Revision of the Genus Tylencholaimus De Man, 1876. Remaining Species
Identifieur interne : 000709 ( Istex/Corpus ); précédent : 000708; suivant : 000710Revision of the Genus Tylencholaimus De Man, 1876. Remaining Species
Auteurs : R. Pena Santiago ; A. CoomansSource :
- Nematologica [ 0028-2596 ] ; 1996.
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DOI: 10.1163/004525996X00037
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ISTEX:E1B42DD75E7414C86FB8B30A4F95E9CD886DE83ALe document en format XML
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<p>REVISION OF THE GENUS TYLENCHOLAIMUS DE MAN, 1876. REMAINING SPECIES BY R. PENA SANTIAGO1) and A. COOMANS2) 1) Departamento de Biología Animal, Vegetal y Ecología, Universidad de Jaén, Virgen de la Cabeza n° 4, 23008-Jaén, Spain; 2)Instituut voor Dierkunde, Rijksuniversiteit Gent, B-9000 Gent, Belgium The present paper deals with the taxonomy of five species belonging to the genus Tylencholaimus de Man, 1876. Two of them are prodelphic: T. stecki Steiner, 1914 and T. vulvulatus Rahman, Jairajpuri, Ahmad & Ahmad, 1987; they are characterized by having perioral region not disc-like, anterior part of the pharynx expanding gradually to the basal bulb, and posterior genital branch more than two body widths long. The other three species are opisthodelphic: T. intermedius n. sp., T. gertii Kruger, 1965 and T. pakistanensis Timm, 1964. These five species are described and illustrated. Moreover, T. rossi Yeates, 1979 is transferred to Tantunema Siddiqi, 1979 as Tantunema rossi (Yeates, 1979) n. comb. Several species recently described or transferred to Tylencholaimus, and others considered species inquirendae are briefly discussed. Keywords: prodelphic and opisthodelphic species, revision, taxonomy, Tylencholaimus. In this sixth paper of the series we have included two prodelphic species, T. stecki Steiner, 1914 and T. vulvulatus Rahman et al., 1987, which have the perioral region not disc-like, the slender part of the pharynx muscular and ex- panding gradually, and posterior genital branch well developed (more than two body widths long); and three opisthodelphic species, viz. T. gertii Kruger, 1965, T. pakistanensis Timm, 1964 and a new species. T. rossi Yeates, 1979 is tras- ferred to Tantunema Siddiqi, 1982 as Tantunema rossi (Yeates, 1979) n. comb.; this genus will be treated in a separate paper. According to Loof (in litt.) T. zee- landicus de Man, 1876 belongs to the genus Doryllium. For methods used, we refer to the first paper of the series (Pena Santiago & Coomans, 1994a). DESCRIPTIONS Tylencholaimus stecki Steiner, 1914 (Fig. 1; measurements in Table I) Tylencholaimus stecki Steiner, 1914 Steiner (1914); Arch. Hydrobiol. Planktonk 9: pp. 428-429.</p>
<p>418 de Man ( 1921 ); Capita zoologica 1 ( 1 ): pp. 40-42. Micoletzky (1922); Arch. Naturg., Berlin 87 (1921): pp. 427-430. Micoletzky (1925); K. Danske Vidensk. Selsk. Skr., naturw. og math. Afd. 8: pp. 183. Fuchs ( 1930); Zool. Jarb., Jena 59: pp. 608-609. de Coninck (1935); Rev. Zool. Bot. Afr. 26: pp. 290-293. Thorne (1939); Capita zoologica 8: p. 58. Gadea (1953); Publ. Inst. Biol. Apl. 14: p. 101. Meyl (1954a); Arch. zool. Ital. 39: p. 244. Meyl (1954b); Mycopath. Mycol. Appl. 7: p. 69. Andrdssy (1958); Acta Zool., Acad. Scient. Hungar. 4: pp. 76-77. Coomans (1962); Nematologica 7: pp. 149-151. Loof & Jairajpuri (1968); Nematologica 14: pp. 342-344. Popovici (1968); St. cerc. Biol., Ser. Zool. 20: p. 258. Sauer (1969); Nematologica 15: pp. 366-367. Vinciguerra (1986); Nematol. medit. 14: pp. 112-113. Winiszewska (1987); Fragmenta faunistica 31: p. 34. Tylencholaimus alpinus Vinciguerra, 1982 Vinciguerra (1982); Animalia 9: pp. 317-318. Female: moderately slender nematodes of small size whose length varies from 0.7 to 1.0 mm. Body practically cylindrical, tapering towards both extrem- ities but somewhat more towards the anterior one. Habitus very variable after fixation: almost straight to clearly ventrally curved adopting G-shape (Fig. 1B). Outer cuticle relatively thin and with very fine transverse striations which often become imperceptible; inner layer irregular and wider than the outer one. Radial refractive elements present but often inconspicuous except for the caudal region. Lateral pores obscure in the material examined. Lateral chord about one-third of the midbody diameter. Lip region (Fig. IE, F) offset by shallow constriction; about twice as wide as high or about one-fourth (? cf. flattened specimens) of the body width at neck base. Lips amalgamated and rounded, with their inner part visibly elevated. Labial and cephalic papillae not interfering with the head contour. Amphid open- ing at level of the cephalic constriction and occupying about two-fifths of the lip region diameter. Odontostyle typical of the genus, its length about two-thirds the lip region width; its aperture about one-third of the total length. Odon- tophore somewhat longer than the odontostyle, with small basal knobs which have become obscure in some specimens. Cheilostoma practically cylindrical or a truncate cone, without a visible perioral sclerotization. Guiding ring simple. Pharynx (Fig. 1 C) consisting of a slender but muscular anterior part which expands gradually into the basal bulb. This latter (Fig. 1 G) is practically cylin- drical, about 5-6 times as long as wide and occupies about two-fifths of the total neck length. Nerve ring located at 37-40% of the neck region. Cardia rounded or</p>
<p>419 Fig. 1. Tylencholaimus stecki Steiner, 1914. A: Male entire. B: Female entire. C: Neck region. D: Female genital system. E: Lip region in median view. F: Same in surface view. G: Pharyngeal bulb and cardia. H, K: Female tail. I: Male, posterior body region. J: Vagina. L: Spicule. M: Lateral guiding piece. N: Sperm.</p>
<p>420</p>
<p>421</p>
<p>422</p>
<p>423</p>
<p>424 somewhat conoid, in general wider than long; partially surrounded by intestinal tissue. Genital system (Fig. 1 D) pseudomonodelphic-prodelphic. Ovary reflexed, with numerous oocytes and rather long since it often reaches and extends beyond the oviduct-uterus junction. The oviduct is a relatively narrow tube without visible lumen and without an offset pars dilatata. A poorly developed sphincter separates oviduct and uterus. The latter is a wider tube with clear lumen but without special modifications. The posterior genital branch consists of a rather long postvulval uterine sac, 2-4 times as long as the corresponding body width, containing abundant sperm and ending in a small cell mass. Vagina (Fig. 1 J) practically cylindrical but sometimes more conical, extending inwards over one- third to two-fifths of the corresponding body diameter; its part adjacent to the vulva weakly sclerotized, the other part surrounded by circular muscles. Vulva apparently a transverse slit. Prerectum (Fig. 1H) about twice the anal body width long. Rectum length almost equal to the anal body diameter. Tail (Fig. 1 K) practically hemispherical and sometimes (probably due to flattening, see Fig. 1 H) more conoid. Caudal pores obscure in the specimens studied but probably two pairs, one more anterior subdorsal and another lateral and more posterior. Male: general morphology (Fig. lA) very similar to the female. Genital sys- tem diorchic. Testes opposed. Apart from the adanal pair, 2-4 widely spaced ven- tromedian supplements are present beginning outside the range of the spicules. Tail (Fig. 11) more conoid than the female, ventrally almost straight, dorsally convex. Spicules (Fig. 1L) ventrally arched, equal or scarcely longer than the anal body width. Lateral guiding pieces (Fig. 1M) relatively short. Spermatozoa (Fig. IN) elliptical to spindle-shaped. Diagnosis: the species can be distinguished by its small size (L = 0.7-1.0), lip region offset by a weak constriction and with amalgamated lips, odontostyle 5-7 pom long, slender part of the pharynx expanding gradually, pharyngeal bulb 90-102 Nm long, female genital system pseudomonodelphic-prodelphic, V = 60-67, posterior genital branch consisting of a long uterine sac and a terminal small cell mass, tail hemispherical, spicules 23-28 pom long and 2-4 ventromedian supplements. Relationships: T. stecki is very similar to T. vulvulatus Rahman, Jairajpuri, Ahmad & Ahmad, 1987 from which is can be separated by the symmetrical lips (vs asymmetrical) of the vulva, shorter prerectum (vs 6-10 times the anal body width) and tail with different morphology (vs convex-conoid).</p>
<p>425 Distribution and habitat: it is a widely distributed species which at present has been recorded in the following sites: i) sphagnum moss, near Arosa, Switzer- land (Steiner, 1914; type population); ii) different habitats in the Netherlands (de Man, 1921); iii) several localities of different countries (Micoletzky, 1922); iv) several localities in Denmark (Micoletzky, 1925); v) Germany (Fuchs, 1930); vi) moss, Nyamuamba, Zaire (De Coninck, 1935); vii) Switzerland (Altherr, 1950); viii) grass roots in Cervera de Pisuerga, Palencia, Spain (Gadea, 1953); ix) Italy (Meyl, 1954a); x) Germany (Meyl, 1954b); xi) several localities, Bul- garia (Andrdssy, 1958); xii) dry sandy soil of an old meadow, around roots of Lolium perenne L. and Cynosurus cristatus L., Belgium (Coomans, 1962); xiii) several localities in Sweden and Switzerland (Loof & Jairajpuri, 1968); xiv) Roumania (Popovici, 1968); xv) uncultivated mallee soil, Hattah, Australia (Sauer, 1969); xvi) several localities in Italy (Vinciguerra, 1982 and 1986); xvii) Swietokrzyskie Mountains, Poland (Winiszewska, 1987); xviii) different habitats in province of Navarra, Spain (Hemández & Jordana, 1990). Type material: probably lost. Material examined: one female and one male from Italy (courtesy of Dr. M. T. Vinciguerra). Seven females and five males from Navarra, Spain (courtesy of Dr. M. A. Hernández). Etymology: the species was named in honor of the entomologist Dr. T. Steck. Tylencholaimus vulvulatus Rahman, Jairajpuri, Ahmad & Ahmad, 1987 (T. vulvulatum emend.) (Fig. 2; measurements in Table I) Tylencholaimus vulvulatus Rahman, Jairajpuri, Ahmad & Ahmad, 1987 Rahman et al. ( 1987); Indian J. Nematol. 16 ( 1986): pp. 201-204. Female: slender nematodes of small size, 0.80-0.93 mm long. Body cylindri- cal, tapering towards both extermities but somewhat more towards the posterior end. Habitus (Fig. 2F) after fixation slightly ventrally arched, especially in the posterior half of the body. Outer layer of the cuticle thin and apparently smooth; inner layer with irregular outline and separated from the outer layer. Radial refractive elements visible along the entire body but clearer at level of neck and tail. Lateral chord about one-third of the midbody diameter. Lateral pores obscure in the single specimen examined. Lip region (Fig. 2G, H) offset by constriction; about 2.5 times as wide as high or one-third of the body diameter at neck base. Lips amalgamated and rounded, inner part projecting. Labial and cephalic papillae not interfering with</p>
<p>426 Fig. 2. Tylencholaimus vulvulatus Rahman et al., 1987. A: Male entire. B: Neck region. C: Pha- ryngeal bulb and cardia. D: Female, posterior body region. E: Female genital system. F: Female entire. G: Lip region in median view. H: Same in surface view. I: Male, posterior body region. J: Female tail. K: Vagina. L: Spicule. M: Lateral guiding piece.</p>
<p>427 the head contour. Cheilostoma a truncate cone; its walls weakly sclerotized but thickening in the perioral region. Odontostyle about three-fourths of the lip region width long, typical of the genus; its aperture one-fourth of the total length. Odontophore scarcely longer than the odontostyle, with small basal knobs. Guiding ring simple. Anterior part of the pharynx (Fig. 2B) slender but muscular; expanding grad- ually to the basal bulb. Pharyngeal bulb (Fig. 2C) cylindrical, about six times as long as wide and occupying one-third to two-fifths of the total neck length. Nerve ring located at 36% of the total neck length. Cardia conoid, almost as long as wide. Gential system (Fig. 2E) pseudomonodelphic-prodelphic. Ovary reflexed, with numerous oocytes and reaching the sphincter level. The oviduct joins the ovary subterminally and consists of a tube-like distal part and a wider proximal part without visible lumen. Oviduct-uterus junction marked off by a poorly devel- oped sphincter. Uterus about three times the corresponding body diameter long; containing an egg which is three times as long as wide. Posterior genital branch reduced to a long uterine sac 3.7 times as long as the body width, and containing sperm. Vagina (Fig. 2K) pyriform and directed forward, extending to half the corresponding body diameter; its part adjacent to the vulva poorly sclerotized, the other part circled by weak musculature. Vulva apparently a transverse slit; its anterior lip swollen and more prominent than the posterior one. Prerectum (Fig. 2D) rather long, about six times the anal body width. Rectum somewhat shorter than anal body width. Tail (Fig. 2J) convex conoid, slightly longer than the anal body width. Caudal pores not very clear but apparently two pairs, one subdorsal and one more lateral. Male: general appearance (Fig. 2A) similar to female but with the posterior body region more ventrally curved. Diorchic with testes opposed. In addition to the adanal pair three spaced ventromedian supplements are present, the anterior- most one poorly developed. Sixteen copulatory muscles present at right side and fifteen at left side of body. Tail (Fig. 21) conoid, ventrally almost straight and dorsally convex. Spicules (Fig. 2L) ventrally arched, 1.6 times the anal body width long. Lateral guiding pieces (Fig. 2M) relatively small. Diagnosis: the species is characterized by its 0.80-0.93 mm long body, an- terior part of the pharynx extending gradually, odontostyle 6-7 pm, female gen- ital system pseudomonodelphic-prodelphic, V = 61-69, vulva lips asymmetrical since the anterior one is clearly swollen, posterior genital branch an uterine sac 87-167 ?m long, tail convex conoid (19-22 pom, c = 40-43, c' = 1.0 in females), spicules 28-32 pom and three ventromedian supplements.</p>
<p>428 Relationships: T. vulvulatus is close to T. stecki Steiner, 1914 from which it can be separated by its asymmetrical vulval lips (vs symmetrical), longer prerectum (vs twice the anal body diameter) and longer tail with different shape (vs hemispherical). Distribution and habitat: the species has been recorded only from the type locality in soil around roots of grasses, Shillong Peak, Meghalaya, India. Type material: six females (holotype and paratypes) and four males (para- types) on slides labelled Tylencholaimus vulvulatum sp.n./1-4 deposited in the Zoology Department, A.M.U., Aligarh, India; a paratype female and a male deposited in the National Nematode Collection, I.A.R.I., New Delhi, India. Material examined: one paratype female and one paratype male on slide no 2 by courtesy of Dr. W. Ahmad. Etymology: unknown. Rem.arks: our description and measurements agree well with the original one, despite small differences in some measurements. However, we have observed only three ventromedian supplements in the male (vs 7-8 after Indian authors) which is an important difference. Tylencholaimus intermedius n. sp. (Fig. 3; measurements in Table I) Female: slender nematodes of very small to small size, 0.44-0.66 mm long. Body cylindrical, tapering slightly towards the anterior end. Habitus (Fig. 3A) after fixation more or less ventrally curved to C-shape. Outer cuticle thin and with very fine transverse striations clearer at both extremities. Inner cuticle wider than the outer layer, with rather regular outline and not separated from the outer cuticle. Radial refractive elements not observed. Lateral chord occupying one-third of the midbody width. Lateral pores obscure. Lip region (Fig. 3E, F) cap-like, offset by a shallow constriction; scarcely more than twice as wide as high and about two-fifths of the body diameter at neck base. Lips amalgamated and rounded, with their inner part less elevated. Small postlabial weakly refractive sclerotization present. Amphid aperture and fovea small, opening at level of the cephalic constriction and occupying one- third of or somewhat less than the corresponding body width. Cheilostoma a truncate cone with slender walls which become thickened at the perioral region. Odontostyle relatively long, practically equal to the lip region width; its aperture one-third of total length. Odontophore slightly longer than the odontostyle, with basal thickenings but no knobs. Guiding ring simple.</p>
<p>429 Fig. 3. Tylencholaimus intermedius n. sp. (female). A: Entire. B: Pharyngeal bulb and cardia. C: Genital system. D: Posterior body region. E: Lip region in median view. F: Same in surface view. G-I; Tail. J: Vagina.</p>
<p>430 Anterior part of the pharynx slender and weakly muscular, expanding suddenly to the basal bulb. Pharyngeal bulb (Fig. 3B) cylindrical, 7-8 times as long as wide and occupying two-fifths of the total neck length. DN and S2N well visible, S IN more obscure; DN situated rather posteriorly. Cardia in general wider than long, surrounded by intestinal tissue. Nerve ring located at 37-44% of the total neck length. Genital system (Fig. 3C) monodelphic-opisthodelphic. Anterior genital branch completely absent. Ovary relatively small, with less oocytes and normally not reaching the oviduct-uterus junction. Posterior gonoduct poorly differentiated; the oviduct is separated from the uterus by a narrowing but no apparent sphincter exists. Vagina (Fig. 4J) somewhat pyriform and extending inward for half the corresponding body diameter; its wall adjacent to the vulva slightly sclerotized, the other part surrounded by weak circular muscles. Vulva a transverse slit. Prerectum (Fig. 3D) 2.7-3.2 anal body widths long. Rectum somewhat longer than anal body width. Tail (Fig. 3G-I) cylindroid, and somewhat variable in length; core outline rather regular. Caudal pores obscure. Male: unknown. Diagnosis: the new species is characterized by its small body (L = 0.43- 0.66), lip region offset by a weak constriction, presence of postlabial scleroti- zation, small amphid aperture, odontostyle 5.5-7.0 p,m long, slender part of the pharynx weakly muscular, female genital system opisthodelphic, anterior uterine sac completely absent, V = 50-54, tail cylindroid (14-20 pm long, c = 31-39, c' = 1.2-1.6), and males absent. Relationships: in being opisthodelphic the new species is close to T. gertii Kruger, 1965 and T. paki.rtanensis Timm, 1964. T. intermedius n. sp. differs from T. gertii by having postlabial sclerotization (vs absence), anterior part of the pharynx expanding abruptly (vs gradually), small amphid aperture (vs four-fifths of the corresponding head diameter), absence of prevulval sac (vs presence), more rounded tail (vs more conoid) with different terminal core, and male unknown (vs known). It can be distinguished from T. pakistanen.sis by having postlabial sclerotization (vs absence), relatively longer pharyngeal bulb (vs 27-33% of the total neck length) and more posterior vulva (vs V = 44.4-44.7). Type habitat and locality: soil around roots of avocado (Persea americana) and mango (Mangifera indica) in Vélez-Málaga, province of Málaga, Spain. Other habitats and localities: soil around roots of Quercus faginea, Q. ro- tundifolia and Fraxinus angustifolia in Sierras de Cazorla y Segura, province</p>
<p>431 of Jaen, Spain. Also in association with fir (Abies pinsapo) in Yunquera and Parauta (Serrania de Ronga), province of Malaga, Spain. Type material: four females (holotype and three paratypes) on slides nos 0147 (holotype), 0148 (two paratypes) and 0149 (one paratype) deposited in the nema- tode collection of the Departamento de Biologia Animal, Universidad de Ja6n, Spain. Etymology: the specific epithet is the latin word "intermedius" = interme- diate, and refers to the possible intermediate taxonomic position of this species between the genera Tylencholaimus and Tantunema (see remarks below). Remarks: general appearance and different diagnostic features such as the presence of postlabial sclerotizations and small but slit-like amphid aperture make T. intermedius n. sp. reminiscent of other species of the same genus as T. chathami Yeates, 1979 (although that species is prodelphic) and of those included in the genus Tantunema Siddiqi, 1982. The latter is, however, characterized by having pore-like amphid aperture. The new species shows that the two genera are closely related. Tylencholaimus gertii Kruger, 1965 (Fig. 4A-E) Tylencholaimus gertii Kruger, 1965 Kruger (1965); Proc. helm. Soc. Wash. 32: p. 1. Sauer (1969); Nematologica 15: p. 365. Description (adapted from Kruger, 1965) Dimensions (2 females): L = 0.61-0.62 mm; a = 17-21; b = 3.6; c = 22-24; V = 14616. Fem.ales: cuticle with fine, nearly inconspicuous transverse striae, two layers thick around the tail with a narrow portion on the terminus. Lip region offset by a deep constriction; the outer lips somewhat angular, the inner portion projecting (inwards and) forwards. Odontostyle 8 /cm long with an aperture of 3 pm,; the width of the spear 1 lem and the odontophore 10 lem. Amphids broad, opening 4/5 of the corresponding head diameter. Body nearly straight when killed by gradual heat. Neck tapering so that the lip region is only 1/3 of the width at base of neck. Tail bluntly convex-conoid. The terminal core of the tail possesses a characteristic bowl-shaped form. Inner cuticle striated, irregularly separated from the outer cuticle. Lateral chord 1/4 as wide as body. Anterior region of pharynx expanding gradually to the basal bulb said to occupy about 1/2 its length. Cardia hemispherical. Intenstine three cells in circumference containing fine granules.</p>
<p>432 Fig. 4. A-E: Tylencholaimus gertii Kruger, 1965. A: Neck region. B: Female genital system. Posterior body region of male. D: Lip region in median view. E: Posterior body region of female. Redrawn after Sauer (1969). F-H: Tylencholaimus pakistanensis Timm, 1964. F: Lip region in median view. G: Female genital system. H: Female tail. Redrawn after Timm (1964).</p>
<p>433 Prerectum almost 6 anal body widths long; rectum as long as anal body width. Female reproductive system monodelphic-opisthodelphic, ovary reflexed about 2/5 back to vulva. Vagina about 1/3 across body. Small anterior uterine branch present. Males: unknown. Australian population described by Sauer (1969): "Females: (n = 10) L = 0.56-0.66 mm; a = 21-25; b = 3.5-4.3; c = 26-35; V= 43-48; spear 6-7 Mm. Male: (n = 1) L = 0.66 mm; a = 24; b = 3.9; c = 26: spear 6 pm; spicules (on chord) 25 pom. Fine lateral body pores arranged in two rows are visible in most specimens. Some radial elements appear refractive in most processed specimens. The only male collected has three ventromedian supplements and the characteristic invagi- nation of the terminal core of the tail." Diagnosis: T. gertii is characterized by its small body 0.56-0.66 mm long, lip region offset by a deep constriction and angular, odontostyle 6-8 pm, anterior part of the pharynx expanding gradually, female genital system opisthodelphic, V = 43-48, anterior branch reduced to a short uterine sac less than the corre- ponding body width long, tail convex conoid with a peculiar invagination of the terminal core of the tail, spicules 25 pom and three widely spaced ventromedian supplements. Relationships: in being opisthodelphic this species is close to T. pakistanensis Timm, 1964 and T. intermedius n. sp. but can be separated from these by the anterior part of its pharynx expanding gradually (vs abruptly), the presence of a short anterior uterine sac (vs absence), more conoid tail (vs more rounded) with a peculiar terminal core, and males known (vs unknown). Moreover, it is distinguished from T. pakistanensis by having longer body (vs L = 0.45-0.50), and from T. intermedius n. sp. by the more anterior vulva (vs V = 50-54), wider amphid opening (vs one-third of the lip region width) and absence of postlabial sclerotizations (vs presence). T. gertii is also similar to T. rossi Yeates, 1979 but observations on type material of this species furthermore revealed a head and amphid structure as in Tantunema Siddiqi, 1982. This species is therefore transferred to the latter genus as Tantunema rossi (Yeates, 1979) n. comb. Distribution and habitat: the species has been collected from soil in the Horticultural Research Institute, Pretoria (South Africa), and from wheat lands, Cullulleraine (Australia).</p>
<p>434 Type material: two females on slide 1386 deposited at the South African National Nematode Collection, Plant Protection Research Institute, Pretoria. Material examined: by courtesy of Dr. E. van den Berg we have seen the slide containing the type material; unfortunately this material is in such deficient state of conservation that it was of no use for our study. Sauer's material was not available to us. Etymology: unknown. Tylencholaimus pakistanensis Timm, 1963 (Fig. 4F-H) Tylencholaimus pakistanensis Timm, 1964 Timm (1964); Proc. helm. Soc. Wash. 31: p. 51. Description (adapted from Timm, 1964) "Females (4): L = 0.42 (0.45-0.50) mm; a = 19 (18-19.6); b = 3.5 (3.4-3.7); c = 31.5 (30.8-32.3); V = 45.8 (44.4-44.7); Ov 2 = 17.8%. Male: unknown. Cuticle moderately thick, especially thickened at the tail end. Distinct trans- verse striation at anterior end, not seen on rest of body, Lip region expanded, offset from rest of body, 30% of body diameter at oesophageal base; lips distinct. Inner circle of 6 prominent labial papillae. Amphids small, indistinct, 30% of body diameter broad, situated opposite spear guiding ring. Odontostyle 7 pm long or one lip region diameter; aperture about I ?em long. Odontophore 8 /im long or one lip region diameter, almost twice as broad as odontostyle, with small but distinct basal knobs. Simple spear guiding ring at level of constriction. Phar- ynx distinctly divided into two portions; basal portion 27-33% of neck length, with distinct radial muscles; anterior portion not distinctly muscular. Nerve ring just anterior to expanded basal portion (?). Intestine ending in long rectum, 1.7 anal body diameters long. Female reproductive system single; ovary postvulvar, reflexed almost to vulva: vagina muscular, inclined posteriorly. Tail subconoid, one anal body diameter long; tail papillae not observed." Diagnosis (modified): T. pakistanensis is characterized by its very small size (L = 0.45-0.50), odontostyle 7 pom long or equal to the lip region diameter, ante- rior part of the pharynx not distinctly muscular (?, extending abruptly), pharyn- geal bulb 27-33% of the pharyngeal total length, female genital system opistho- delphic, V = 44.4-46.7, vagina posteriorly directed, anterior uterine branch ab- sent, tail conoid (c' = 1 ), and males unknown.</p>
<p>435 Relationships: this species is very similar to T. gertii Kruger, 1965 and T. in- termedius n. sp. From T. gertii it differs in having shorter body (vs L = 0.56-066), absence of prevulval sac (vs presence), more rounded tail (vs more conoid) with rounded terminal core (vs a bowl-shaped terminal core), and male unknown (vs known). It can be separated from T. intermedius n. sp. by the absence of post- labial sclerotization (vs presence), relatively shorter pharyngeal bulb (vs 37-44% of the total neck length), and more anterior vulva (vs 50-54). Distribution and habitat: this species has been collected only from the type locality in soil around jute roots, Mymensingh, Bangladesh. Type m.aterial: four females (holotype and paratypes) whose deposition site is unknown to the authors. Material examirved: no specimens available for study. Etymology: the specific epithet derives form Pakistan, former name of the country where the species was collected. SPECIES RECENTLY DESCRIBED OR TRANSFERRED TO THE GENUS 1. Tylencholaimus accentuatus (Thome & Swanger, 1936) Andrdssy, 1991 (syn. Dorylaimus accentuatus Thorne & Swanger, 1936; Eudorylaimus accen- tuatus (Thorne & Swanger, 1936) Andrdssy, 1959; Thonus accentuatus (Thorne & Swanger, 1936) Andrdssy, 1986). This species presents several characters that are unusual for the genus, e.g. body 1.6 mm long, odontostyle somewhat longer than lip region width with the aperture occupying almost one-half its length. Furthermore nothing is known about the shape of the odontophore, hence we cannot find arguments to sub- stantiate the transfer of this species to the genus Tylencholaimus as proposed by Andrdssy (1991), however without any argumentation. In our opinion it is better to consider it as species inquirenda. 2. Tylencholaimus elongatus Jain, Saxena & Sharma, 1993 (T. elongatum emend.). We have some doubt about the identity of this species because the original description and illustration are insufficient: there is contradiction between text and figures (see in relation to the stylet), nothing is known of the odontophore morphology, and the pharyngeal bulb apparently occupies two-thirds (?) of the total neck length, an atypical feature in the genus; moreover, although four specimens were collected ("measurements (n = 4)", is mentioned in the text), only the holotype morphometry is given being L = 1.4 mm, an exceptionally</p>
<p>436 long body since all the known species of the genus are less than 1.2 mm long. So, it is better, in our opinion, to consider T. elongatum as species inquirenda. 3. Tylencholaimus loofi Dhanachand, 1994 (syn. T. minutus apud Dhanachand, Mohilal & Joymati, 1992 nec T. minutus Vinciguerra, 1986). This species has been inadequately described since some important taxonom- ical details (e.g. morphology of the anterior part of the pharynx) are not men- tioned ; the measurement of the stylet (22.5-25.5 pm) is abnormally high in the genus for a very small species, and if correct, a remarkable feature. Moreover, in the original description the species was not compared with closely related ones. If we assume that the anterior part of the pharynx is weakly muscular and it expands abruptly to the basal bulb (Indian authors' Fig. 1B, C seems to indicate it), T. loofi would belong to out second group of prodelphic species (Pena Santiago & Coomans, 1994b) being close to T. rumjhumi Khan & Laha, 1982 (other species insufficiently known) from which it can be distinguished by its more anterior vulva (V = 65.2-67.5 vs 70-78) and relatively shorter tail (c' = 0.7 vs 0.8-1.0); but if the anterior part of the pharynx expands gradually, this species can be classified under our third group of prodelphic species (Pena Santiago & Coomans, 1994c) being very similar to T. maritus Loof & Jaira- jpuri, 1968 from which it can be distinguished by its hemispherical (vs rounded conoid) and shorter tail (9-11 vs 15-16 pm; c = 50-54 vs 30-35; and c' = 0.6-0.7 vs 1.1-1.3). In any case, T. loofi is apparently a clearly distinct species although a more detailed description would be necessary. OTHER SPECIES INQUIRENDAE 1. Tylencholaimus kirifuri Kaburaki & Imamura, 1933 Lip region and stylet morphology in this species do not correspond to the genus. We agree with Thome (1939), Loof & Jairajpuri (1968) and Jairajpuri & Ahmad (1992) who considered this species as probably not belonging to Tylencholaimus. 2. Tylencholaimus nikkoensis Kaburaki & Imamura, 1933 Lip region, stylet and pharyngeal bulb morphology separate this species from those included in Tylencholaim.us. 3. Tylencholaimus parateres Meyl, 1957 This species was described on the basis of a single male. Lip region and stylet morphology indicate that it could belong to Tylencholaimus; however, the pharyngeal bulb occupies about' one-fifth of the total neck length, and the ven- tromedian supplements are numerous beginning within the range of the spicules; these are two features otherwise unknown in the genus.</p>
<p>437 4. Tylencholaimus zimmermanni (Bally & Reydon, 1931) Thorne & Swanger, 1936 (syn. Dorylaimus zimmermanni Bally & Reydon, 1931) Thorne (1939) considered this species close to T. nanus, but some doubt exists about the structure of the stylet since the original illustration shows the odon- tostyle relatively broad and as long as or longer than the lip region width, two atypical features in the genus. The first author thank the projects DGICYT PB92-0121 (Fauna Iberica III) and PB91-0712 for financial support. ZUSAMMENFASSUNG Revision der Gattung Tylencholaimus de Man, 1876. Die restlichen Arten Diese Arbeit beschäftigt sich mit fünf Arten, die zur Gattung Tylencholaimus de Man, 1876 gehören. Zwei von ihnen sind prodelphisch: T. stecki Steiner, 1914 und T. vulvulatus Rahman, Jairajpuri, Ahmad & Ahmad, 1987. Sie sind gekennzeichnet durch eine nichtscheibenförmige periorale Region, einen sich graduell zum Basal-bulbus ausdehnenden vorderen Pharynxabschnitt und durch einen hinteren Genitalast, der mehr als zwei Körperbreiten lang ist. Die übrigen drei Arten sind opistodelphisch: T. intermedius n. sp., T. gertii Kruger, 1965 und T. pakistanensis Timm, 1964. Die genannten fünf Arten werden beschrieben und abgebildet. Außerdem wird T. rosii Yeates, 1979 in die Gattung Tantunema Siddiqi, 1979 überführt als Tantunema rosii (Yeates, 1979) n. comb.. Kürzlich beschriebene oder in Tylencholaimus überführte Arten sowie Arten, die als species inquirendae eingestuft werden, werden kurz diskutiert. REFERENCES ALTHERR, E. (1950). Les Nématodes du Parc National Suisse. Ergebnisse der wissenschaftlichen Untersuchung des schweizerischen Nationalparks 22, 1-46. ANDRÁSSY, I. (1958). Erd- und Süsswassernematoden aus Bulgarien. Acta Zoologica Academiae Scientiarum Hungaricae 4, 1-88. ANDRÁSSY, I. (1959). Taxonomische Uebersicht der Dorylaimen (Nematoda). I. Acta Zoologica Academiae Scientiarum Hungaricae 5, 191-240. ANDRÁSSY, I. (1986). The genus Eudorylaimus Andrássy, 1959 and the present status of its species (Nematoda: Qudsianematidae). Opuscula Zoologica Budapest 22, 3-42. ANDRÁSSY, I. (1991). The superfamily Dorylaimoidea (Nematoda) - a review. Family Qudsiane- matidae, II. Opuscula Zoologica Budapest 24, 3-55. COOMANS, A. (1962). Some species of Dorylaimoidea found in Belgium. I. Members of the Tylencholaiminae Filipjev, 1934. Nematologica 7, 146-154. DE CONINCK, L. (1935). Contribution à la connaissance des nématodes libres du Congo belge. I. Les nématodes libres des marais de la Nyamuamba (Ruwenzori) et des sources chaudes du Mont Banze (Lac Kivu). Revue de Zoologie et de Botanique Africaines 26, 249-326. DE MAN, J. G. (1921). Nouvelles recherches sur les nématodes libres terricoles de la Hollande. Capita Zoologica 1, 3-62. DHANACHAND, C. (1994). Tylencholaimus loofi nomen novum for T. minutus Dhanachand et al., 1992. Current Nematology 4 (1993), 243.</p>
<p>438 DHANACHAND, C., MOHILAL, N. & JOYMATI, L. (1992). Tylencholaimus minutus n. sp. and Oriverutus prodelphus n. sp. (Nematoda: Dorylaimida) from Manipur. Current Nematology 3, 149-152. FUCHS, G. (1930). Neue an Borken- und Rüsselkäfer gebundene Nematoden, halbparasitische und Wohnungseinmieter. Freilebende Nematoden aus Moos und Walderde in Borken- und Rüsselkäfergängen. Zoologische Jahrbücher Jena 59, 505-646. GADEA, E. (1953). Nematodos libres terrestres de los Montes Cántabros. Publicaciones del Instituto de Biología Aplicada 14, 71-108. HERNANDEZ, M. A. & JORDANA, R. (1990). Estudio cualitativo de la fauna de nematodos de los suelos de tres bosques de Quercus en Navarra, España. In: Sesión Homenaje al Profesor García de Viedma. pp. 63-85. Ed. Fundación Conde del Valle de Salazar. JAIN, S. K., SAXENA, R. & SHARMA, R. K. (1993). New species of the genus Tylencholaimus de Man, 1876 (Nematoda: Dorylaimida). Current Nematology 4, 89-92. JAIRAJPURI, M.S. & AHMAD, W. (1992). Dorylaimida - free-living, predaceous and plant parasitic nematodes. Leiden: E. J. Brill. 458 pp. KRUGER, S. P. (1965). New species of the genera Tylencholaimus and Dorylaimellus from South Africa. Proceedings of the Helminthological Society of Washington 32, 1-7. LOOF, P. A. A. & JAIRAJPURI, M. S. (1968). Taxonomic studies on the genus Tylencholaimus de Man, 1876 (Dorylaimoidea) with a key to the species. Nematologica 14, 317-350. MEYL, A. H. (1954a). Die bisher in Italien gefundenen freilebenden Erd und Süsswasser- Nematoden. Archivio Zoologico Italiano 39, 161-264. MEYL, A. H. (1954b). Die Nematodenfauna höherer Pilze in Laub und Nadelwäldern zwischen Braunschweig und dem Harz. Mycopathologica et Mycologia Applicata 7, 1-180. MICOLETZKY, H. (1922). Die freilebenden Erd-Nematoden mit besonderer Berücksichtigung der Steiermark und der Bukowina, zugleich mit einer Revision samtlicher nicht mariner, freileben- der Nematoden in Form von Genus - Beschreibungen und Bestimmungsschlüsseln. Archiv für Naturgeschichte, Berlin 87 (1921), 1-320. MICOLETZKY, H. (1925). Die freilebenden Süsswasser- und Moornematoden Dänemarks. Nebst Anhang: Ueber Amöbosporidien und andere Parasiten bei freilebenden Nematoden. Mémoires de l'Academie Royale des Sciences et des Letres de Danemark, Copenhague 10, 57-310. PEÑA SANTIAGO, R. & COOMANS, A. (1994a). Revision of the genus Tylencholaimus de Man, 1876. Didelphic species. Nematologica 40, 32-68. PEÑA SANTIAGO, R. & COOMANS, A. (1994b). Revision of the genus Tylencholaimus de Man, 1876. Prodelphic species: Part II. Nematologica 40, 186-213. PEÑA SANTIAGO, R. & COOMANS, A. (1994c). Revision of the genus Tylencholaimus de Man, 1876. Prodelphic species: Part III. Nematologica 40, 348-368. POPOVICI, I. (1968). Nematode din sol noi pentru fauna Romaniei. Studii si cercetari de Biologie, Seria Zoologie 20, 255-259. RAHMAN, M. F., JAIRAJPURI, M. S., AHMAD, W. & AHMAD, I. (1987). Three new species of dorylaim nematodes from the Northeastern Region of India. Indian Journal of Nematology 16 (1986), 197-204. SAUER, M. R. (1969). Species of Leptonchus, Tylencholaimus and Chitwoodius from River Murray areas, Australia. Nematologica 15, 361-369. SIDDIQI, M. R. (1982). Seven new genera of dorylaimid nematodes form Colombia rain forest. Systematic Parasitology 6, 69-87. STEINER, G. (1914). Freilebende Nematoden aus der Schweiz. 2. Teil einer vorläufigen Mitteilung. Archiv Hydrobiologie und Planktonkunde 9, 420-438. THORNE, G. (1939). A monograph of the nematodes of the superfamily Dorylaimoidea. Capita Zoologica 8, 1-261.</p>
<p>439 THORNE, G. & SWANGER, H. H. (1936). A monograph of the nematode genera Dorylaimus Du- jardin, Aporcelaimus n. g., Dorylaimoides n. g. and Pungentus n. g. Capita Zoologica 6, 1-223. TIMM, R. V. (1964). Nematodes of the superfamily Dorylaimoidea from East Pakistan. Proceedings of the Helminthological Society of Washington 31, 144-153. VINCIGUERRA, M. T. (1982). Two new species of Dorylaimida (Nematoda) from Italian Alps. Animalia 9, 313-319. VINCIGUERRA, M. T. (1986). New and known species of Tylencholaimus de Man, 1876 (Dorylaim- ida, Nematoda) from Italian beech forest with a key to the species. Nematologia Mediterranea 14, 107-116. WINISZEWSKA, G. (1987). The free-living soil nematodes (Nematoda) of the Swietokrzyskie Moun- tains. Fragmenta Faunistica 31, 11-41.</p>
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<mods version="3.6"><titleInfo><title>Revision of the Genus Tylencholaimus De Man, 1876. Remaining Species</title>
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<titleInfo type="alternative" contentType="CDATA"><title>Revision of the Genus Tylencholaimus De Man, 1876. Remaining Species</title>
</titleInfo>
<name type="personal"><namePart type="given">R.</namePart>
<namePart type="family">Pena Santiago</namePart>
<role><roleTerm type="text">author</roleTerm>
</role>
</name>
<name type="personal"><namePart type="given">A.</namePart>
<namePart type="family">Coomans</namePart>
<role><roleTerm type="text">author</roleTerm>
</role>
</name>
<typeOfResource>text</typeOfResource>
<genre type="research-article" displayLabel="research-article"></genre>
<originInfo><publisher>BRILL</publisher>
<place><placeTerm type="text">The Netherlands</placeTerm>
</place>
<dateIssued encoding="w3cdtf">1996</dateIssued>
<dateCreated encoding="w3cdtf">1996</dateCreated>
<copyrightDate encoding="w3cdtf">1996</copyrightDate>
</originInfo>
<language><languageTerm type="code" authority="iso639-2b">eng</languageTerm>
<languageTerm type="code" authority="rfc3066">en</languageTerm>
</language>
<physicalDescription><internetMediaType>text/html</internetMediaType>
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<subject><genre>keywords</genre>
<topic>prodelphic and opisthodelphic species</topic>
<topic>Tylencholaimus</topic>
<topic>taxonomy</topic>
<topic>revision</topic>
</subject>
<relatedItem type="host"><titleInfo><title>Nematologica</title>
<subTitle>International Journal of Nematological Research</subTitle>
</titleInfo>
<titleInfo type="abbreviated"><title>NEMA</title>
</titleInfo>
<genre type="journal">journal</genre>
<identifier type="ISSN">0028-2596</identifier>
<identifier type="eISSN">1875-2926</identifier>
<part><date>1996</date>
<detail type="volume"><caption>vol.</caption>
<number>42</number>
</detail>
<detail type="issue"><caption>no.</caption>
<number>4</number>
</detail>
<extent unit="pages"><start>417</start>
<end>439</end>
</extent>
</part>
</relatedItem>
<identifier type="istex">E1B42DD75E7414C86FB8B30A4F95E9CD886DE83A</identifier>
<identifier type="DOI">10.1163/004525996X00037</identifier>
<identifier type="href">18752926_042_04_s003_text.pdf</identifier>
<accessCondition type="use and reproduction" contentType="copyright">© 1996 Koninklijke Brill NV, Leiden, The Netherlands</accessCondition>
<recordInfo><recordContentSource>BRILL Journals</recordContentSource>
<recordOrigin>Koninklijke Brill NV, Leiden, The Netherlands</recordOrigin>
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