taxonID	type	description	language	source
4B7387B5FFBBFFF70CDDD43BA88A4A34.taxon	materials_examined	Material examined. Eighty-two females from seven different localities, most of them in good state of preservation.	en	Ahmad, Wasim, Ahad, Sumaya, Islam, Md. Niraul, Sturhan, Dieter (2018): Taxonomy of the genus Doryllium Cobb, 1920 (Nematoda: Dorylaimida) with description of two new and a known species. Zootaxa 4441 (2): 261-278, DOI: 10.11646/zootaxa.4441.2.4
4B7387B5FFBBFFF70CDDD43BA88A4A34.taxon	description	Description. For measurements, see Table 1. Female: Slender nematodes of small size; body cylindrical, slightly curved ventrad upon fixation, tapering slightly towards the anterior extremity, posteriorly ending in a rounded tail. Cuticle with two distinct layers, 2.0 µm thick at midbody and 3 – 4 µm on tail. Outer cuticle with fine transverse striation; inner layer with distinct transverse striations. Lateral chords occupying about one-fourth to one-third of the midbody diameter. Lip region slightly offset by a weak constriction, 2.0 – 2.5 times as wide as high and about one-third of body diameter at neck base; lips amalgamated, with rounded contour, inner portion transformed into a perioral disc. Amphids cup-shaped, aperture occupying about one-half of the lip region diameter. Stoma a truncate cone. Odontostyle 0.8 – 1.1 times the lip region diameter long, aperture about one-fourth of its length. Odontophore with distinct basal knobs, 1.4 – 2.0 times the odontostyle length. Guiding ring simple, refractive, at 0.8 – 1.0 times lip region diameter from anterior end. Pharynx consists of a slender and weakly muscular anterior part, separated from basal bulb by a constriction. Pharyngeal bulb cylindroid, occupying about 22 – 27 % of total neck length. Pharyngeal glands often visible. Cardia short, rounded to conoid, about one-fifth to one-fourth of midbody diameter long. Nerve ring located at 42 – 49 % of neck length from anterior end. Genital system monodelphic-opisthodelphic. Ovary reflexed, measuring 39 – 127 µm long; oocytes arranged in single row except near tip. Oviduct joining the ovary subterminally, separated from the uterus by a narrowing, but with no apparent sphincter seen. Anterior genital branch absent or a rudimentary sac, less than one-third of corresponding body diameter. Vagina cylindrical; pars proximalis vaginae 4 – 8 µm long, wall encircled by muscles; pars distalis vaginae short, 1.5 – 2.0 µm long with slightly curved walls; pars refringens absent. Vulva apparently a transverse slit. Prerectum 3.3 – 4.9 anal body diameters long. Intestine-prerectum junction guarded by three cells. Rectum 0.8 – 1.0 anal body diameter long. Tail cylindroid, hemispheroid to slightly clavate terminus, 1.3 – 2.0 anal body diameters long, cuticle at tail terminus thick, hyaline portion of the tail variable; caudal pores two on each side. Male: Not found Habitats and localities. Soil samples collected from around rhizosphere of forest trees from: 1 Dachigam National Park, Srinagar, Jammu & Kashmir, India; 34 o 8 ’ 36 ” N 75 o 2 ’ 16 ” E 2 Baramullah, Kashmir valley, Jammu & Kashmir, India; 34 o 11 ’ 52.8 ” N 74 o 21 ’ 50.4 ” E 3 Kangna valley, Ganderbal district, Jamuu & Kashmir state, India; 34 o 9 ’ 36 ” N 75 o 33 ’ 0 ” E iv). Kigwema, Kohima district, Nagaland, India; 25 o 60 ’ 62 ” N, 94 o 12 ’ 70 ” E 4 Gudalur, Theni district, Tamil Nadu, India; 11 o 50 ’ 30 ” N, 76 o 49 ’ 17 ” E 5 Ooty, Nilgiris district, Tamil Nadu, India; 11 ° 24 ' 0 " N, 76 ° 42 ' 0 " E 6 vii). Freiburg, Germany; 47 ° 59 ' 56 ” N, 7 ° 50 ' 31 " E TABLE]. Measurements of Doryllium minor Jairajpuri, 1963. (All measurements in µm) …… continued on the next page TABLE]. (Continued)	en	Ahmad, Wasim, Ahad, Sumaya, Islam, Md. Niraul, Sturhan, Dieter (2018): Taxonomy of the genus Doryllium Cobb, 1920 (Nematoda: Dorylaimida) with description of two new and a known species. Zootaxa 4441 (2): 261-278, DOI: 10.11646/zootaxa.4441.2.4
4B7387B5FFBBFFF70CDDD43BA88A4A34.taxon	discussion	Remarks. Jairajpuri (1963) described Doryllium minor from Srinagar, Kashmir, India. Goseco et al. (1975) redescribed this species based on a study of the holotype as well as additional material from Indiana. Goseco et al. (1981) further redescribed this species from Panama while Nasira et al. (2005) redescribed it from Pakistan. The present specimens, collected from three entirely different geographical areas in India, as well as a population from Germany provide a much wider range of morphometric data on this widely distributed species. Interestingly, the morphometric data of all seven of these populations are quite similar without any significant differences; however, some morphological differences were recorded. The three populations from Kashmir valley, the region from where the type population was collected fits well with the original morphometric data provided by Jairajpuri (1963). However, the amphid is rather cup-shaped in the Dachigam National Park and Kangana valley populations, whereas it is stirrup-shaped in the Baramullah population. The outer cuticle is smooth in all three Kashmir populations as well as in the Nagaland population. Jairajpuri (1963) and Goseco et al. (1975) described outer cuticle as smooth. The Nagaland population also has a cup-shaped amphid. The two populations from Tamil Nadu, as well as the German population, have a distinctly striated outer cuticle and cup-shaped amphid. The tail in the Gudalur, Tamil Nadu population is similar to that of the type population, whereas it is rather more clavate in the Ooty population as well as in the German population. None of these morphological differences show a definite combination of characters, hence it is concluded that these differences are simple geographical variations and one has to be very cautious when using these characters individually when differentiating species.	en	Ahmad, Wasim, Ahad, Sumaya, Islam, Md. Niraul, Sturhan, Dieter (2018): Taxonomy of the genus Doryllium Cobb, 1920 (Nematoda: Dorylaimida) with description of two new and a known species. Zootaxa 4441 (2): 261-278, DOI: 10.11646/zootaxa.4441.2.4
4B7387B5FFBCFFFB0CDDD204A9034EF4.taxon	materials_examined	Material examined. Fourteen females in good state of preservation.	en	Ahmad, Wasim, Ahad, Sumaya, Islam, Md. Niraul, Sturhan, Dieter (2018): Taxonomy of the genus Doryllium Cobb, 1920 (Nematoda: Dorylaimida) with description of two new and a known species. Zootaxa 4441 (2): 261-278, DOI: 10.11646/zootaxa.4441.2.4
4B7387B5FFBCFFFB0CDDD204A9034EF4.taxon	description	Description. Measurements, see Table 2 Female: Small sized nematodes, slightly curved upon fixation; body cylindrical tapering gradually towards both the extremities. Cuticle with two distinct layers, 3 – 4 µm thick at midbody and 4 – 5 µm on tail. Outer cuticle finely striated; inner layer smooth. Lateral chords occupying about one-fourth to one-third of the midbody diameter. Lateral, dorsal and ventral body pores indistinct. Lip region continuous, 1.6 – 2.0 times as wide as high or about one-third of the body diameter at neck base; lips rounded, amalgamated; papillae barely raised. Amphids cup-shaped, their aperture occupying about one-half of lip region diameter. Stoma a truncate cone. Odontostyle asymmetrical, 0.8 – 1.0 times the lip region diameter in length, with distinct lumen and aperture. Odontophore flanged, 1.8 – 2.1 times the odontostyle length. Guiding ring simple, refractive, at 0.6 times lip region diameter from anterior end. Pharynx consists of a slender and weakly muscular anterior part, expanding into a slightly constricted pyriform basal bulb, occupying about 16 – 18 % of total neck length. Cardia rounded-conoid, about one-third of the corresponding body diameter long. Nerve ring located at 54 – 57 % of neck length from anterior end. Genital system mono-opisthodelphic. Ovary reflexed, measuring 51 – 78 µm long, reaching the oviduct-uterus junction (surpassing the oviduct-uterus junction in one specimen); oocytes arranged in single row except near tip. Oviduct joining the ovary subterminally, measuring 65 – 76 µm, proximal and distal parts not differentiated. Oviduct-uterus junction marked by weak sphincter. Uterus short and tubular, measuring 15 – 20 µm long. Anterior genital branch small, 0.3 – 0.5 times midbody diameter long. Vagina cylindrical; pars proximalis vaginae 5 – 6 µm long, wall encircled by muscles; pars distalis vaginae short, 3 – 4 µm long with slightly curved walls; pars refringens absent. Vulva a transverse slit. Prerectum 2.3 – 2.5 and rectum 0.9 – 1.2 anal body diameters long. Tail rounded-hemispheroid, 1.2 – 1.5 anal body diameters long. Male: Not found.	en	Ahmad, Wasim, Ahad, Sumaya, Islam, Md. Niraul, Sturhan, Dieter (2018): Taxonomy of the genus Doryllium Cobb, 1920 (Nematoda: Dorylaimida) with description of two new and a known species. Zootaxa 4441 (2): 261-278, DOI: 10.11646/zootaxa.4441.2.4
4B7387B5FFBCFFFB0CDDD204A9034EF4.taxon	materials_examined	Type habitat and locality. Meadow soil, Nienhagen near Kassel, Germany; collected by Dr. Dieter Sturhan on 29.05.1985. GPS Coordinates 51 ° 18 ' 46 '' N and 9 ° 28 ' 53 '' E. Type specimens. Holotype female on slide Doryllium enigmatum sp. n. / 1; paratype females on slides Doryllium enigmatum sp. n. / 2 – 8; deposited with the nematode collection of the Department of Zoology, Aligarh Muslim University, India and on slide Doryllium enigmatum sp. n. / 9; deposited with the German nematode collection.	en	Ahmad, Wasim, Ahad, Sumaya, Islam, Md. Niraul, Sturhan, Dieter (2018): Taxonomy of the genus Doryllium Cobb, 1920 (Nematoda: Dorylaimida) with description of two new and a known species. Zootaxa 4441 (2): 261-278, DOI: 10.11646/zootaxa.4441.2.4
4B7387B5FFBCFFFB0CDDD204A9034EF4.taxon	etymology	Etymology. The new species is named D. enigmatum because of its unusual asymmetrical odontostyle and flanged odontophore, which is unique for Doryllium.	en	Ahmad, Wasim, Ahad, Sumaya, Islam, Md. Niraul, Sturhan, Dieter (2018): Taxonomy of the genus Doryllium Cobb, 1920 (Nematoda: Dorylaimida) with description of two new and a known species. Zootaxa 4441 (2): 261-278, DOI: 10.11646/zootaxa.4441.2.4
4B7387B5FFBCFFFB0CDDD204A9034EF4.taxon	diagnosis	Diagnosis and relationships. Doryllium enigmatum sp. n. is characterized by a body length of 0.40 – 0.52 mm; continuous lip region; rounded lips; asymmetrical odontostyle, 5 – 6 µm long, flanged odontophore 9 – 11 µm long (total spear length: 14 – 17 µm); pharynx with slender anterior part, which expands into a slightly constricted pyriform basal bulb occupying about 16 – 18 % of total neck length; mono-opisthodelphic female genital system, anterior uterine sac 7 – 9 µm; transverse vulva and rounded-hemispheroidtail. The new species is distinctive because of its asymmetrical odontostyle which is unusual for a Doyllium species. In the shape of its lip region and nature of pharyngeal expansion, the new species comes close to D. uniforme Cobb, 1920 but differs in its smaller body size (vs L = 0.78 – 0.85 mm); more posterior vulva position (vs V = 33 – 35); very short anterior uterine sac (vs anterior uterine branch 32 µm); comparatively shorter tail (c = 24 – 32 vs 38 – 41) and in the absence of males (vs present).	en	Ahmad, Wasim, Ahad, Sumaya, Islam, Md. Niraul, Sturhan, Dieter (2018): Taxonomy of the genus Doryllium Cobb, 1920 (Nematoda: Dorylaimida) with description of two new and a known species. Zootaxa 4441 (2): 261-278, DOI: 10.11646/zootaxa.4441.2.4
4B7387B5FFB0FFFC0CDDD7C8A8B24FA3.taxon	materials_examined	Material examined. Eight females and five males in good state of preservation.	en	Ahmad, Wasim, Ahad, Sumaya, Islam, Md. Niraul, Sturhan, Dieter (2018): Taxonomy of the genus Doryllium Cobb, 1920 (Nematoda: Dorylaimida) with description of two new and a known species. Zootaxa 4441 (2): 261-278, DOI: 10.11646/zootaxa.4441.2.4
4B7387B5FFB0FFFC0CDDD7C8A8B24FA3.taxon	description	Description. Measurements, see Table 3 Female: Small sized nematodes, slightly curved ventrad upon fixation; body cylindrical tapering slightly towards the extremities. Cuticle with two distinct layers, 3 – 4 µm thick at midbody and 5 – 6 µm on tail. Outer cuticle finely striated; inner layer thick in tail region. Lateral chords occupying about one-fourth to one-third of the midbody diameter. Ventral and lateral body pores indistinct. A dorsal body pore slightly posterior to nerve ring visible in a few specimens. Lip region slightly offset, 1.7 – 2.0 times as wide as high or about one-third of body diameter at neck base; lips rounded and amalgamated; labial papillae barely raised. Amphids cup-shaped, aperture occupying about one-half of lip region diameter. Stoma a truncate cone. Odontostyle asymmetrical, about as long as lip region diameter, with distinct lumen and aperture. Odontophore slightly flanged, 1.6 – 1.8 times the odontostyle length. Guiding ring simple, refractive, located 0.6 times lip region diameter from anterior end. Pharynx consists of a slender and weakly muscular anterior part, expanding gradually into a pear-shaped basal bulb, occupying about 16 – 18 % of total neck length. Cardia rounded-conoid, about one-third corresponding body diameter long. Nerve ring at 50 – 58 % of neck length from anterior end. Genital system mono-opisthodelphic. Ovary reflexed, measuring 98 – 161 µm long, reaching the oviduct-uterus junction; oocytes arranged in single row except near tip. Oviduct joining the ovary subterminally, measuring 116 – 165 µm, its proximal and distal parts not differentiated. Oviduct-uterus junction marked by weak sphincter. Uterus short and tubular, measuring 35 – 47 µm, filled with sperms; uterine egg, measuring 65 x 19 µm, observed in one specimen. Anterior genital branch small, 0.7 – 1.5 midbody diameters long, filled with sperms. Vagina cylindrical; pars proximalis vaginae 4 – 5 µm long, wall encircled by muscles; pars distalis vaginae short, 3 µm long with slightly curved walls; pars refringens absent. Vulva a transverse slit. Prerectum 2.2 – 2.5 and rectum 0.7 – 1.2 anal body diameters long. Tail short, rounded hemispheroid, 0.8 – 1.1 anal body diameters long. Male: General morphology similar to female except for the posterior region being more ventrally curved. Sperms spindle-shaped. Only the adcloacal pair of supplements present, located at 6 µm anterior to cloacal aperture. Spicules, slender, slightly curved ventrad, 1.5 times cloacal body diameter long. Lateral guiding pieces, simple, rod-like, one-fourth of the spicules length. Tail short, rounded hemispheroid, 1.1 anal body diameter long.	en	Ahmad, Wasim, Ahad, Sumaya, Islam, Md. Niraul, Sturhan, Dieter (2018): Taxonomy of the genus Doryllium Cobb, 1920 (Nematoda: Dorylaimida) with description of two new and a known species. Zootaxa 4441 (2): 261-278, DOI: 10.11646/zootaxa.4441.2.4
4B7387B5FFB0FFFC0CDDD7C8A8B24FA3.taxon	materials_examined	Type habitat and locality. Oat field, Lintach near Amberg, Germany; collected by Dr. Dieter Sturhan on 0 7.10. 1981. GPS Coordinates 49 ° 26 ′ 34 ″ N and 11 ° 51 ′ 45 ″ E Type specimens. Holotype female on slide Doryllium asymmetricum sp. n. / 1; paratypes females on slides Doryllium asymmetricum n. sp. / 2 – 5; paratype males on slides Doryllium asymmetricum sp. n. / 6 – 8 deposited with the nematode collection of the Department of Zoology, Aligarh Muslim University, India. Two paratype female and males each deposited with German nematode collection.	en	Ahmad, Wasim, Ahad, Sumaya, Islam, Md. Niraul, Sturhan, Dieter (2018): Taxonomy of the genus Doryllium Cobb, 1920 (Nematoda: Dorylaimida) with description of two new and a known species. Zootaxa 4441 (2): 261-278, DOI: 10.11646/zootaxa.4441.2.4
4B7387B5FFB0FFFC0CDDD7C8A8B24FA3.taxon	etymology	Etymology. The new species is named D. asymmetricum sp. n. because of its asymmetrical odontostyle.	en	Ahmad, Wasim, Ahad, Sumaya, Islam, Md. Niraul, Sturhan, Dieter (2018): Taxonomy of the genus Doryllium Cobb, 1920 (Nematoda: Dorylaimida) with description of two new and a known species. Zootaxa 4441 (2): 261-278, DOI: 10.11646/zootaxa.4441.2.4
4B7387B5FFB0FFFC0CDDD7C8A8B24FA3.taxon	diagnosis	Diagnosis and relationships. Doryllium asymmetricum sp. n. is characterized by having a body length of 0.52 – 0.63 mm; cap-like, slightly offset lip region; asymmetrical odontostyle, 6 µm long; weakly flanged odontophore 9 – 11 µm long (total spear length: 16 – 17 µm); pharynx with slender anterior part, which expands gradually into pear-shaped basal bulb occupying about 16.4 – 18.5 % of total neck length; mono-opisthodelphic female genital system; anterior uterine sac 17 – 30 µm, filled with sperms; transverse vulva and roundedhemispheroid tail. With its unconstricted pharyngeal bulb, presence of anterior uterine sac and tail shape, the new species comes close to Doryllium jamesi (Goseco, Ferris & Ferris, 1975) Andrássy, 2009 and Doryllium nudum Thorne, 1964. From D. jamesi, it differs in having a slightly longer body (vs 0.45 – 0.50 mm); asymmetrical odontostyle (vs symmetrical); lower c ratio (vs 56 – 64); more anteriorly located vulva (vs 41 – 47 %); longer anterior uterine sac (vs 6.4 µm) and longer tail (vs 7 – 9 µm). From Doryllium nudum, the new species differs in having a slightly longer body (vs 0.50 mm); higher a, b and c ratios (vs 21, 3.5 and 34, respectively); vulva more anteriorly located (vs 38 %); shorter odontostyle (vs 7 µm); smaller combined odontostyle and odontophore length (vs 19 µm); asymmetrical odontostyle and odontophore with weak flanges (vs odontostyle symmetrical and odontophore without flanges) and males present (vs absent).	en	Ahmad, Wasim, Ahad, Sumaya, Islam, Md. Niraul, Sturhan, Dieter (2018): Taxonomy of the genus Doryllium Cobb, 1920 (Nematoda: Dorylaimida) with description of two new and a known species. Zootaxa 4441 (2): 261-278, DOI: 10.11646/zootaxa.4441.2.4
4B7387B5FFB0FFFC0CDDD7C8A8B24FA3.taxon	discussion	Discussion. Both new species distinctly differ from all the known species of the genus Doryllium in their characteristic odontostyle asymmetry (vs short, tubular) and in the odontophore bearing flanges instead of round basal knobs. However, there is a very fine line of differentiation between distinct knobs and flanges and sometimes these two terms are used synonymous and this character should not be used for generic differentiation. Even though the odontostyle nature appears different, proposing a new genus on the basis of such a slight difference does not seem reasonable. Hence, for the present, both new species are placed in the genus Doryllium and the diagnosis of the genus is slightly emended to accommodate the additional variation. A diagnostic compendium (table 4) of the species of the genus Doryllium is also provided. …… continued on the next page Lrd = lip region diameter Ods = Odontostyle Ph. bulb = Pharyngeal bulb Vms = # of ventromedian supplements * = Calculated value from illustrations Geo. dist. = Geographical distribution	en	Ahmad, Wasim, Ahad, Sumaya, Islam, Md. Niraul, Sturhan, Dieter (2018): Taxonomy of the genus Doryllium Cobb, 1920 (Nematoda: Dorylaimida) with description of two new and a known species. Zootaxa 4441 (2): 261-278, DOI: 10.11646/zootaxa.4441.2.4
4B7387B5FFB5FFE20CDDD28DAE864BFB.taxon	diagnosis	Diagnosis (emended). Small nematodes, between 0.5 and 1.1 mm in length. Lip region continuous, or offset by constriction or depression. Lips amalgamated, with the inner part sometimes forming a perioral disc. Cheilostom a truncate cone, without any sclerotization. Odontostyle short, tubular or asymmetrical with distinct lumen and aperture. Odontophore with distinct basal knobs or flanges. Anterior portion of pharynx slender and weakly muscular, basal expansion more or less pyriform, with or without a constriction separating it from the anterior portion. Female genital system mono-opisthodelphic, with or without pre-vulval uterine sac. Male with dorylaimoid spicules. Ventromedian supplements only one, adcloacal. Tail similar in both sexes, short and rounded.	en	Ahmad, Wasim, Ahad, Sumaya, Islam, Md. Niraul, Sturhan, Dieter (2018): Taxonomy of the genus Doryllium Cobb, 1920 (Nematoda: Dorylaimida) with description of two new and a known species. Zootaxa 4441 (2): 261-278, DOI: 10.11646/zootaxa.4441.2.4
4B7387B5FFB5FFE20CDDD28DAE864BFB.taxon	materials_examined	Type species: Doryllium uniforme Cobb, 1920	en	Ahmad, Wasim, Ahad, Sumaya, Islam, Md. Niraul, Sturhan, Dieter (2018): Taxonomy of the genus Doryllium Cobb, 1920 (Nematoda: Dorylaimida) with description of two new and a known species. Zootaxa 4441 (2): 261-278, DOI: 10.11646/zootaxa.4441.2.4
