Vol. 3, No. 1, pp. 81-89 Afro-Asian Journal of Nematology June, 1993

Descriptions of Paratylenchus pernoxius sp. n. and Paralongidorus duncani sp. n, from Senegal, and the Synonymization of Longidoroides with Paralongidorus

M. R. Siddiqi", P.Baujard" and D. Mounport'?"

• International Institute or Parasitology, 395a Hatfield Road, SL A1bans, Herts. AlA OXU, UK •• ORSTOM Laboratoire de Nematologie, B.P. 1386, Dakar, senegal; present address: Museum national d'Histoire natu­ relie, Laboratoire de Biologie Parasitaire, ProtistoJogie, Helminthologie, 61 rue Buffon, 75231 Paris cedex 05, France ••• Laboratoire de Biologie Animale, Faculte des Sciences, Dakar, Senegal

AbstracL Paratylenchus pemoxius sp. n. and Paralongidorus duncani sp. n. from Senegal are described and illustrated. Paratylenchus pemoxius collected and reared on Sorghum vulgare L. is less than 0.4 mm long and has lateral fields with 4 incisures, 4 distinct submedian lobes around oral opening, a short stylet 13.5-16.0 (14.7) um long, excretory pore 0-2 annules anterior to hemizonid, vulva at 80-86 (83.3) per cent of body, female tail straight, 18-25 (21.3) urn long having a smooth, conoid-rounded terminus and males with 18-21 (20) urn long spicules and a regularly tapering, straight tail with conoid-rounded terminus. Paralongidorus duncani collected around roots of fallow plants and reared on Sorghum vulgare L. has females with a small body 1.67-2.29 (2.03) mm long, small odontostyie and odontophore (40-45 (43.5) urn and 37-46 (41) urn long, respectively), vulva at 45-49 (46.8) per cent of body, tail 42-54 (47.6) urn long and regularly tapering to a rounded terminus, males with spicuJes 28-31 (29.7) urn long and juveniles with only three developmental stages. Longidoroides Khan, Chawla & Sa ha, 1978 has been synonymized with Paralongidorus and thirteen new combinations have been proposed.

Keywords: Longidoroides synonymy, new combinations, new species, Paralongidorus duncani; Paratylenchuspernox­ ius, Senegal, Sorghum vulgare, .

INTRODUCTION and fixed in FP 4 : 1 (Netscher & Seinhorst, 1969), and mounted in dehydrated glycerine on glass slides. During studies on ecology of from Sahelian For scanning electron microscopy the nematodes were zone of West Africa (Baujard, 1986), two new species of killed by gentle heating at 60°C for 30 seconds and processed plant-parasitic nematodes were identified in Senegal. They by the method of Baujard & Pariselle (1987). have been reared on Sorghum vulgare L. in the laboratory and are described here as Paratylenchus pernoxius sp. n. and Para­ SYSTEMATICS longidorus duncani sp. n. The latter species was first identified Paratylenchus pemorius sp. n. as belonging to the genus Longidoroides Khan, Chawla & (Figs 1 & 2) Saha, 1978, but a comparison with the paratypes of Paralongi­ dorus sali Siddiqi et al., 1963, the type species of Paralongido­ Measurements rus Siddiqi, Hooper & Khan, 1963, convinced us that it is a Holotype female: L 0.36 mm; a 24; b 4.6; c 15; c' species of Paralongidorus. We propose here to synonymize = = = = Longidoroides with Paralongidorus. = 2.6; V = 25-83.3; stylet = 15 urn. 25 paratype females: L = 0.28-0.38 (0.337±0.024) mm; a = 20-28 (23.8±1.8); b = 3.7-4.9 (4.2±O.3); c = 12-18 (15.5±1.59); c' = 2-2.9 (2.47±0.25); V = 80-86 (83.3±1.23); MATERIALS AND METHODS stylet = 13.5-16.0 (14.7±0.58) urn; tail = 18-25 (21.3±1.9) urn. The two species described here were collected 15 paratype males: L = 0.29-0.38 (0.335±0.021) mm; a = during surveys of the peanut cropping areas in Senegal, West 24-34 (27.6±2.7); b = ?; c = 12.8-14.3 (13.5±O.54); c' = Africa (Baujard & Martiny, 1991). They were reared in the 2.3-3.1 (2.68±0.3); T = 19-37 (28±5.8); tail = 22-28 laboratory on Sorghum vulgare L., extracted by elutriation (23.7±1.2) urn.

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Afro-Asian Journal of Nematology Vol. 3, No. 1, 1993 1

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IOpm KL f------=------jl A,B, E G, 1, 1 - N 1 20p m f--_----e:-._--i1 C-E, H

Fig. 1. Paraty/eru:hus pemoxius sp. n. A- F, J- N. Females. G-1. Males. A, B, G. Head ends. 1 C, K. Oesophageal regions. D, E, 1, L. Posterior ends. F. Lateral field. H. Reproductive tract. J. Entire mature female. M, N. Vulval regions. 1 82 1 1 •

1 Paraty/enchus pemoxius and Paralongidorus duncani: Siddiqi, Baujard, Mounport

1 Description (23.7±1.2) ~m long; terminus conoid-rounded, smooth. lYpe host and locality: Collected in March, 1985 during the Female. Body cylindrical, usually tapering regularly towards dry season in a lowland cropped with sorghum (Sorghum vu/­ both ends but with no marked tapering behind vulva except in 1 gare L.) in the previous rainy season, at Keur Boumi, about 5 mature females which become a little fatter in the reproduc­ km on Louga to Gueoul Road, Senegal, and then reared in tive region, slightly arcuate ventrally or C-shaped. Cuticle dis­ the laboratory on Sorghum vu/gare cv. 51 69 in Senegal. tinctly striated; striae 1.1-1.3 (1.2) ~m apart near middle. Lat­ 1 eral fields with 4 incisures forming 3 bands of which the lYpe specimens: Holotype female, 140 female and 10 male middle one is slightly narrower than the others; inner two inci­ paratypes on permanent glycerine mounted slides and numer­ sures sometimes difficult to see. Cephalic region with straight ous females and juveniles in fixative, at International Institute to convex sides, tapering anteriorly, with 4 distinct submedian of Parasitology, St. Albans, England; 10 female and one male lobes bordering a slightly convex central area, as seen in lat­ paratypes at Rothamsted Experimental Station, Harpenden, eral view. Face view in SEM shows 4 large round submedian England. Numerous females, males and juveniles in fixative at lobes, an 1-shaped oral opening guarded by 2 large lateral the Laboratoire de Biologie Parasitaire, Protistologie, HeI­ Iiplets and 4 annules (Fig. 2B,C). 1 minthologie, Muséum national d'Histoire naturelle, Paris, Stylet strong, 13.5-16.0 (14.7±0.58) ~m long; conus 8.5-10.5 France. Five female paratypes, 1 male and 2 juveniles at the (9.7) ~m in length; basal knobs prominent, rounded, c10sely National Nematode Collection, Indian Agricultural Research packed together, 2.3-2.7 ~m acrossand 2-2.5 ~m high. Orifice Institute, New Delhi, India. 1 of dorsal oesophageal gland 4-6 ~m behind stylet. Excretory pore 60-90 (69±7.5) ~m from anterior end, at or up to two an­ Diagnosis and'relationship: Paratylenchus pemoxius sp. n nules anterior to hemizonid but usually at its anterior edge. is recognized by its body 0.28-0.38 mm long, female stylel Hemizonid distinct, 2-3 annules long. Deirids at level ofhemi­ 13.5-16 (14.7) ~m long, distinct submedian lobes, four inci· 1 zonid, 0-4 ~m behind excretory pore. Oesophagus typical, sures in lateral field, distinct hemizonid with excretory pore 70-89 (85±4.9) ~m long; distance from anterior end ofbody to located at or near its anterior margin, vulva at 80-86 (83.3) base of median oesophageal bulb almost as long as that from ~r cent, regularly tapering posterior end with straight to vulva to tail end. 1 shghtly arcuate tail ending in a conoid-rounded terminus and Vulva distinct, 230-318 (279±22.5) ~m from anterior end ot abundance of males. body; lateral vulval flaps straight-sided to slightly rounded It cornes close to P. italiensis Raski, 1975 and P. co/brani when seen laterally, 5 annules long (Fig. 2E). Vagina directed Raski, 1975, paratypes ofwhich have been studied and com­ 1 inward and forward; no post-uterine sac. Spermatheca round pared with il. It differs from P. ita/iensis in having slightly to slightly oval longitudinally, usually 11-13 ~m long, 10-11 longer body (31 females: 0.23-0.34 (0.28) mm long in the ~m wide in young and 30-31 ~m long and 20-21 ~m wide in latter), prominent submedian lobes, spermatheca with sperm, old females; packed with small rounded sperm in most fe­ straight-sided vulval flaps and tail tip conoid-rounded and not 1 males, 5-7 sperm occupying its width. Ovary anteriorly out­ acute as in the latter and the presence of males. From P. co/­ stretched, often irregularly bent, with 8-15 (11) oocytes in brani it differs in having prominent submedian lobes, slightly young and 17-29 in old females. smaller stylet (15-18 (16) ~m long in the latter), distinct hemi­ Rectum short, difficult to see. Anus pore-like. Tail straight 1 zonid, more posterior vulva (V = 79-82 (81) in the latter), to slightly dorsally convex but never arcuate, regularly taper­ large straight-sided vulval flaps, large spermatheca with smail mg to a small, smooth, conoid-rounded terminus 18-25 sized sperm, and male tail being straight, sometimes slightly (21.3±1.9) ~m long, with 16-25 (20) annules ventrally; lateral arcuate ventrally (curved a1most to 90 degrees in the latter). 1 field reaching near to tail terminus (Fig. 2D).

Male. Cephalic region conoid-rounded to truncate, about 5 ~m wide at base and 3 ~m high. Excretory pore at anterior ParalongùJorus dum:ani sp. n. 1 halfof hemizonid which is distinct and 2-3 annules long, 64-75 (Figs 3 & 4) (68±5.6) ~m from anterior end of body. Testis anteriorly out­ stretched. Spicules slightly arcuate ventrally, strongly cepha­ Measurements 1 lated, slightly flanged in posterior half except near distal end w~ich may be knobbed or notched with terminal aperture Holotype remale: L = 2.28 mm; a = 72.4; b = 9.5; c = 45; (Fig. 2F-H); 18-21 (20±1) ~m long. Gubernaculum linear or c' = 2.6; V = 45.2; odontostyle = 40 ~m; odontophore = 39 trough-shaped, fixed, 3.0-5.5 (4.2±0.6) ~m long. Penial tube ~m. 1 formed by c10acal lips, about 1.5 ~m long, posteriorly with a 2S paratype remales: L = 1.67-2.29 (2.03±O.096) mm; a = single, projecting hypoptygma (Fig. 2F,H). Tail regularly 47-82 (70±9.8); b = 6.8-9.6 (8.27±O.88); c = 39-50 (43±2.6); tapering, straight, sometimes slightly arcuate ventrally, 22-28 c' =2.3-2.9 (2.56±O.l6); V =45-49 (46.8±1.05): odontostyle

1 83 1 • 1

Afro-Asian Journal of Nematolugy Vol. 3, No. 1,1993 1 1 1 1 -1 1 1 1 1 1 1 1 1 ·1 1 1 Fig. 2. Paratylenchus pemoXlus sp. n. A- E. Females. F- H. Males. A-c. Head ends. D, F. Tail ends. E. Vulval region. G, H. Spicules. 1 84 1 1 •

1 Paratylenchur pemoxiur and Paralongidorur duncani: Siddiqi, Baujard, Mounport

1 = 40-45 (43.5±1) ~m; odontophore = 37-46 (41±2.4) ~m; Odontophore with slightly swollen posterior region, 37-46 total stylet = 78-89 (83.2±3) ~m; stylet guiding ring = 20-25 (41±2.4) ~m long. Total stylet length 78-89 (83.2±3) ~m. (22.7±1.2) ~m; tail = 42-54 (47.6±3.97) ~m. Stylet guiding ring single, 20-25 (22.7±1.2) ~m or 2.1-2.5 head 1 10 paratype males: L = 1.87-2.09 (1.95±0.08) mm; a = widths from anterior end. Oesophagus dorylaimoid, 205-255 65-79 (70±4.2); b = 7.8-9.0 (8.2±0.43); c = 41-45 (43±1.24); (225±7) ~m long; posterior enlarged part cylindroid, 50-63 c'= 1.9-2.6 (2.18±

Afro-Asian Journal of Nematology Vol. 3, No. l, 1993 1

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Fig. 3. Paralongidoms duncani sp, n. A, C-E, J, K. Females. B, F. Male. G, L. Second-stage juveniles, H, M, Third-stage juveniles. l & N. Fourth -stage juveniles, O. Paralongidoms sali, female paratype, head 1 end. A-C, O. Head ends. D. Basal region of oesophagus. E. Anterior reproductive branch. F-K. Tail ends. L-N. Genital primordia on ventral side of intestine. 1 86 1 1

1 Para tylenchus pemoxius and Paralongidorus duncani: Siddiqi, Baujard, Mounport 1 1 1

1 1 1 1 1 1 1 1 1· 1 1 Fig. 4. Paralongidorus duncani sp. n. A-O. Females. E-J. Males. A, B, E-H. Head ends. C, J. Tail ends. 1 O. Vulval region. 1. Spicules. 87 1 1

Afro-Asian Journal of Nematology Vol. 3, No. 1, 1993 1

This species is named in honour of Dr. L. W. Duncan in ap­ structure, but in P. duncani, the sensillum is not visible al­ 1 preciation of his valuable contribution to Sahelian nemato­ though there is a small slit in its place; the flap in P. duncani logy when he visited ORSTOM Laboratory, Dakar, Senegal. appears to overhang a depression (Fig. 2A,E,G).

Type host and locality: Collected in August, 1991 during 1 the rainy season from sandy soil at a depth of 50-100 cm around roots of fallow plant~ (Andropogon gayanus Hack., Synonymy of the genus Longidoroicks Khan, Chawla Guiera senegalensis Lam.) at Nebe, 5 km on Diourbel to Kao­ & Saha, 1978 1 lack Road, Senegal and then reared in the laboratory on Sor­ ghum vulgare cv. 5169 in Scnegal. Coomans (1985) analysed main morphological characters of various genera of the and discussed their Type specimes: Holotype femaJe, 75 female, 10 male and phylogenetic relationships. He gave diagnostic characters of 100 juvenile paratypes on permanent glycerine mounted Xiphinema Cobb, 1913, Xiphidorus Monteiro, 1976, Longido­ sI ides and numerous females and juveniles in fixative, at In­ rus Micoletzky, 1922, and Paralongidorus Siddiqi, Hooper and ternational Institute of Parasitology, St. Albans, England; 5 Khan, 1963, but for the genusLongidoroides Khan, Chawla & 1 female and two male paratypes at Rothamsted Experimental Saha, 1978, he wrote: ''As Paralongidorus and Longidorus ex­ Station, Harpenden, England. Numerous fema1cs, males and cept for the amphids that have a pouch-like fovea and wide juveni1cs in fixative at the Laboratoire de Biologie Parasitaire, slit-like apertures." At present Longidoroides is differentiated Protistologie, Helminthologie, Muséum national d'Histoire from Paralongidorus Siddiqi, Hooper & Khan, 1963 only by 1 naturelle, Paris, France. Two female paratypes at the National the size and shape of the amphidial pouch (fovea) which in Nematode Collection, Indian Agricultural Research Institute, the former is stirrup-shaped with large slit-like aperture, and New Delhi, India. pouch-like with small to large slit-like aperture in the latter. This difference largely depends on the narrowing of the head 1 Diagnosis, relationship and discussion: Paralongidorus end and the degree ofamphidial duct pushing into the base of duncani sp. n. is recognised by its small body size, odontostyle the pouch. and odontophore. In having a small odontostyle it cornes Species included in Longidoroides show considerable vari­ 1 close to Paralongidorus pulcheroides (Jacobs & Heyns, 1987) ation in the shape and size of amphidial pouch and the gen­ comb. n. (syn. Longidoroides pulcheroides Jacobs & Hcyns, eral pattern is the same as shown by the type species of Para­ 1987) from which it differs in having a smaller body, smaller longidorus, P. sali Siddiqi, Hooper & Khan, 1963 (see Fig. 3, adantostyle, smaller c ratio and a narrower and longer tail 0). Longidoroides, therefore, cannot be differentiated from 1 (body Iength = 3.03-3.72 (3.38) mm, odontostyle 55-67 (59) Paralongidorus and is proposed here as ajunior synonym ofit. Accordingly, the following species ofLongidoroides are trans­ !lm long, c = 93-124 (111), c' = 1.4-1.7 (1.5) in P. pulcher­ ferred to Paralongidorus and those either originally described oides). Longidorus juveniloides Jacobs & Heyns, 1987 has in Paralongidorus, or later transferred to it from other genera 1 similar sized odontostyJc, but it has large bilobed amphidial are retumed to il. pouches with indistinct apertures and hence belongs ta a dif­ ferent genus. Paralongidorus bikanerensis (Lai & Mathur, 1986) comb. n. We have found that Paralongidorus duncani has flap-like 1 syn. Longidoroides bikanerensis Lai & Mathur, 1986 structures at the site of four cephalic sensilla. Such structures P. cedari (Khan, Chawla & Saha, 1978) comb. n. have been described in P. deborae, P. sp. from Port Elizabeth, syn. Longidoroides cedari Khan, Chawla & Saha, 1978 Cape Province and P. sp. from Knysna, Cape Province, South P. clavicaudatus (Jacobs & Heyns, 1982) comb. n. 1 Africa by Swart and Heyns (1987). A similar species described syn. Longidoroides clavicaudatus Jacobs & Heyns, 1982 recently, Paralongidorus bullatus by Sharma and Siddiqi P. costatus (Jacobs & Heyns, 1987) comb. n. (1990) alsa has similar flaps at the cephalic sensilla. The three syn. Longidoroides costatus Jacobs & Heyns, 1987 species examined by Swart and Heyns have offset head and P. eugeni (Khan, 1987) comb. n. 1 may belong to Siddiqia Khan, Chawla & Saha, 1978 which syn. Inagreius eugeni Khan, 1987 Luc and Doucet (1984) synonymized with Paralongidorus. Longidoroides eugeni (Khan, 1987) Jairajpuri & The typical flap-like structure at the cephalic sensilla which Ahmad,1992 1 may be modified cephalic setae in this group gives a further P. gloriosus (Khan, 1982) comb. n. diagnostic character, besides the knob-like head offset by a syn. Inagreius gloriosus Khan, 1982 constriction, to the genus Siddiqia but its occurrence in P. Longidoroides gloriosus (Khan, 1982) Luc & duncani suggests that it may be a cornmon character of the Doucet, 1984 1 Paralongidorus as presently recognized. In these species the P. latilabiatus (Jacobs & Heyns, 1987) comb. n. cephalic sensillum is located just anterior to the flap-like syn. Longidoroides latilabiatus Jacobs & Heyns, 1987 1 88 1 1

1 Paratylenchus pemoxius and Paralongidorus duncani: Siddiqi, Baujard, Mounport

1 Rlongiurus (Chawla & Samathanam, 1981) comb. n. Fortuner, 1987 (Nemata: Belonolaimidae). I. Etudes au champ. syn. Longidoroides longiurus Chawla & Samathanam, Afro-Asian Journal ofNematology 1, 94-100. 1981 Baujanl, P. and A. Pariselle 1987. Fabrication de microtamis et pré­ 1 R pmi (Jacobs & Heyns, 1987) comb. n. paration des nématodes pour l'observation au microscope élec­ syn. Longidoroides pini Jacobs & Heyns, 1987 tronique à balayage. Revue de Nématologie 10,477-481. R pulcher (Jacobs & Heyns, 1982) comb. n. Coomans, A. 1985. A phylogenetic approach to the classification of syn. Longidoroides pulcher Jacobs & Heyns, 1982 the Longidoridae (Nematoda : ). Agriculture, Eco­ 1 R pulcheroïdes (Jacobs & Heyns, 1987) comb. n. systems and Environment, 12 (1984/85), 335-354. syn. Longidoroides. pulcheroides Jacobs & Heyns, 1987 Khan, E., M. L. Chawla and M. Saha 1978. Comments on thc classi­ R seclepsi (Khan, Singh & Singh, 1981) comb. n. fication of the Longidoroidea (Nematoda) with descriptions of syn. Siddiqia seclipsi Khan, Singh & Singh, 1981 three ncw species. lndÙJn Journal ofNematology 6 (1976), 47-68. 1: Longidoroides seclipsi (Khan, Singh & Singh, 1981) Netscher, C. and J. \v. Seinhorst 1969. Propionic acid better than Luc & Doucet, 1984 acetic acid for killing nematodes. Nematologica 9, 286. R teres (Khan, 1987) comb. n. Sharma, S. B. and M. R. Siddiqi 19911. Paralongidorus bullatus n. sp. syn. Inagreius teres Khan, ]987 1 from groundnut soils in Niger and comments on Xiphinema Longidoroides teres (Khan, 1987) Jairajpuri & Ahmad,1992 parasetariae Luc. Journal ofNematology 22, 579-584. Siddiqi, M. R., D. J. Hooper and E. Khan 1963. A new nematode 1 LITERATURE CITEU genus Paralongidorus (Nematoda: Dorylaimoidea) with descrip­ tions of Iwo new species and observations on Paralongidorus Baujard, P. 1986. Ecologie des nématodes dans le bassin arachidier citri (Siddiqi, 1959) n. comb. Nematologica 9, 7-14. du Sénégal. Revue de Nématologie 9, 288 (Abstr.). Swart, A. and J. Heyns 1987. Comparative morphology of the head 1 Baujard, P. and B. Martiny 1991. Données nouvelles sur le néma­ regions of sorne longidorid nematodes from South Africa using tode Tylenchorhynchus germani (Germani & Luc, 1984) Luc & the SEM.Phytophylactica 19,99-101. 1 1 1 1 1 1 1 1 1 89 1