0.5 ( ) 8.3±0.6 ( ) 52.5±1.8 ( ) 31.8±0.9 ( ) 26.0±1.0 ( ) 39.5±1.2 ( ) 203±6.9 ( ) 608±32.4 ( ) 12.2±0.6 ( ) 47.4±1.1 ( ) 76.6±6.3 ( ) 84.3±7.3 ( ) 51.9±4.0 ( ) 45.5±2.8 ( ) 43.2±2.8 ( ) 160±14.0 ( ) 2703±184 ( ) 24.2±0.7 ( ) 4.0±0.1 ( ) 15.0±0.9 ( ) 3.1±0.1 ( ) 62.8±1.3 ( ) 12.4±1.0 ( ) 12.8±0.9 ( ) 55.0±0.7 ( ) 30.2±0.6 ( ) 25.3±0.9 ( ) 40.2±0.9 ( ) 230±6.4 ( ) 670±28.5 ( ) 11.5±0.5 ( ) 51.8±1.4 ( ) 99,2±8.4 ( ) 111.8±9.7 ( ) 59.3±3.5 ( ) 42.6±3.4 ( ) 48.3±2.2 ( ) 181±18.6 ( ) 2574 ( ) 24.7 ( ) 2800 ( ) 25.0 ( ) ( ) 4.2 ( ) ( ) 14.4 ( ) ( ) 3.3 ( ) ( ) 13.1 ( ) 13.8 ( ) 55.8 ( ) 31.5 ( ) 23.7 ( ) 41.4 ( ) 208 ( ) 610 ( ) 18.0 ( ) 48.7 ( ) 89.1 ( ) ( ) 57.8 ( ) 37.1 ( ) 47.7 ( ) 180 ( ) 63.5 ( ) 12.4 ( ) 13.2 ( ) 54.0 ( ) 31.0 ( ) 23.1 ( ) 39.0 ( ) 229 ( ) 671 ( ) 15.9 ( ) 51.4 ( ) 96.8 ( ) ( ) 59.6 ( ) 37.8 ( ) 49.3 ( ) 194 ( ) Table 4. Morphometrics of Prionchulus zelli nom. nov. (mean ± standard deviation, with range in parentheses, standard deviation was calculated when n>5) oocytes in one row. Oviduct with very well marked pars dilatata 53.0±11.0 ( ) µm in width (without egg). Uterus-oviduct junction looks like of strong contraction, surrounded by pars dilatata. Uterus with thick walls, in shape of relatively short, often curled tube. Pars proximalis vaginae longer than wide, with more or less sigmoid contours, surrounded by strong circular muscles, making up 42.8±4.9 ( )% of corresponding body diameter. Pars refringens vaginae with two relatively large and close rounded triangular or trapezoid sclerotizations (8.6±1.1 ( ) µm long and 5.7±0.7 ( ) µm wide. Pars distalis vaginae well expressed, with rounded walls. Vulva a transverse slit. In most females vulval lips are asymmetrical, one of them is swollen and protruding. In such

12 570 G. WINISZEWSKA and A. SUSULOVSKY Figures Prionchulus zelli nom. nov. (39) Entire female, (40) female anterior region, (41) female gonad (anterior), (42) vaginal region, (43) female tail. 39, 41, 43 holotype female of Prionchulus bastiani Zell, 1985, 40, 42 paratype female of P. bastiani, (44) female buccal cavity, (45) vaginal region, (46) female tail paratype female of Prionchulus oxycercus Zell, Scale bars: 39 = 500 µm, 40, 42, 44, 45 = 20 µm, 41, 43, 46 = 50 µm. case sclerotizations shift in relation to one another, contour of pars proximalis vaginae curves and musculature of vaginae grows asymmetrical. Male. Not found. Material examined. Type specimens: Holotype female (slide: QHA-L-979 (3)) and paratype female (slide: QHA-L-977 (3)), Landessammlungen für Naturkunde, Karlsruhe, Germany. Type material is flattened; internal structures are faintly visible. Other material. 1. Germany. 4 females from type locality (internal structures are faintly visible); 2 Ukraine: (I). Zakarpats ka prov., Peretchyn distr., Turia Poliana village, Rivna Mountains, the Carpathians, beech forest, dust under the bark of dead beech, , leg. A. Susulovsky; (II). Zakarpats ka prov., Volovets distr., Bukovets village, Beskydy Mountains, the Carpathians, beech forest, June 1991, leg. A. Sususlovsky; (III). L viv Prov., Yavoriv Distr., Ivano- Frankove, Roztochya National Reserve, beech forest, litter, , leg. A. Susulovsky; 3. Poland, Łomna Las (Warsaw neighbourhood), Sphagnum sp., , leg. G. Winiszewska; 4. Czech Republic, Boh. C. 5950, Benešŭv luh. 1, Carpinus/Mel. Nem.-Quercetum, , leg L. Hánĕl. Diagnosis. P. zelli. is characterized by a medium body length; weakly rounded lip region; buccal cavity spacious and oval in its anterior part and cylindrical below the dorsal tooth; low position of dorsal tooth apex;

13 PRIONCHULUS MUSCORUM AND RELATED SPECIES Figures Prionchulus zelli nom. nov. (47) Entire female, (48) female gonad (posterior), (49 51) vaginal regions, different position of sclerotizations popul. (I) from Ukraine. Scale bars: 47 = 200 µm; 48 = 50 µm; = 20 µm. sigmoid shape of pars proximalis vaginae; large and angular sclerotization; lack of the hyaline part of tail and tail terminus having the tendency to bend dorsally. Remarks. Zell (1985) described new species using specific name, which was preoccupied by Mononchus bastiani after its synonymization with Prionchulus muscorum by Clark (1960). On the basis of Art. 60 of the International Code of Zoological Nomenclature we propose the name P. zelli for this species. Detailed study of paratype female P. oxycercus (slide: QHA-H-180 (2)) has been done. This species cannot be differentiated from P. zelli nom. nov. and is therefore synonymized with it. Prionchulus ferox sp. nov. (Figs 61 69) Description. Measurements: see Table 5. Female. Relaxed specimens arcuate more curved ventrad at posterior end. Body tapering towards both ends. Maximum body diameter at a level of vulva. Cuticle

14 572 G. WINISZEWSKA and A. SUSULOVSKY Figures Prionchulus zelli nom. nov. (52) Female anterior region, (53 60) diversity of females tail shape popul. from Ukraine I; popul. from Germany; 60 popul. from Poland. Scale bars: 52 = 200 µm; = 50 µm.

15 PRIONCHULUS MUSCORUM AND RELATED SPECIES ±0.4 ( ) µm thick, with faint transverse striae more visible in caudal region, subcuticule with longitudinal lines. Cuticular pores very small, not numerous, arranged along the body (excluding tail) in two rows on each side: one subdorsal and one subventral. Tail with five pores present on each side: two subdorsal, two lateral and one subventral. Lip region rounded, slightly offset by a depression, almost the same width as adjacent body. Labial and cephalic papillae conical, protruding. Cephalic papillae bigger than labial. Amphid cup-like, its aperture 4.3±0.7 ( ) µm wide, located approximately half way between the anterior end of buccal cavity and tooth apex, 16.3±1.1 ( ) µm from head end. Buccal cavity thin-walled, ovate, 1.7±0.04 ( ) times as long as wide, with funnel-shaped base. Dorsal wall (at a level of dorsal tooth apex) 2.4±0.5 ( ) µm thick. Dorsal tooth of medium size, its apex located 10.0±0.9 ( ) µm from anterior end of the sclerotized buccal cavity, its upper edge, 5.4±0.5 ( ) µm long, little bent and pointing forward or perpendicular to the body axis. Subventral denticles, in number, relatively large, pointing forward, separated, arranged almost regularly, covering 62.8±3.5 ( )% of the Character Holotype Paratypes n 1 female 24 females L (µm) ±150 ( ) a ±2.5 ( ) b ±0.13 ( ) c ±0.8 ( ) c ±0.2 ( ) V (%) ±1.2 ( ) G1 (%) ±1.2 ( ) G2 (%) ±1.3 ( ) Buccal cavity length (µm) ±1.2 ( ) Buccal cavity width (µm) ±0.8 ( ) Position of tooth apex (%) ±1.1 ( ) Length of ventral wall (µm) ±1.4 ( ) Excretory pore from anterior end (µm) ±10 ( ) Pharynx length (µm) ±29 ( ) Lip region height (µm) ±1.0 ( ) Lip region width (µm) ±1.6 ( ) Body width at pharynx base (µm) ±5.4 ( ) Maximum body width (µm) ±7.8 ( ) Anal body width (µm) ±2.3 ( ) Vagina length (µm) ±2.7 ( ) Rectum length (µm) ±2.4 ( ) Tail length (µm) ±12 ( ) Hyaline part of tail length (µm) ±1.0 ( ) Table 5. Morphometrics of Prionchulus ferox sp. nov. from Russia (mean ± standard deviation, with range in parentheses). ventral wall. Distance between first and last denticle 21.7±1.5 ( ) µm. Pharynx cylindrical and muscular. Excretory pore weakly marked. Rectum thickwalled, almost straight, 0.7±0.04 ( ) times of the anal body diameter. Tail conical, slightly ventrally bent, regularly tapering. Tail terminus narrow, elongated, slightly conical, with weakly rounded tip. Hyaline area of tail 1.8±0.2 ( ) times longer than wide. Some spherical glandular bodies present at the base of tail just behind the anus. Genital branches almost symmetrical, anterior branch 216±29.7 ( ) µm, posterior 217±29.7 ( ) µm long. Ovaries, on alternate sides of intestine, well developed, anterior ovary 129±21.3 ( ) µm, posterior 137±30.0 ( ) µm long, not reaching the uterus-oviduct junction. Gravid females with maximum 9 oocytes in one row. Oviduct with very well marked pars dilatata 55.4±8.6 ( ) µm in width (without egg). Uterus-oviduct junction looks like narrow, elongated tube, partly surrounded by pars dilatata. Uterus with thick walls, in shape of relatively short, often curled tube. Pars proximalis vaginae longer than wide, with straight contours, surrounded by strong circular muscles, making up 38.0±4.4 ( )% of corresponding body width. Pars refringens vaginae with two very close rounded dropshaped or arcuate semicircular sclerotizations (7.2±0.8 ( ) µm long and 4.8±0.7 ( ) µm wide), with smooth surface. Pars distalis vaginae short with rounded walls. Vulva a transverse slit. Egg (in pars dilatata) oval with thin (2.1 µm) and smooth shell, measures µm. Male. Not found. Type locality. Russia, Rep. Gornyi Altai, vil. Tungur, near Belukha Mt., moraine in front of glacier, soil under bush, , collected by I. Dykyy. Type material. Holotype female and eleven paratype females on permanent mounts in glycerine, deposited in the nematode collection of the State Museum of Natural History, L viv, Ukraine; six females at the Museum and Institute of Zoology, PAS, Warszawa, Poland; three females at the Nematology Department of Wageningen University, The Netherlands; two females at the Instituut voor Dierkunde, Universiteit Gent, Belgium; two females at United States Department of Agriculture Nematode Collection, Beltsville, U.S.A.. Diagnosis and relationships. P. ferox sp. nov. is characterized by the relatively small body dimensions; relatively long pharynx; rounded lip region; protruding cephalic and labial papillae; buccal cavity spacious

16 574 G. WINISZEWSKA and A. SUSULOVSKY Figures Prionchulus ferox sp. nov. (61) Entire female, (62) female anterior region, (63) female genital system, (64) vaginal region, (65 69) diversity of females tail shape holotype female; paratype females. Scale bars: 61 = 200 µm; 62, 64 = 20 µm; 63, = 50 µm.

17 PRIONCHULUS MUSCORUM AND RELATED SPECIES 575 and ovate with funnel-shaped base; lower position of dorsal tooth apex; close arranged sclerotizations, rounded, drop-shaped or arcuate semicircular, with smooth surface; tail conical and slightly ventraly bent; hyaline part of tail conical, longer than wide with slightly rounded tip. A combination of the above mentioned characters separates P. ferox sp. nov. from most species in the genus. In general appearance it is most similar to P. muscorum from which is distinguishes by having shorter body (L = vs L = ), relatively longer pharynx (b = vs b = ), lower position of amphid, structure of surface of sclerotizations (smooth vs granular), the shape of tail (weakly curved ventraly vs strongly curved ventraly) and the shape of the hyaline part of tail (conical and slightly rounded vs cylindrical and more broadly rounded). Etymology. This species was named in honour of the collector and our colleague, Ihor Dykyy, whose Ukrainian family name is equal to English wild or Latin ferox, acknowledging his kindness when collecting samples in Altai Mountains. Prionchulus oleksandri sp. nov. (Figs 70 78) pointing forwards, separated, sometimes arranged irregularly, covering 59.0±5.6 ( )% of the ventral wall. Distance between first and last denticle 21.3±1.9 ( ) µm. Pharynx cylindrical and muscular. Excretory pore weakly marked. Rectum thick-walled, almost straight, 0.8±0.05 ( ) times the anal body diameter. Tail conical, ventrally bent, regularly tapering, its distal part more curved. Tail terminus narrow and elongated, slightly cylindrical, with weakly rounded tip. Hyaline part of tail 1.4±0.2 ( ) times longer than wide. Some spherical glandular bodies present at the base of tail just behind the anus. Genital branches almost symmetrical, anterior branch 301±37.0 ( ) µm, posterior 296±34.5 ( ) µm long. Ovaries, on alternate sides of intestine, well developed, anterior ovary 176±28.3 ( ) µm, posterior 186±28.3 ( ) µm long, not reaching the uterusoviduct junction. Gravid females with maximum 11 oocytes in one row. Oviduct with very well marked pars dilatata 48.2±7.5 ( ) µm in width (without egg). Uterus-oviduct junction looks like a strong contraction. Uterus with thick walls, in shape of relatively long, often curled tube. Pars proximalis vaginae longer than wide, Character Holotype Paratypes Description. Measurements: see Table 6. Female. Anterior part of body in relaxed specimens straight, growing arcuate below the base of pharynx, more curved ventrad at posterior end. Body tapering towards both ends. Maximum body width at a level of vulva. Cuticle 3.1±0.4 ( ) µm thick, with faint transverse striae. Cuticular pores very small, numerous, scattered on the body surface. Each pore connected with big vacuolated cell through the very thin duct. Tail with five pores present on each side: three subdorsal, one lateral and one subventral. Lip region truncate, almost the same width as adjacent body. Labial and cephalic papillae small and conical. Only labial papillae protruding. Amphid cup-like, its aperture 5.2±0.8 ( ) µm wide, located approximately half way between the anterior end of buccal cavity and dorsal tooth apex, 16.3±1.0 ( ) µm from head end. Buccal cavity massive, thickwalled, cylindrical, 1.7±0.1 ( ) times as long as wide, with funnel-shaped base. Dorsal wall (at a level of dorsal tooth apex) 2.2±0.3 ( ) µm thick. Dorsal tooth of medium size, its apex located 11.3±0.7 ( ) µm from anterior end of the sclerotized buccal cavity; its upper edge 5.1±0.5 ( ) µm long is little bent and pointing forward. Subventral denticles in number, of middle size, n 1 female 21 females L (µm) ±104 ( ) a ±2.1 ( ) b ±0.7 ( ) c ±0.8 ( ) c ±0.3 ( ) V (%) ±1.4 ( ) G1 (%) ±1.4 ( ) G2 (%) ±1.4 ( ) Buccal cavity length (µm) ±0.9 ( ) Buccal cavity width (µm) ±1.1 ( ) Position of tooth apex (%) ±1.3 ( ) Length of ventral wall (µm) ±1.2 ( ) Excretory pore from anterior end (µm) ±5.9 ( ) Pharynx length (µm) ±15.3 ( ) Lip region height (µm) ±1.1 ( ) Lip region width (µm) ±1.7 ( ) Body width at pharynx base (µm) ±4.3 ( ) Maximum body width (µm) ±6.9 ( ) Anal body width (µm) ±2.8 ( ) Vagina length (µm) ±2.6 ( ) Rectum length (µm) ±1.9 ( ) Tail length (µm) ±7.5 ( ) Hyaline part of tail length (µm) ±0.9 ( ) Table 6. Morphometrics of Prionchulus oleksandri sp. nov. from Ukraine (mean ± standard deviation, with range in parentheses).

18 576 G. WINISZEWSKA and A. SUSULOVSKY Figures Prionchulus oleksandri sp. nov. (70) Entire female, (71) female anterior region, (72) female genital system, (73) vaginal region, (74 78) diversity of females tail shape holotype female; paratype females. Scale bars: 70 = 200 µm; 71, 73 = 20 µm; 72, = 50 µm.

19 PRIONCHULUS MUSCORUM AND RELATED SPECIES 577 with straight or slightly sigmoid contours, surrounded by strong circular muscles, making up 40.3±3.2 ( )% of corresponding body width. Pars refringens vaginae with two separated arcuate drop-shaped sclerotizations (6.7±0.6 ( ) µm long and 4.5±0.6 ( µm wide) with smooth surface. Pars distalis vaginae short with rounded walls. Vulva a transverse slit. Eggs in genital branches oval in shape, measuring µm (n=15) with smooth egg shell ( µm). Male. Not found. Type locality. Ukraine, Jalta District, The Crimea Peninsula, Ai-Petri Mountain; moss, July 1998, leg. O. Holovachov. Type material. Holotype female and ten paratype females on permanent mounts in glycerine, deposited in the nematode collection of the State Museum of Natural History, L viv, Ukraine; four females at the Museum and Institute of Zoology, PAS, Warszawa, Poland; three females at the Nematology Department of Wageningen University, Wageningen, The Netherlands; two females at the Instituut voor Dierkunde, Universiteit Gent, Belgium; two females at United States Department of Agriculture Nematode Collection, Beltsville, U.S.A. Diagnosis and relationships. P. oleksandri sp. nov. is characterized by a medium size of body; truncate lip region with not protruding cephalic papillae; thick-walled and cylindrical buccal cavity with funnel-shaped base; lower position of dorsal tooth and amphid; well separated sclerotizations of pars refringens vaginae, which are arcuate drop-shaped with smooth surface; conical and regularly tapering tail with cylindrical and weakly rounded tip; well expressed hyaline part of the tail. A combination of the above characters, especially lip and vaginal regions structure, separates P. olexandri sp. nov. from all other species in the genus. Etymology. The species is named after our friend Oleksandr Holovachov who collected the specimens. ACKNOWLEDGEMENTS We thank Dr. H. Zell for lending the type specimens of P. fagi, P. bastiani and P. oxycercus; Prof. P. Arpin, Dr. P. Baujard and Dr. L. Hánél for putting the specimens of Prionchulus from their collection at our disposal. Constructive referees criticism is also acknowledged. The paper was partly sponsored by the State Committee for Scientific Research grant no. 6 PO4C (G. Wniszewska). REFERENCES Andrássy, I Über das System der Mononchiden (Mononchidae Chitwood, 1937; Nematoda). Annales Historico- Naturales Musei Nationalis Hungarici, (s. n. 9) 50: Andrássy, I On the genera Mononchus Bastian, 1865 and Prionchulus (Cobb, 1916) Wu & Hoeppli, 1929 (Nematoda: Mononchidae). Opuscula Zoologica Universitatis Budapestinensis, 21: Arpin, P Ecologie et systématique des nématodes Mononchides des zones forestières et herbacées sous climat tempéré humide. I. Types de sol et groupements spècifiques. Revue de Nématologie, 2: Arpin, P., Samsoen, L., Ponge, J-F. and S. H. Khan Ecology and systematics of the Mononchid nematodes from wood-and grassland areas in wet temperate climate. II. The genus Prionchulus Cobb, Revue de Nématologie, 7: Bastian, H.C Monograph on the Anguillulidae, or free nematoids, marine, land, and freshwater; with descriptions of 100 new species. Transactions of the Linnean Society of London, 25: , pls Clark, W. C Redescription of Mononchus truncatus Bastian, M. papillatus Bastian and Prionchulus muscorum (Dujardin) (Enoplida, Nematoda). Nematologica, 5: Cobb, N.A Subdivisions of Mononchus. Journal of Parasitology, 2: Dujardin, F Histoire naturelle des helminthes ou versintestinaux. Paris, 648 pp., 22 pls. Eroshenko, A.S Desjat novych vidov nematod otrjada Mononchida Jairajpuri, 1969, iz chvojnych lesov Primorskogo Kraja. Gelmintologičeskie Issledovanija Životnych i Rastenij. Trudy Biologo-Počvennogo Instituta. Novaja Serja, 26(129): Man, de J.G Onderzoekingen over vrij in de aarde levende Nematoden. Tijdschrift der Nederlandsche Dierkundige Vereeniging, 2: Mulvey, R.H The Mononchidae: a family of predaceous nematodes VII. Genus Prionchulus (Nematoda: Mononchidae). Canadian Journal of Zoology, 45: Samsoen, L., Arpin, P., Khan, S. H. and A. Coomans Differentiation of Prionchulus muscorum (Dujardin, 1845) Wu & Hoeppli, 1929 and P. punctatus (Cobb, 1917) Andrássy, 1958 by egg-shell structure. Revue de Nématologie,7: Susulovsky, A. and G. Winiszewska Prionchulus fistulosus sp. nov. (Nematoda: Mononchina) from Ukraine. Annales Zoologici, 52: Tysowski, A Wolnożyjące nicienie (nematodes) zebrane w powiecie sokalskim. Rozprawy i Wiadomości z Muzeum im. Dzieduszyckich, 1: , pl. 3. Winiszewska, G Prionchulus brevicaudatus sp. nov., (Nematoda: Mononchina) from Poland. Annales Zoologici, 52: Wu, H.W. and R. J. C. Hoeppli Free-living nematodes from Fookien and Chekiang. Teil III. Beihefte (1). Archiv für Schiffsund Tropen-Hygiene, Pathologie und Terapie Exotischer Krankheiten, 33: 35 43, pls Cobb N.A Subdivisions of Mononchus. Journal of Parasitology, 2: Zell, H Nematoden eines Buchenwaldbodens 3. Prionchulus muscorum (Nematoda, Mononchida). Carolinea, 42: Received: April 10, 2003 Accepted: June 20, 2003